993 resultados para fossil record


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Recovery from the end-Permian mass extinction is frequently described as delayed, with complex ecological communities typically not found in the fossil record until the Middle Triassic epoch. However, the taxonomic diversity of a number of marine groups, ranging from ammonoids to benthic foraminifera, peaked rapidly in the Early Triassic. These variations in biodiversity occur amidst pronounced excursions in the carbon isotope record, which are compatible with episodes of massive CO2 outgassing from the Siberian Large Igneous Province. Here we present a high-resolution Early Triassic temperature record based on the oxygen isotope composition of pristine apatite from fossil conodonts. Our reconstruction shows that the beginning of the Smithian substage of the Early Triassic was marked by a cooler climate, followed by an interval of warmth lasting until the Spathian substage boundary. Cooler conditions resumed in the Spathian. We find the greatest increases in taxonomic diversity during the cooler phases of the early Smithian and early Spathian. In contrast, a period of extreme warmth in the middle and late Smithian was associated with floral ecological change and high faunal taxonomic turnover in the ocean. We suggest that climate upheaval and carbon-cycle perturbations due to volcanic outgassing were important drivers of Early Triassic biotic recovery.

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The Miocene is the last warm episode in Earth history, and this episode was well recorded in Turkey as shown by plant distribution and inferred numerical temperature values. In this study, Ören-Kultak, Hüssamlar and Karacaagac palynofloras from western Turkey, which are characterized by the thermophilous plants (Engelhardia, Sapotaceae, Cyrillaceae, Avicennia, Arecaceae, Palmae), are described. Age determinations of these palynofloras (middle Burdigalian-Langhian) are strengthened by the mammalian fossil record (MN4-5) and strontium isotope results. Palaeoclimate is humid and warm subtropical during the middle Burdigalian-Langhian time interval in Europe and Turkey. However, temperature difference has been observed between Europe and Turkey during this time interval and it could be explained by the palaeogeographic position of countries. Despite some discrepancies in the climatic values and palaeovegetation groups, warm climatic conditions are recorded, based on the palynofloras, in Turkey (Cayyrhan, Havza, Can, Etili, Gönen, Bigadic, Emet, Kirka and Kestelek, Sabuncubeli, Soma, Tire, Kulogullary, Bascayyr, Hüssamlar and Karacaagac), Greece and elsewhere in Europe throughout the middle Burdigalian-Langhian period. This warming is related to the Middle Miocene Climatic Optimum period. Carbon and oxygen isotope values obtained from tooth enamel of Gomphotherium sp. from Kultak and Hüssamlar indicate similar ecological condition during the Burdigalian-Langhian time. This isotopic result and high MAPDRY value from the Kultak locality are in agreement with ecological interpretation of mammalian fossils. Besides, according to the precipitation values, central and northwestern Anatolian sites provide more rainfall during the Burdigalian-Langhian time interval than the western Anatolian sites.

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Downcore cyclic variation in high-resolution nannofossil abundance records from mid-Pliocene equatorial Atlantic ODP Sites 662 and 926 demonstrate the direct response by several Pliocene taxa (notably Discoaster, Sphenolithus and Florisphaera profunda) to orbitally forced climatic variation. In particular, these records display strong obliquity and precessional signals reflecting primarily high latitude, Southern hemisphere changes influencing upwelling intensity and local low-latitude, insolation-driven climatic changes (via the productivity and/or turbidity influence of Amazon-sourced terrigenous material) at Sites 622 and 926 respectively. In seasonal studies of coccolithophorid assemblages, only part of the variation observed can be explained by abiotic processes, so it is perhaps not surprising that in this study few Pliocene nannofossil taxa demonstrate significant correlations with each other or with physical environmental parameters. Only some variance in nannofossil abundances can be explained by the primary controls of temperature and productivity. The rest is attributed to nonlinear responses to climatic changes; biotic processes such as grazing, predation, viral infection and competition, and/or, abiotic factors for which there is no readily available proxy (e.g. salinity). The lack of strong, consistent intra- and inter-relationships of the nannoflora and the environment reflects an ecologically complex, differentiated original community producing a complex integrated signal transmitted into the fossil record.

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40Ar-39Ar step-heating dating was applied to a basalt from Hole 462 and to basalt and dolerite samples from Hole 462A. Only a basalt sample at Hole 462A yielded a reasonable isochron age, 110 ± 3 million years. The radiometric age is consistent with the fossil record (Cenomanian) in the sediments, into which the basalt sill intruded. However, the age is much less than that of the oceanic basement as deduced from the magnetic anomaly (M-26).

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Investigations at a Late Weichselian freshwater basin in northwestern Jutland, Denmark, yielded a fairly rich assemblage of vertebrate remains, mostly bones and teeth of small mammals. The remains are primarily allochthonous and the bones have been subjected to different taphonomic pathways and agents. AMS 14C-dates on terrestrial organic remains provided ages of Middle to Late Allerød time. Identifications revealed the first fossil record in Scandinavia of Rana arvalis, Sorex minutus, Ochotona cf. pusilla, Microtus gregalis, Microtus oeconomus, and Sicista cf. betulinu. Spermophilus cf. major and Desmana moschata, previously found only once and twice respectively, were retrieved, and Sorex araneus and Arvicola terrestris were recovered for the first time beyond the Atlantic chronozone. Ecologically, the Nørre Lyngby small mammal fauna can be characterized by its very high and almost equal proportions of boreal forest and steppe elements followed by a relatively high proportion of tundra elements. The fossil species share a modern area of sympatry north of the Caspian Sea from the river Volga in the west to the southern and western slopes of the Urals. If, however, the large Allerød mammals are added, the fauna is without modern analogues. The Nørre Lyngby fauna can be seen as a last expansion of the North European glacial fauna. Provided that an absolute chronology and a differentiated sea-level curve for the area can be established, the Nørre Lyngby fauna could become important for studies in mammalian dispersal and migration rates.

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NW African climate shows orbital and millenial-scale variations, which are tightly connected to changes in marine productivity. We present an organic-walled dinoflagellate cyst (dinocyst) record from a sediment core off Cape Yubi at about 27°N in the Canary Basin covering the time period from 47 to 3ka before present (BP). The dinocyst record reflects differences in upwelling intensity and seasonality as well as the influence of fluvial input. Sea-level changes play an important role for the upwelling pattern and productivity signals at the core site. Within the studied time interval, four main phases were distinguished. (1) From 45 to 24ka BP, when sea-level was mostly about 75m lower than today, high relative abundances of cysts of heterotrophic taxa point to enhanced upwelling activity, especially during Heinrich Events, while relatively low dinocyst accumulation rates indicate that filament activity at the core location was strongly reduced. (2) At sea-level lowstand during the LGM to H1, dinocyst accumulation rates suggest that local filament formation was even more inhibited. (3) From the early Holocene to about 8ka BP, extraordinary high accumulation rates of most dinocyst species, especially of Lingulodinium machaerophorum, suggest that nutrient supply via fluvial input increased and rising sea-level promoted filament formation. At the same time, the upwelling season prolongated. (4) A relative increase in cysts of photoautotrophic taxa from about 8ka BP on indicates more stratified conditions while fluvial input decreased. Our study shows that productivity records can be very sensitive to regional features. From the dinocyst data we infer that marine surface productivity off Cape Yubi during glacial times was within the scale of modern times but extremely enhanced during deglaciation.

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In the late Paleocene to early Eocene, deep sea benthic foraminifera suffered their only global extinction of the last 75 million years and diversity decreased worldwide by 30-50% in a few thousand years. At Maud Rise (Weddell Sea, Antarctica; Sites 689 and 690, palaeodepths 1100 m and 1900 m) and Walvis Ridge (Southeastern Atlantic, Sites 525 and 527, palaeodepths 1600 m and 3400 m) post-extinction faunas were low-diversity and high-dominance, but the dominant species differed by geographical location. At Maud Rise, post-extinction faunas were dominated by small, biserial and triserial species, while the large, thick-walled, long-lived deep sea species Nuttallides truempyi was absent. At Walvis Ridge, by contrast, they were dominated by long-lived species such as N. truempyi, with common to abundant small abyssaminid species. The faunal dominance patterns at the two locations thus suggest different post-extinction seafloor environments: increased flux of organic matter and possibly decreased oxygen levels at Maud Rise, decreased flux at Walvis Ridge. The species-richness remained very low for about 50 000 years, then gradually increased. The extinction was synchronous with a large, negative, short-term excursion of carbon and oxygen isotopes in planktonic and benthic foraminifera and bulk carbonate. The isotope excursions reached peak negative values in a few thousand years and values returned to pre-excursion levels in about 50 000 years. The carbon isotope excursion was about -2 per mil for benthic foraminifera at Walvis Ridge and Maud Rise, and about -4 per mil for planktonic foraminifera at Maud Rise. At the latter sites vertical gradients thus decreased, possibly at least partially as a result of upwelling. The oxygen isotope excursion was about -1.5 per mil for benthic foraminifera at Walvis Ridge and Maud Rise, -1 per mil for planktonic foraminifera at Maud Rise. The rapid oxygen isotope excursion at a time when polar ice-sheets were absent or insignificant can be explained by an increase in temperature by 4-6°C of high latitude surface waters and deep waters world wide. The deep ocean temperature increase could have been caused by warming of surface waters at high latitudes and continued formation of the deep waters at these locations, or by a switch from dominant formation of deep waters at high latitudes to formation at lower latitudes. Benthic foraminiferal post-extinction biogeographical patterns favour the latter explanation. The short-term carbon isotope excursion occurred in deep and surface waters, and in soil concretions and mammal teeth in the continental record. It is associated with increased CaC03-dissolution over a wide depth range in the oceans, suggesting that a rapid transfer of isotopically light carbon from lithosphere or biosphere into the ocean-atmosphere system may have been involved. The rapidity of the initiation of the excursion (a few thousand years) and its short duration (50 000 years) suggest that such a transfer was probably not caused by changes in the ratio of organic carbon to carbonate deposition or erosion. Transfer of carbon from the terrestrial biosphere was probably not the cause, because it would require a much larger biosphere destruction than at the end of the Cretaceous, in conflict with the fossil record. It is difficult to explain the large shift by rapid emission into the atmosphere of volcanogenic CO2, although huge subaerial plateau basalt eruptions occurred at the time in the northern Atlantic. Probably a complex combination of processes and feedback was involved, including volcanogenic emission of CO2, changing circulation patterns, changing productivity in the oceans and possibly on land, and changes in the relative size of the oceanic and atmospheric carbon reservoirs.

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The fossil floras described by Dieter MAI and Harald WALTHER are invaluable for understanding the past plant diversity in Europe, and provide important information on the occurrence of taxa in the fossil record that is critical for evolutionary studies. Among the taxa they recognized were seeds assigned to the extant genus Alpinia ROXB. (Zingiberaceae, Zingiberales). We reinvestigated 28 specimens that were assigned to Alpinia arnensis (CHANDLER) MAI, Alpinia cf. arnensis, and Alpinia bivascularis MAI from the Ypresian (lower Eocene) of the UK, upper Eocene of Germany, and lower Miocene of Germany using non-destructive synchrotron-based X-ray tomography to reveal internal anatomy. None of the samples studied show an anatomy consistent with extant Alpinia or even Zingiberales. The fossils lack the globose shape, often striate external surface, seed coat structure, operculum, and micropylar collar seen in all Alpinia, and lack the chalazal chamber seen in many Alpinia species. Two specimens from the lower Miocene of Germany showed the structure of fruits of Caricoidea CHANDLER (Cyperaceae) with a single-layered exocarp, thick mesocarp, and sclerified endocarp. The other specimens are recognized as Carpolithes albolutum nom. nov. (incertae sedis) from the Ypresian of the UK, C. phoenixnordensis sp. nov. (incertae sedis) from the upper Eocene of Germany, C. bivascularis comb. nov. (incertae sedis) from the lower Miocene of Germany as well as indeterminate tegmens from the lower Miocene of Germany. This reinvestigation demonstrates that there is, as yet, no confirmed fossil record for the extant genus Alpinia. Furthermore, at least four different taxa are recognized from what had been two extinct species, enhancing our understanding of these important European Cenozoic carpofloras.

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Extinction is a remarkably difficult phenomenon to study under natural conditions. This is because the outcome of stress exposure and associated fitness reduction is not known until the extinction occurs and it remains unclear whether there is any phenotypic reaction of the exposed population that can be used to predict its fate. Here we take advantage of the fossil record, where the ecological outcome of stress exposure is known. Specifically, we analyze shell morphology of planktonic Foraminifera in sediment samples from the Mediterranean, during an interval preceding local extinctions. In two species representing different plankton habitats, we observe shifts in trait state and decrease in variance in association with non-terminal stress, indicating stabilizing selection. At terminal stress levels, immediately before extinction, we observe increased growth asymmetry and trait variance, indicating disruptive selection and bet-hedging. The pre-extinction populations of both species show a combination of trait states and trait variance distinct from all populations exposed to non-terminal levels of stress. This finding indicates that the phenotypic history of a population may allow the detection of threshold levels of stress, likely to lead to extinction. It is thus an alternative to population dynamics in studying and monitoring natural population ecology.

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Using shells collected from a sediment trap series in the Madeira Basin, we investigate the effects of seasonal variation of temperature, productivity, and optimum growth conditions on calcification in three species of planktonic Foraminifera. The series covers an entire seasonal cycle and reflects conditions at the edge of the distribution of the studied species, manifesting more suitable growth conditions during different parts of the year. The seasonal variation in seawater carbonate saturation at the studied site is negligible compared to other oceanic regions, allowing us to assess the effect of parameters other than carbonate saturation. Shell calcification is quantified using weight and size of individual shells. The size-weight scaling within each species is robust against changes in environmental parameters, but differs among species. An analysis of the variation in calcification intensity (size-normalized weight) reveals species-specific response patterns. In Globigerinoides ruber (white) and Globigerinoides elongatus, calcification intensity is correlated with temperature (positive) and productivity (negative), whilst in Globigerina bulloides no environmental forcing is observed. The size-weight scaling, calcification intensity, and response of calcification intensity to environmental change differed between G. ruber (white) and G. elongatus, implying that patterns extracted from pooled analyses of these species may reflect their changing proportions in the samples. Using shell flux as a measure of optimum growth conditions, we observe significant positive correlation with calcification intensity in G. elongatus, but negative correlation in G. bulloides. The lack of a consistent response of calcification intensity to optimum growth conditions is mirrored by the results of shell size analyses. We conclude that calcification intensity in planktonic Foraminifera is affected by factors other than carbonate saturation. These factors include temperature, productivity, and optimum growth conditions, but the strength and sign of the relationships differ among species, potentially complicating interpretations of calcification data from the fossil record.

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The origins of some species of economic importance occurring over the Mediterranean Basin have been a traditional matter of debate that has important implications for land management. The case of Pinus pinea L. (Stone pine) is probably one of the most controversial, due to its documented long-term interaction with humans and its presence as a symbolic tree in certain areas of the Mediterranean (e.g., southwestern Iberia and Tuscany). Among the rest of the Mediterranean pines, several features make this pine unique (it has a characteristic crown shape, an edible kernel, cones that require three years to mature, and a very depauperate genetic diversity across its range). In addition, its palaeoecological information is rather limited, as the taxonomic precision attained by pollen analysts is insufficient for this tree and macroremains (such as kernels or anatomically well preserved wood) are needed to unequivocally detect the species in the fossil record. Recent findings of macrofossils of Pinus pinea in inland Iberia (Duero Basin) extend the late- Holocene range of the species, but the palaeobiogeographical information and the exhaustive genetic data available still suggest a very limited natural area (but still not sufficiently well defined) and a long and intense history of linkage to humans.

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En esta memoria de Tesis Doctoral se aborda el estudio paleobotánico de seis yacimientos tobáceos situados en las localidades burgalesas de Tubilla del Agua, Sedano, Herrán, Tobera y Frías, y en la alavesa de Ocio. El registro fósil encontrado en estos afloramientos se analiza de forma conjunta con el objetivo de conocer la evolución de la vegetación en el sector biogeográfico Castellano Cantábrico. Este sector se considera el territorio para el cual los hallazgos paleobotánicos son representativos y extrapolables, en tanto que constituye una región homogénea desde el punto de vista florístico, que abarca todos los yacimientos prospectados. El contexto temporal en el que se enmarca este estudio es el final del Cuaternario, desde el Pleistoceno Medio hasta la actualidad. Este intervalo se ha establecido a partir de la edad de los depósitos estudiados, la cual ha sido determinada —para los yacimientos de los que no se disponía de edades fiables— mediante la datación de muestras extraídas de las diferentes unidades litológicas identificadas. Para ello han sido empleadas las técnicas de carbono-14, desequilibrio de las series del uranio y racemización de aminoácidos. Los resultados geocronológicos obtenidos junto con el análisis geomorfológico de los yacimientos han permitido vincular la génesis de las 13 unidades litológicas identificadas con diferentes estadíos climáticos. Estos abarcan un amplio rango de condiciones ambientales, desde las más extremas del Último Máximo Glacial, hasta las más benignas de los Estadíos Isotópicos Marinos interglaciares 1 y 5. Como resultado de la prospección de los depósitos de toba fueron recuperados 1.820 fósiles, la mayoría impresiones foliares, pero también moldes de estróbilos femeninos, ramas y corteza, así como 42 carbones y restos subfósiles de Pinus sp. La identificación taxonómica de estos restos se ha realizado fundamentalmente a partir del análisis de caracteres diagnósticos morfológicos. Como resultado de ello, han sido descritos 28 taxones pertenecientes a las subclases Bryidae, Polypodiidae, Pinidae y Magnoliidae. La flora de los yacimientos estudiados se puede clasificar en tres grupos en función de sus requerimientos ecológicos: (i) uno formado por dos especies de alta tolerancia a la continentalidad —Pinus nigra y Quercus faginea—, las cuales aparecen bien representadas en la mayoría de los depósitos; (ii) otro constituido fundamentalmente por un conjunto de árboles y arbustos que habitualmente tienen el papel de especies acompañantes en los bosques ibéricos submediterráneos y eurosiberianos; y (iii) un tercer grupo compuesto por taxones hidrófitos o edafohigrófilos asociados al ecosistema del fitohermo activo y la vegetación de ribera. En el capítulo de Discusión se propone y analiza la hipótesis de que P. nigra y Q. faginea habrían sido las especies protagonistas de la vegetación zonal del sector Castellano Cantábrico durante el Cuaternario Final. Estas podrían haber persistido como tal incluso durante las épocas más frías, debido a su amplia valencia ecológica y a la capacidad de reproducirse vegetativamente en el caso del quejigo. Por el contrario, los taxones mesofíticos y eurosiberianos pudieron haber sufrido la expansión y retracción de sus poblaciones al ritmo de las oscilaciones climáticas. Sin embargo, la orografía diversa del sector Castellano Cantábrico proporciona emplazamientos en los que se combinan las diversas variables fisiográficas, de tal forma que pudieron haber existido microrrefugios en los que encontraron cobijo algunos taxones mesotérmicos y eurosiberianos durante los periodos glaciales. Por último, la historia evolutiva reciente de la vegetación de este territorio ha estado marcada por la acción antrópica, la cual empezó a ser manifiesta a partir del Neolítico. Esta se tradujo en la degradación y reducción de la cubierta forestal, así como en la extinción del pino laricio del Sector Castellano Cantábrico en los dos últimos milenios. ABSTRACT This PhD Dissertation focuses in the study of six tufa formations located nearby the villages of Tubilla del Agua, Sedano, Herrán, Tobera y Frías, all of them in the province of Burgos, and Ocio, which belongs to the province of Álava. We analyze the palaeobotanical archives of these sites with the purpose of unveiling and understanding the evolution of the vegetation of the Castilian Cantabrian biogeographical sector. This area is considered to be the territory that is represented in the palaeobotanical sample of the studied tufa archives. It is the homogeneous phitogeographical area with the lowest rank that include all the sites. The time frame of this study is the last part of the Quaternary, since the Middle Pleistocene to the present time. This interval is defined by the age of the tufa deposits, which were dated —for the ones that there were not available datings— throughout the analysis of 20 tufa samples taken from the 13 identified lithostratigraphic units. The age of the samples has been determined by using the methods of radiocarbon, U-Th dating and amino acid racemization. Chronological results, along with the chronostratigraphic study of the sites has allowed us to relate the build-up of the 13 identified lithostratigraphic units with different climatic stages. These structures were deposited in a wide range of climatic conditions, from the most extreme ones of the Last Glacial Maximum, to the warmer ones of the Marine Isotopic Stages 1 and 5. A total of 1,820 fossils were recovered from the tufa deposits, most of them were leaf impressions, but also pine cones, branches and bark moulds, along with charcoal and Pinus nigra macro remains. The taxonomical identification of these remains has been done mainly through the analysis of morphological traits. As a result of this process, 28 taxa belonging to the subclass of Bryidae, Polypodiidae, Pinidae and Magnoliidae were identified. The persistency of some taxa can be traced along different climatic stages in this fossil record. This fossil flora can be classified in three different groups: (i) the first one would be composed of two species with high continental climate tolerance —Pinus nigra y Quercus faginea—, which can be found in most of the deposits, (ii) the second group would be mostly formed by trees and shrubs that usually grow in the Iberian forests as an accessory species and (iii) the third one is composed of hydrophytes or hydrophilic taxa associate to the streams, riparian zones or the active tufa ecosystem. In the Discussion chapter we propose and analyse the hypothesis that P.nigra and Q. faginea were the main species of the zonal vegetation of the Castilian Cantabrian biogeographical sector during the last part of the Quaternary. This species could have persisted due to their wide ecological amplitude and also due to the capacity of asexual reproduction in the cases of the oak. On the other hand, mesophitic taxa could have suffered the retraction and expansion of their population following the climate oscillations. However, the diverse orography of the Castilian Cantabrian biogeographical sector provides a variety of combinations of physiographic variables, which could have been suitable refuges for some of the mesophitic taxa. The recent evolutionary history of the vegetation in this territory has been affected by human activities, which started to be relevant since the Neolithic. This led to a reduction of the forests and eventually, to the extinction of P. nigra in the Castilian Cantabrian biogeographical sector in the last two thousands of years.

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Analysis of several Salmonella typhimurium in vivo-induced genes located in regions of atypical base composition has uncovered acquired genetic elements that cumulatively engender pathogenicity. Many of these regions are associated with mobile elements, encode predicted adhesin and invasin-like functions, and are required for full virulence. Some of these regions distinguish broad host range from host-adapted Salmonella serovars and may contribute to inherent differences in host specificity, tissue tropism, and disease manifestation. Maintenance of this archipelago of acquired sequence by selection in specific hosts reveals a fossil record of the evolution of pathogenic species.

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Polycystine radiolaria are among few protistan groups that possess a comprehensive fossil record available for study by micropaleontologists. The Polycystinea and the Acantharea, whose skeletons do not become fossilized, were once members of the class “Radiolaria” (“Radiolaria” sensu lato: Polycystinea, Phaeodarea, and Acantharea) originally proposed by Haeckel but are now included in the superclass Actinopoda. Phylogenetic relationships within this superclass remain largely enigmatic. We investigated the evolutionary relationship of the Acantharea and the Polycystinea to other protists using phylogenetic analyses of 16S-like ribosomal RNA (rRNA) coding regions. We circumvented the need to culture these organisms by collecting and maintaining reproductive stages that contain many copies of their genomic DNA. This strategy facilitated extraction of genomic DNA and its purification from symbiont and prey DNA. Phylogenetic trees inferred from comparisons of 16S-like coding regions do not support a shared history between the Acantharea and the Polycystinea. However, the monophyly of the Acantharea and the separate monophyly of the Polycystinea (Spumellarida) are well supported by our molecular-based trees. The acantharian lineage branches among crown organisms whereas the polycystine lineage diverges before the radiation of the crown groups. We conclude that the Actinopoda does not represent a monophyletic evolutionary assemblage and recommend that this taxonomic designation be discarded.

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The past two decades have greatly improved our knowledge of vertebrate skeletal morphogenesis. It is now clear that bony morphology lacks individual descriptive specification and instead results from an interplay between positional information assigned during early limb bud deployment and its “execution” by highly conserved cellular response programs of derived connective tissue cells (e.g., chondroblasts and osteoblasts). Selection must therefore act on positional information and its apportionment, rather than on more individuated aspects of presumptive adult morphology. We suggest a trait classification system that can help integrate these findings in both functional and phylogenetic examinations of fossil mammals and provide examples from the human fossil record.