bluuebunny/crossref_metadata_2025 huggingface dataset:
Morphological complexity increase in metazoans
Authors: Valentine James W. , Collins Allen G. , Meyer C. Porter
Published: July 2016

The number of cell types required for the construction of a metazoan body plan can serve as an index of morphological (or anatomical) complexity; living metazoans range from four (placozoans) to over 200 (hominids) somatic cell types. A plot of the times of origin of body plans against their cell type numbers suggests that the upper bound of complexity has increased more or less steadily from the earliest metazoans until today, at an average rate of about one cell type per 3 m.y. (when nerve cell types are lumped). Computer models in which increase or decrease in cell type number was random were used to investigate the behavior of the upper bound of cell type number in evolving clades. The models are Markovian; variance in cell type number increases linearly through time. Scaled to the fossil record of the upper bound of cell type numbers, the models suggest that early rates of increase in maximum complexity were relatively high. The models and the data are mutually consistent and suggest that the Metazoa originated near 600 Ma, that the metazoan “explosion” near the Precambrian/Cambrian transition was not associated with any important increase in complexity of body plans, and that important decreases in the upper bound of complexity are unlikely to have occurred.
Comparative biogeographic analysis of planktic foraminiferal survivorship across the Cretaceous/Tertiary (K/T) boundary
Authors: MacLeod Norman , Keller Gerta
Published: July 2016

It is now widely recognized that a large number of Cretaceous planktic foraminiferal species are commonly found associated with fully Danian faunas in many K/T boundary sections and deep-sea cores. This “Cretaceous” fauna has traditionally been regarded as representing the reworking of older Cretaceous sediments into younger strata, though recent isotopic data from some species indicates that, at least in these instances, the reworking hypothesis is false. To further test this reworking hypothesis the biogeography of this “Cretaceous” fauna is compared to the underlying uppermost Maastrichtian biogeography and to the biogeography of lowermost Danian planktic foraminiferal faunas. Results show that there is no regular decline in species richness, extinction, or faunal co-occurrence values for this “Cretaceous” fauna at progressively higher (=younger) Danian stratigraphic horizons. Moreover, there is no compelling association between the stratigraphic persistence of this “Cretaceous” fauna and shallow depositional settings. Instead, this fauna is characterized by: (1) a close (and predictive) association between “Cretaceous” and indigenous Danian species richness values throughout the lower Danian, (2) a close numerical and geographic correspondence between Danian speciation and the disappearance of “Cretaceous” species from the Danian fossil record, and (3) a pronounced similarity between changes in the general biogeographic structures of the “Cretaceous” and associated Danian faunas throughout the study interval. These data suggest that the K/T planktic foraminiferal extinction event exhibited a marked geographic structure with low and middle latitude faunas experiencing differentially high extinction rates in the lowermost Danian zones P0 and P1a and high latitude survivor faunas persisting more or less unchanged into the overlying zone, P1b and P1c. Taken together, these results challenge the traditional concept of an instantaneous uppermost Cretaceous planktic foraminiferal mass extinction and its proposed causal connection to bolide impact.
Appearance event ordination: a new biochronologic method
Authors: Alroy John
Published: July 2016

The fundamental goal of biochronology is ordering taxonomic first and last appearance events. The most useful biochronologic data are of the form “the first appearance event of one taxon predates the last appearance event of a second taxon” (FAE < LAE). FAE < LAE data sets are unusually reliable because they converge on a unique solution with greater sampling. The fact that the FAE of one taxoni< the LAE of another taxonjalways can be inferred either ifiis found lower thanjin a stratigraphic section, or ifiandjco-occur in at least one taxonomic list. Thus, FAE < LAE data accurately synthesize two disparate sources of information: routine biostratigraphic observations and taxonomic lists that may have no stratigraphic context. Appearance event ordination, the new method introduced here, is intended to summarize FAE < LAE data. The algorithm is founded on the following parsimony criterion: arrangements of FAEs and LAEs should always imply FAEi< LAEjwhen this is known, and otherwise imply LAEj< FAEiwhenever possible. The technique differs from others related to correspondence analysis in its use of FAE < LAE data and explicit definition as a parsimony method. The algorithm is even more unique in that it uses different subsets of FAEi< LAEjstatements at each iterative step, converging on separate sets of scores for the FAEs and LAEs. After arranging either the FAEs or the LAEs on the basis of their scores, the other set of scores can be discarded and the best arrangement of the remaining events can be inferred directly. An analysis of the Plio-Pleistocene mammalian record in the Lake Turkana region is used to illustrate the method. Biochronologic resolution on the order of 0.2-1.5 m.y. is achieved. The Turkana species lists by themselves demonstrate enough FAEi< LAEjrelationships to resolve the basic biochronologic pattern, but stratigraphic information is still of great use.
A biostratigraphic method based on a quantification of the characters of Devonian tracheophytes
Authors: Gerrienne Philippe , Streel Maurice
Published: July 2016

Biostratigraphic applications of Devonian plant macrofossils are unusual because of the rarity of these fossil plants. Moreover, the taxonomic determination of these plants is too often either arguable or impossible. Here is proposed a biostratigraphic method based on a quantification of individual plant characters and not on determination of whole fossils. The method is the following: (1) each biocharacter found at a given locality receives a score according to whether it is primitive or derived, so that the biocharacter score increases with increasing derivation; (2) the biostratigraphic coefficients of well-dated localities are calculated, the biostratigraphic coefficient of a given plant locality being the mean of all the biocharacter scores of this locality; (3) the scores of these well-dated localities are used to build a reference scale to which localities under investigation will be compared. A detailed example is developed for Early Devonian times. Various plant biocharacters of seven fossiliferous outcrops have been quantified and the subsequent scores of these localities are presented. This example shows that biostratigraphic results obtained by using the quantification method are comparable in precision with dating based on palynological assemblage zones. Evolutionary aspects of the method are also discussed.
A catastrophic kill of ice-trapped coots: time-averaged versus scavenger-specific disarticulation patterns
Authors: Oliver James S. , Graham Russell W.
Published: July 2016

More than 300 coots (Fulica americana) became frozen in Spring Lake, Tazewell County, Illinois, on December 1, 1985. This catastrophic event permitted 8 weeks of taphonomic observations, which showed that ice forms a stable substrate which permits terrestrial taphonomic processes to be imprinted on lacustrine deposits. Bird and mammal scavengers attacked coot carcasses in different manners, resulting in distinct disarticulation sequences. Bird scavengers preferentially fed on the head, neck, and breast-wing complex, causing early disarticulation of bones in these areas, late loss of hindlimb joints, and minimal bone damage. In contrast, mammal scavengers concentrated their attention on the hindlimb and tail region, resulting in bone breakage and early disarticulation of these body parts, but late disarticulation of the breast-wing complex. These data demonstrate that scavenger-specific feeding behaviors significantly influence disarticulation patterns early in assemblage formation, while anatomy may exert increasingly greater influence on disarticulation patterns as carasses become less attractive to scavengers. Finally, because taphonomic processes change in intensity and type through time, bone frequency and modification patterns will vary according to the time at which the patterns are arrested by burial. Thus, bone frequency and modification patterns should provide an index to the relative importance of specific biotic agents and of anatomy in fossil disarticulation patterns as well as an estimate of the time between death and burial.
Estimating numbers of whole individuals from collections of body parts: a taphonomic limitation of the paleontological record
Authors: Gilinsky Norman L. , Bennington J Bret
Published: July 2016

Paleoecologists have long sought to obtain estimates of the sizes of extinct populations. However, even in ideal cases, accurate counts of individuals have been hampered by the fact that many organisms disarticulate after death and leave their remains in the form of multiple, separated parts. We here analyze the problem of estimating numbers of individuals from collections of parts by developing a general counting theory that elucidates the major contributing variables. We discover that the number of unique individuals of a particular species that are represented in a fossil collection can be described by an intricate set of relationships among (1) the number of body parts that were recovered, (2) the number of body parts that were possessed by organisms belonging to that species, and (3) the number of individuals of that species that served as the source of the parts from which the paleontological sample was obtained (the size of the “sampling domain”). The “minimum number of individuals” and “maximum number of individuals” methods currently used by paleontologists to count individuals emerge as end members in our more general counting theory. The theory shows that the numbers of individuals of a species that are represented in a sample of body parts is fully tractable, at least in a theoretical sense, in terms of the variables just mentioned. The bad news is that the size of the “sampling domain” for a species can never be known exactly, thus placing a very real limit on our ability to count individuals rigorously. The good news is that one can often make a reasonable guess regarding the size of the sampling domain, and can therefore make a more thoroughly informed choice regarding how to estimate numbers of individuals. By isolating the variables involved in determining the numbers of individuals in paleontological samples, we are led to a better appreciation of the limits, and the possibilities, that are inherent in the fossil record.
Recognizing morphospecies in colonial reef corals: I. Landmark-based methods
Authors: Budd Ann F. , Johnson Kenneth G. , Potts Donald C.
Published: July 2016

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.
A study of stasis and change in two species lineages from the Middle Devonian of New York state
Authors: Lieberman Bruce S. , Brett Carlton E. , Eldredge Niles
Published: July 2016

More than 5000 measurements were taken on over 1000 specimens of two species of brachiopods,Mediospirifer audaculusandAthyris spiriferoides,from the Middle Devonian Hamilton Group of New York state. Statistical analyses were performed on these data, with specimens partitioned by their occurrence in one of many paleoenvironments and stratigraphic horizons. Neither species showed substantial morphological departures between first appearance and extinction (the range of the Hamilton Group, roughly 5 m.y.). However, oscillations in morphology were discovered in both taxa.

For the two species we studied, groups of organisms occurring in a single paleoenvironment undergo moderate morphological change through time; however, the net sum of changes through time in all paleoenvironments in which these species occur is essentially zero. Therefore, stasis may be partly a property of the organization of species into different environmental populations. Different “environmental populations” may evolve, but they will typically do so in several different “directions,” generally producing no net change. The difference between the morphology of species in different environments over the whole interval of the Hamilton Group is also nil, thereby ruling out any major role that ecophenotypic effects could play in the patterns recognized herein.
Genetic divergence and geographic diversification in Nautilus
Authors: Wray Charles G. , Landman Neil H. , Saunders W. Bruce , Bonacum James
Published: July 2016

Despite exhaustive investigation of present-day Nautilus, the phylogenetic relationships of the five or six recognized species within this genus remain unclear. Mitochondrial and nuclear DNA sequence data plus a suite of morphological characters are used to investigate phylogenetic relationships. Systematic analysis of the morphological variation fails to characterize described species as independent lineages. However, DNA sequence analysis indicates that there are three geographically distinct clades consisting of western Pacific, eastern Australian/Papua-New Guinean, and western Australian/Indonesian forms. The morphologically and genetically distinct species Nautilus scrobiculatus falls outside the three geographically recognized assemblages. Members of the genus Nautilus also exhibit low levels of sequence divergence. All these data suggest that Nautilus is currently undergoing diversification, which may have begun only several million years ago. These data also suggest that some of the morphological features used to define Nautilus species may simply represent fixed variations in isolated populations within the same species.
Extinction selectivity among lower taxa: gradational patterns and rarefaction error in extinction estimates
Authors: McKinney Michael L.
Published: July 2016

Documenting past environmental disturbances will provide a very incomplete explanation of extinctions until more data on intrinsic (e.g., phylogenetic) responses to disturbances are collected. Taxonomic selectivity can be used to infer phylogenetic inheritance of extinction-biasing traits. Selectivity patterns among higher taxa, such as between mammals and bivalves, are well documented. Selectivity patterns among lower taxa (genus, species) have great potential for understanding the dynamics underlying higher taxic turnover. Two echinoid data sets, of fossil and living taxa, indicate that species extinctions do not occur randomly within genera. Reverse rarefaction estimates of past species extinction rates assume random species extinction within higher taxa, so these widely cited extinction estimates may be inaccurate. Revised estimates based on a simulated curve imply that past species extinctions rates may be 6%–15% lower than previously cited. Possible causes for the observed selectivity patterns are discussed. These include nonrandom phylogenetic nesting of species with traits often cited as enhancing extinction vulnerability, into certain taxa. Such traits include low abundance, large body size, narrow niche breadth, and many others. Phylogenetic nesting of extinction-biasing traits at many taxonomic levels does not predict that a dichotomy of mass-background selectivity based on a few traits will occur. Instead, it predicts patterns of selectivity at many taxonomic levels, and at many spatio-temporal scales.
Observed phenotypic variation in a Paleozoic bryozoan
Authors: Hageman Steven J.
Published: July 2016

Documentation of morphologic variation within and among fossil species (and larger clades) provides fundamental data needed for studies of evolution, paleoecology, and the systematic foundation required for most fields of paleobiology. In paleontological (and, frequently, biological) studies, morphologic variation is used as a general proxy for genetic variation. Although the occurrence of ecophenotypic variation is well appreciated in these studies, it is only with the use of colonial (clonal) organisms that the scope and significance of phenotypic variation can be evaluated directly. Systematic evaluation of intracolonial morphologic variation (transects through growth series) can yield insights about ecophenotypic variation in bryozoans and suggest the most appropriate methods for data collection in paleobiologic and taxonomic studies.

In this study, morphological conservatism is documented within local segments of bryozoan colonies; each zooid is generally more similar to adjacent zooids than to distant zooids within the same colony. One region of a colony, therefore, can be more similar to a region of a different colony than to a distant region of its own colony. Variation within one colony does not, however, represent the total variation among a group of specimens, indicating a colonial level of morphologic control (genetic or macroenvironmental) over morphogenesis. Directional morphogenetic gradients (associated with successive ontogenetic histories) are not recognized in these specimens, but fluctuating trends within colonies (some cyclic), were observed and are indicative of changing microenvironmental influence during skeletal formation. In order to best document morphologic variation within a population, for any type of paleobiological study, individual measurements should be widely distributed over large colony fragments and (or) a minimal number of measurements collected from each of a large number of smaller fragments.

Direct extrapolation of these results to non-colonial organisms is not appropriate at this time. However, additional, related studies with bryozoans and other colonial organisms (e.g., corals, graptolites), should provide a greater, general appreciation of relationships between morphology and genetics.
Sexual versus asexual dispersal in clonal animals: examples from cheilostome bryozoans
Authors: Thomsen Erik , Håkansson Eckart
Published: July 2016

The relative numbers of sexually and asexually recruited colonies and the proportion of brooding zooids were determined in 26 species of cheilostome bryozoans of Late Cretaceous and Early Tertiary ages. Asexual reproduction seems to be much more widespread than previously realized, although its relative importance is related to growth habit. Arborescent species of these fossil assemblages reproduce mainly asexually via fragmentation; encrusting species reproduce sexually via motile larvae. Free-living species use both methods; some reproduce sexually, whereas other species have enhanced the ability to break and reproduce predominantly asexually. Mode of reproduction was stable over a period of 3 m.y. in all species except the vinelike Columnotheca cribrosa. In this species both the proportion of asexual recruits and brooding zooids varied in accord with environmental parameters. In all cases populations dominated by asexual propagation had a significantly lower proportion of brooding zooids than populations dominated by sexual propagation.
Why do lower plants and animals biomineralize?
Authors: Brasier Martin
Published: July 2016

Widespread concern about environmental pollution is putting a new question to the fossil record: How has the biosphere reacted to chemical changes in the past? Monera and Protoctista might be expected to provide valuable clues in this quest since their biomineral remains are generally formed in conditions closely related to the environment. But why do unicells biomineralize at all? It was with such questions in mind that an international symposium of the Systematics Association on “Biomineralization in Lower Plants and Animals” was held at Birmingham on April 15–19, 1985. Monerans, protoctistans, lichens, calcareous algae, and bryozoans were discussed in 36 papers, of which 23 are to be published in a volume by Oxford University Press. This volume, edited by Leadbeater and Riding (1986), will form a natural sequel to the papers in Miller et al. (1984) on mineral phases in biology and in Westbroek and de Jong (1983) on biomineralization and biological metal accumulation.
Clinal geographic variation in mammals: implications for the study of chronoclines
Authors: Koch Paul L.
Published: July 2016

Mammalian species often exhibit clinal geographic variation in body size: individuals tend to be larger in areas with lower mean annual temperature. Climatic change involving increasing or decreasing mean annual temperature may cause clines to shift geographically, resulting in a phenotypic shift at all affected locales within a species' range. I assess the potential of shifting geographic clines to produce morphological trends in the fossil record. Five extant North American mammalian species (Didelphis virginiana, Mephitis mephitis, Odocoileus virginianus, Scalopus aquaticus, and Sciurus carolinensis) are examined to quantify size change along latitudinal clines and to estimate the geographic range and temperature difference commonly associated with a given difference in body size. Relative to body size, the observed size range of skeletal characters within each of these five species is comparable to that seen in a much larger sample of North American mammals. Thus patterns of variation documented for the five species may be used to assess the likelihood of dine translocation as an explanation of size change in the mammalian fossil record. As a case study, I examine three lineages from the Early Eocene of the Bighorn Basin, Wyoming. I determine that size change in these chronoclines represents evolutionary change and is not merely the result of shifting geographic clines.
Stasis in Homo erectus defended
Authors: Rightmire G. Philip
Published: July 2016

Wolpoff's (1984) recent discussion of evolutionary rates in Homo erectus deserves careful study. Whether Homo erectus or other hominid species exhibit gradual, continuous change in key characters or whether instead there is evidence for morphological stasis in the fossil record is an important question. By allocating all Homo erectus specimens to three groups of early, intermediate, and later geological age and by comparing group means for 13 measurements, Wolpoff attempts to show that gradualism is the rule for this mid-Pleistocene taxon. This method is straightforward, but it is crucial that the samples be composed in a manner which is biologically reasonable. I argue here that Wolpoff has not done this. While there may be legitimate doubt concerning sorting of the fossils, especially where specimens are incomplete, several of the individuals said to be representative of Homo erectus are simply inappropriate for use in this analysis. Wolpoff insists that he has employed a “conservative” definition of the species, but instead he has measured everything in sight. This approach to the record does influence his results.
Stasis in the interpretation of evolution inHomo erectus: a reply to Rightmire
Authors: Wolpoff Milford H.
Published: July 2016

Focusing on the question of trends inHomo erectuscranial capacity evolution, Rightmire contends to have reanalyzed the data and demonstrated “a pattern of change … quite different from that described by Wolpoff.” Accepting the procedures used in this earlier analysis (Wolpoff 1984), he does this by changing the composition of theHomo erectussample.
The Feeding Habits of the Shovel-Tusked Gomphotheres: Evidence From Tusk Wear Patterns
Authors: Lambert W. David
Published: July 2016

The shovel-tusked gomphotheres are normally portrayed scooping up water plants with their shovellike mandibular tusks. This portrayal is based on speculation about the possible functions of the lower tusks and misinterpretation of mandibular-tusk wear patterns that goes back to the 1920s and 1930s. In addition, most literature concerning shovel-tusker feeding behavior ignores the possibility that the upper tusks may have had a role in feeding, as they do in modern African elephants. Because wear patterns on tusks provide direct evidence as to how tusks were used during feeding, I examined and interpreted patterns on the lower and upper tusks of shovel-tusked gomphotheres on the basis of theoretical models of tusk wear to determine how the tusks were used in feeding.

This examination led to the following conclusions. (1) The genus Amebelodon was characterized by feeding opportunism, gathering food with both its lower and upper tusks in a variety of ways. (2) There is evidence for similar flexibility within the genus Serbelodon. (3) Wear patterns on mandibular tusks of Platybelodon grangeri and Torynobelodon barnumbrowni indicate that these taxa did not practice mandibular shoveling but used these tusks to cut tough vegetation in a specialized fashion. (4) The flaplike trunks (probosces) generally attributed to shovel-tusked gomphotheres in restorations are shown almost certainly to be erroneous; morphological and functional evidence suggests that these animals had trunks similar to those of modern elephants. (5) Strong tusk sexual dimorphism is unknown in the shovel-tusked gomphotheres except for Platybelodon and should not be found in any taxa that relied on their tusks for procuring food. New restorations of both Amebelodon and Platybelodon are provided that are more consistent with the actual evidence bearing on their feeding behavior.
Contributions of individual taxa to overall morphological disparity
Authors: Foote Mike
Published: July 2016

Two methods are discussed for assessing the contributions of subgroups to the morphological disparity of the larger group containing them. (1) Given an ordination of points representing specimens or species in morphological space, morphological disparity of the entire group is measured as the average squared distance of points from the centroid. The contribution that a subgroup makes to morphological disparity is measured as the average squared distance of its points from the overall centroid (not the subgroup centroid), weighted by the subgroup sample size relative to the total group sample size. Thus, morphological disparity of a group can be additively partitioned into the disparity components of its subgroups, and the relative contributions of these subgroups can be assessed quantitatively. (2) An alternative approach is to compare morphological disparity of a group to the disparity it would have if a certain subgroup were omitted. If the resulting disparity differs substantially from the original disparity, then the subgroup in question is considered to have a significant effect on morphological disparity. Because some subgroups are very centralized in morphological space, omitting them can cause an increase in morphological disparity when disparity is measured as the average dissimilarity among species. In general, relatively large subgroups that are located peripherally in morphospace make the greatest contributions to morphological disparity, and failure to sample smaller groups often has little effect on disparity estimates. The two methods are applied to morphological disparity in trilobites, partitioned at different levels in the taxonomic hierarchy. Results of the two methods are intuitively reasonable and largely in agreement, and point to the predominance of Early Cambrian olenelloids, Cambro-Ordovician Libristoma, Ordovician Asaphina and Cheirurina, Siluro-Devonian Phacopida and Phacopina, and Devonian Proetida.
Patterns of drilling predation on gastropods of the family Turritellidae in the Gulf of California
Authors: Tull Debra S. , Böhning-Gaese Katrin
Published: July 2016

Predatory marine snails and their prey provide a unique look at many aspects of predation events, allowing behavioral inference and studies of coevolution. This study examines differential predation patterns, rates, and success of two co-occurring gastropod predator families which drill two co-occurring species ofTurritella(Turritellidae: Gastropoda) in the Gulf of California. Both naticid and muricid predators, identified by the shapes of their respective boreholes, attacked the thinner-shelledTurritella leucostomamore frequently than the thicker-shelledTurritella gonostoma.Both species were drilled more frequently and more successfully by naticid, as compared to muricid, predators. Naticids drilled prey in the 40- to 70-mm size class most frequently. Prey over 100 mm in length were relatively safe from all drilling predators. Predator size (estimated by borehole diameter) in naticids was correlated with prey size in both species, but for a given-sized prey, predators onT. gonostomawere proportionally larger. There was no size correlation for muricid predators. Unsuccessful attempts (incomplete drilling) were started on the suture more often than were completed holes, for both predator families on both prey species. Naticids began drillingT. leucostomaon the suture significantly less than expected by chance. We looked for possible changes over evolutionary time by analyzing prey shells from Pleistocene and Recent storm deposits. We found no evidence of change in any aspect of implied predatory behavior over the past 100 k.y.
Taxonomic evolution in North American Neogene horses (subfamily Equinae): the rise and fall of an adaptive radiation
Authors: Hulbert Richard C.
Published: July 2016

The 18 m.y. history of the subfamily Equinae (exclusive of Archaeohippus and “Parahippus”) in North America consisted of a 3-m.y. radiation phase, a 9-m.y. steady-state diversity phase, and a 6-m.y. reduction phase. During the steady-state phase, species richness varied between 14 and 20, with two maxima at about 13.5 and 6.5 Ma. Species richness of the tribes Hipparionini and Equini was about equal through the middle Miocene, but hipparionines consistently had more species in the late Miocene and early Pliocene. Overall mean species duration was 3.2 m.y. (n = 50), or an average extinction rate of 0.31 m.y.-1 During the radiation phase, speciation rates were very high (0.5 to 1.4 m.y.-1), while extinction rates were low (<0.10 m.y.-1). Speciation and extinction rates both averaged about 0.15 m.y.-1 during the steady-state phase, with extinction rates having more variation. Extinction rates increased fourfold during the reduction phase, while speciation rates declined slightly. Late Hemphillian extinctions affected both tribes severely, not just the three-toed hipparionines, and were correlated with global climatic change.
Biogeographic and stratigraphic evidence for rapid speciation in semionotid fishes
Authors: McCune Amy R.
Published: July 2016

In this study I take advantage of an unusual system of fossil lakes in eastern North America to estimate the time for speciation of endemic semionotid fishes. Twenty-one species are all found in sedimentary cycle P4, the deposits of a single Early Jurassic lake, in the Towaco Formation of the Newark Basin in New Jersey. To determine the degree of endemism in the fauna from this fossil lake and estimate time for speciation, I surveyed more than 2000 museum specimens from 45 named localities in the Newark Basin and related basins of the Late Triassic to Early Jurassic Newark Supergroup. Six species not found in deposits equal in age to P4 or older are considered to be endemics, eight species occurring in older deposits presumably colonized Lake P4, and evidence for whether the remaining seven species were endemics or colonists is equivocal. The time for the formation, decline, and evaporation of Lake P4, in which P4 sediments were deposited, has been estimated at 21,000-24,000 years. Because all endemic Semionotus first occur in the first third of lake history, the estimated time for speciation of endemics is six species in 5000-8000 years. This rate is remarkably similar to that estimated for the five cichlids in Lake Nabugabo that diverged from Lake Victoria cichlids in about 4000 years.
Trilobite biogeography in the Middle Devonian: geological processes and analytical methods
Authors: Lieberman Bruce S. , Eldredge Niles
Published: July 2016

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.
Comparative taphonomy of bivalves and foraminifera from Holocene tidal flat sediments, Bahia la Choya, Sonora, Mexico (Northern Gulf of California): taphonomic grades and temporal resolution
Authors: Martin Ronald E. , Wehmiller John F. , Harris M. Scott , Liddell W. David
Published: July 2016

We compare the preservation (taphonomic grade) and age ofChione(bivalve) and foraminifera from modern siliciclastic tidal flat sediments of Bahia la Choya, Sonora, Mexico (northern Gulf of California). Disarticulated shells ofChionecollected from the sediment-water interface of Choya Bay exhibit a substantial range in taphonomic grade and age, several hundred years to ∼80–125 ka based on Accelerator Mass Spectrometer14C dates and D-Alloisoleucine/L-Isoleucine values. There is not, however, a one-to-one correspondence between age and taphonomic alteration ofChione:old (or young) valves may be highly altered or they may be relatively pristine. In contrast toChione, most foraminiferal tests at Choya Bay are quite pristine, which suggests a quite young age, but tests are surprisingly old (up to ∼2,000 calendar years based on Accelerator Mass Spectrometer14C dates).

We suggest that following seasonal pulses in reproduction, some foraminiferal tests are rapidly incorporated into a subsurface shell layer by “Conveyor Belt” deposit feeders and preserved there, while the rest of the reproductive pulse rapidly dissolves. Ultimately, some of these buried tests, along withChione, are transported back to the surface by biological activity and storms. The much greater range of taphonomic grades and ages amongChioneshells suggests that they, unlike foraminifera, are sufficiently large and preservable (low surface/volume ratio and chemical reactivity) to undergo many cycles of degradation, burial, and exhumation before complete destruction. The age of foraminiferal tests indicates that time-averaging of microfossil assemblages at Choya Bay is much more insidious than would be expected considering the relatively pristine state of the tests alone.

Based on our studies, the lower limit of temporal resolution of shallow shelf microfossil assemblages appears to be ∼1000 years. We caution, however, that each depositional setting (taphofacies) should be evaluated on a case-by-case basis before gross generalizations are made. Indeed, the discrepancy between age and taphonomic grade of fossil assemblages at Choya Bay suggests that neither hardpart size or taphonomic grade are infallible indicators of test preservability or likely temporal resolution of the host assemblage, and that the dynamics of hardpart input and loss must also be evaluated.
Trophic group and the end-Cretaceous extinction: did deposit feeders have it made in the shade?
Authors: Levinton Jeffrey S.
Published: July 2016

It has been suggested that a sudden event at the end of the Cretaceous period caused a major extinction that was felt disproportionately by creatures in the water column. It also has been argued that benthic deposit feeders, being relatively independent of abundance of organic particles in the water column, should have survived the crisis more readily than suspension feeders, which depended more upon feeding upon phytoplankton. I argue that the hypothesis of relative immunity of deposit feeders is insufficient, because deposit feeders by and large depend upon a supply of organic matter from the water column and would have succumbed to food shortage nearly as rapidly as suspension feeders, possibly within a maximum of three to six months. This near simultaneity of extinction would have been especially true of continental shelf environments. Even in some parts of the deep sea, it is likely that a dependence upon the water column above might have caused deep-sea deposit feeders to succumb rapidly. Therefore, deposit feeders would not necessarily have outsurvived suspension feeders during a crisis of depleted water-column phytoplankton, increased shading by inert particles, or poisoning of the water column and killing of phytoplankton. The relatively lower rate of extinction of nuculoid bivalves may relate instead to their presence in deeper-water refuge habitats, their apparent relative ability to diversify in higher latitudes, or their resistance to factors other than food shortage.
Fossil preservation and the stratigraphic ranges of taxa
Authors: Foote Mike , Raup David M.
Published: July 2016

The incompleteness of the fossil record hinders the inference of evolutionary rates and patterns. Here, we derive relationships among true taxonomic durations, preservation probability, and observed taxonomic ranges. We use these relationships to estimate original distributions of taxonomic durations, preservation probability, and completeness (proportion of taxa preserved), given only the observed ranges. No data on occurrences within the ranges of taxa are required. When preservation is random and the original distribution of durations is exponential, the inference of durations, preservability, and completeness is exact. However, reasonable approximations are possible given non-exponential duration distributions and temporal and taxonomic variation in preservability. Thus, the approaches we describe have great potential in studies of taphonomy, evolutionary rates and patterns, and genealogy.

Analyses of Upper Cambrian-Lower Ordovician trilobite species, Paleozoic crinoid genera, Jurassic bivalve species, and Cenozoic mammal species yield the following results: (1) The preservation probability inferred from stratigraphic ranges alone agrees with that inferred from the analysis of stratigraphic gaps when data on the latter are available. (2) Whereas median durations based on simple tabulations of observed ranges are biased by stratigraphic resolution, our estimates of median duration, extinction rate, and completeness are not biased. (3) The shorter geologic ranges of mammalian species relative to those of bivalves cannot be attributed to a difference in preservation potential. However, we cannot rule out the contribution of taxonomic practice to this difference. (4) In the groups studied, completeness (proportion of species [trilobites, bivalves, mammals] or genera [crinoids] preserved) ranges from 60% to 90%. The higher estimates of completeness at smaller geographic scales support previous suggestions that the incompleteness of the fossil record reflects loss of fossiliferous rock more than failure of species to enter the fossil record in the first place.
Fourier analysis applied toStephanomys(Rodentia, Muridae) molars: nonprogressive evolutionary pattern in a gradual lineage
Authors: Renaud Sabrina , Michaux Jacques , Jaeger Jean-Jacques , Auffray Jean-Christophe
Published: July 2016

Size and shape are analyzed for Pliocene lineages of the rodent genusStephanomysSchaub 1938. Previous phylogenetic studies were based mainly on size variation and descriptive comparisons, without any attempt to quantify shape changes. Hence, on the basis of regular size increase,Stephanomyshas been considered a prime example of phyletic gradualism. In order to quantify morphological variation within the lineage, a method for analyzing complex outlines, the elliptic Fourier transform, was applied to tooth contour (upper and lower first molars). It was then possible to compare evolution in size, estimated by tooth area, as well as evolution of shape, represented by Fourier coefficients.

While size seems to change gradually through time, morphology gives a rather discontinuous evolutionary pattern for both the upper and lower molar. Such a discrepancy between the evolution of size and shape of a single structure suggests that different genetic determinisms and mechanical constraints may act on size and shape. Hence it may be misleading to infer generalized evolutionary processes from either size or shape alone.
Isotopic evidence for interspecies habitat differences during evolution of the Neogene planktonic foraminiferal clade Globoconella
Authors: Schneider Cynthia E. , Kennett James P.
Published: July 2016

Oxygen and carbon isotopic analyses have been conducted on Miocene to Pliocene (6.0 to 2.9 Ma) members of the gradually evolving, deep-dwelling planktonic foraminiferal clade, Globoconella, in temperate waters of the southwest Pacific, Deep Sea Drilling Project (DSDP) Site 593. In the late Miocene the clade began with Globoconella conoidea, and continued through G. conomiozea, G. sphericomiozea, and G. puncticulata to the extant form G. inflata. Isotopic analyses were performed on ancestor-descendant species within the clade to determine if isotopic differences exist between these species which would, in turn, suggest depth and/or seasonal habitat differences and perhaps segregation, as well as ecological changes in the clade. Isotopic analyses were also conducted on the relatively shallow-dwelling planktonic foraminifer Orbulina universa and the benthic form Cibicidoides wuellerstorfi to determine if any relationships exist between the evolution of Globoconella and paleoceanographic/paleoclimatic change.

Small (usually 0.1–0.15 ‰ but up to 0.3 ‰) oxygen isotopic differences exist between ancestor and descendant forms that we believe represent small (∼1°) temperature differences. These temperature differences are inferred to indicate depth and/or seasonal habitat differences and possible segregation between the species during the gradual evolution. The largest differences in oxygen isotopic values occur between ancestor and descendant forms during the most conspicuous morphological transition within the clade near the Miocene/Pliocene boundary. During this interval, the clade underwent a transformation from conical to spherical forms and there was a loss of the keel. No consistent differences were observed between ancestor and descendant carbon isotopic values.

Both morphological and ecological evolution appear to have been associated with paleoceanographic/paleoclimatic changes. Intervals marked by warming of surface-to-upper intermediate waters are associated with evolution of forms with a spherical test and inferred adaptation to cooler waters relative to ancestral forms. We propose two alternative models for the evolution of the Globoconella clade. In the first model, we assume that depth and/or seasonal segregation between ancestor and descendant forms provided a partial barrier to gene flow and that evolution resulted from genetic drift and/or differential selective pressures acting on each morphotype. In the second model, we assume that coeval members of Globoconella formed a vertical and/or seasonal morphocline in the water column and that segregation did not provide an effective barrier to gene flow. Evolution proceeded as directional selection acted on morphotypes of Globoconella inhabiting selectively advantageous positions in the water column.
The early Mesozoic radiation of dinoflagellates
Authors: Fensome R. A. , MacRae R. A. , Moldowan J. M. , Taylor F. J. R. , Williams G. L.
Published: July 2016

Dinoflagellates are a major component of the marine microplankton and, from fossil evidence, appear to have been so for the past 200 million years. In contrast, the pre-Triassic record contains only equivocal occurrences of dinoflagellates, despite the fact that comparative ultrastructural and molecular phylogenetic evidence indicates a Precambrian origin for the lineage. Thus, it has often been assumed that the dearth of Paleozoic fossil dinoflagellates was due to a lack of preservation or recognition and that the relatively sudden appearance of dinoflagellates in the Mesozoic is an artifact of the record. However, new evidence from a detailed analysis of the fossil record and from the biogeochemical record indicates that dinoflagellates did indeed undergo a major evolutionary radiation in the early Mesozoic.