Hominid paleoecology: The competitive exclusion principle and determinants of niche relationships

Some paleonanthropologists invoke the competitive exclusion principle programmatically in support of a single-species (or lineage) hypothesis of hominid evolution. Others apparently accept the association between competitive exclusion and a single species view, and develop multi-species interpretations using only taxonomic concepts. This paper demonstrates that the competitive exclusion principle itself is too assumption-bound to be appropriate for analyzing questions of hominid coexistence. Especially, the principle does not establish the ecological validity of the single-species hypothesis. Contrary to its immediate appearances, the principle has developed historically and analytically to predict and explain the evolution and maintenance of diversity in communities. Used in this manner it constitutes an important basis for the study of hominid paleoecology. Niche concepts necessary to demonstrate this are introduced, and competition and other factors that may have influenced the realized niches of sympatric hominid species are discussed. Nearly all modifications of the competitive exclusion principle that make it more realistic also illuminate factors that generate stable coexistence among competing species. These factors and their organizing theory establish the relevance of a broad data base to the analysis of hominid paleoecology; they should help to guide research on the ecology of early homonid species and their interrelationships.     

Hominid paleoecology and competitive exclusion: Limits to similarity,niche differentiation,and the effects of cultural behavior

Fossil evidence from the Plio-Pleistocene of Africa apparently has confirmed a multi-lineage interpretation of early hominid evolution. Empirical refutation of the single species hypothesis must now be matched to the evolutionary ecology theory, which can underwrite taxonomic assessment and help to explain sympatric hominid coexistence. This paper contributes to that goal by reassessing the ecological rationale provided for the single-species hypothesis. Limiting similarity concepts indicate that the allowable ecological overlap between sympatric competitors is greater than the degrees of metric overlap often advanced as standards for identifying fossil species. Optimal foraging theory and the compression hypothesis show that the initial ecological reaction of a hominid to a sympatric competitor would likely be micro-habitat divergence and possibly also temporal differentiation of resource use. The long-term, evolutionary response is niche divergence, probably involving diet as well. General niche partitioning studies suggest that diet and habitat are the most common dimensions of niche separation, although temporal separation is unusually frequent in carnivores. The equation of niche with culture, basic to the single-species hypothesis, has no analytic meaning. Finally, four minor points are discussed, suggesting that (a) extinction is not unlikely, even for a long-lived and competitively competent hominid lineage, (b)parsimony is fickle, (c)interspecific mutualism may jeopardize survival, and (d)generalists are subordinate competitors, but for hominids, seemingly, successful ones. I argue that analog models of hominid paleoecology should be replaced by the use of zoological and anthropological observations to assess the generality and reliability of ecological theory and comcepts that may encompass early hominids.     

The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment

The reconstructed taphonomic and paleoenvironmental contexts of a ca. 4 million-year-old partial hominid skeleton (Stw 573) from Sterkfontein Member 2 are described through presentation of the results of our analyses of the mammalian faunal assemblage associated stratigraphically with the hominid. The assemblage is dominated by cercopithecoids (Parapapio and Papio) and felids (Panthera pardus, P. leo, Felis caracal, and Felidae indet.), based on number of identified specimens, minimum number of elements and, minimum number of individuals. In addition, the assemblage is characterized by a number of partial skeletons and/or antimeric sets of bones across all taxonomic groups. There is scant indication of carnivore chewing in the assemblage. These observations, in addition to other taphonomic data, suggest that the remains of many animals recovered in Member 2 are from individuals that entered the cave on their own-whether accidentally by falling through avens connecting the cave to the ground surface above or by intentional entry-and were then unable to escape, rather than primarily through systematic collection by a biotic, bone-accumulating agent. The taphonomic conclusion that animals with climbing proclivities (i.e., primates and carnivores) are preferentially preserved over other taxa, ultimately because of those proclivities, urges caution in assessing the fidelity of the assemblage for reconstruction of the Member 2 paleoenvironment. With that caveat, we infer that the Member 2 paleoenvironment was typified by rolling, rock-littered and brush- and scrub-covered hills, indicated by the abundant F. caracal and cercopithecoid fossils recovered and the identified presence of the extinct Caprinae Makapania broomi. In addition, the valley bottom may have retained standing water year-round, perhaps supporting some tree cover--a setting suitable for the well-represented ambush predator P. pardus and suggested by the presence of Alcelaphini. Finally, the reconstructed taphonomic and paleoenvironmental settings of Sterkfontein Member 2 are compared to penecontemporaneous sites in South and East Africa.     

Evolution of Hominid Femur and Tibia: A morphometric approach to the evolutionary research in anthropology

Some theoretical and methodological morphometrical approaches in evolutionary anthropology and paleoanthropology are reviewed in this study. It is shown which are the contemporary possibilities of sophisticated biometrical and biostatistical methods and the role of the morphometrical approach. A new approach, experimental morphometrics, is presented, reflecting recent trends in evolutionary morphology as well as sophisticated biostatistical methods. The approach emphasizes the complex inter-related approach to the data processing and a double nature of morphometric data, i.e. biological and biostatistical one. The practical use of experimental morphometry is given for the two examples of analyses of the evolution of the hominoid and hominid femur and tibia. The hypothesis on a two stage restructuring of morphology of the hominid femur and tibia is supported by experimental results. Two different steps during this restructuring could be recognized: 1) Structural remodelling typical for the origin of hominids and australopithecine evolution, and 2) proportional remodelling of lower limb long bones which is connected with the Australopithecus/Homo transition (i.e. mainly Homo habilis stage). The results confirm the increasing trend of bipedal adaptations on the early hominid lower limb skeleton. Analysis of microevolutionary trends on the Homo sapiens femur and tibia indicates at least three different morphological patterns, Paleolithic, Neolithic and Recent, with numerous specific features in morphology and proportions. Neanderthal morphology is very derived. Upper Paleolithic/Mesolithic/Neolithic transition has a key character for the understanding of post-Paleolithic morphology. A very high sexual dimorphism of the femur and tibia has been demonstrated for Upper Paleolithic and Neolithic populations. Presented at the Foundation of Different Approaches to the Study of Human Evolution edited by B. Sigmon & V.V. Leonovicova-Liblice, September 1–3, 1989     

Biostratigraphy and biochronology of the Late Miocene of central Argentina: Evidence from rodents and taphonomy

Biostratigraphy and biochronology of the Late Miocene of central Argentina is analyzed through evolutionary patterns of octodontoid rodents. The studied faunas were found in continental sediments assigned to the Cerro Azul and Saldungaray formations (La Pampa and Buenos Aires provinces, respectively). Taphonomic attributes of recovered mammals, linked mostly to the development of paleosols or to predators’ activities, constraint the deposition lapse to short intervals. A biostratigraphic scheme based mainly on the anagenetic pattern of change in the octodontoid rodent lineage Chasichimys-Xenodontomys is proposed. Since there is no stratigraphic superposition among the studied levels, relationships among deposits are established through the polarity evidenced by the stage of evolution of chronomorphs of this lineage. Five new biozones for the Late Miocene of central Argentina are recognized. Evidence of other anagenetic sequences, and general trends of increasing hypsodonty in octodontoids, support this proposal. The results enhance the value of the “stage of evolution” concept as a correlation tool in biostratigraphic and biochronologic studies, mainly when applied through phyletic sequences.     

Vertebrate taphonomy in loess-palaeosol deposits: A case study from the late Miocene of central Argentina

This paper deals with the taphonomic analysis of the late Miocene bone assemblage from the Cerro Azul Formation at Telén (La Pampa Province, Argentina). The faunal assemblage was assigned to the Huayquerian mammal age (late Miocene). The fossiliferous section shows a homogeneous lithology, and is interpreted as a loess deposit with two similar and slightly developed palaeosols, classified as calcic vertisols. The studied sample comprises 5598 remains anatomically and taxonomically determined. They were collected from an area of about 48,000 m², appearing randomly distributed through the section and with low density. Most remains are small- to very small-sized, disarticulated, and very fragmented. Different taphonomical histories are inferred for microvertebrates and macromammals. The microvertebrate assemblage is interpreted as the result of predator activities. After a brief period of pre-burial exposure, remains were dispersed from the original depositional area. On the other hand, a natural and gradual death process is envisaged for macromammals, followed by a long period of exposure to weathering and dispersal by physical agents. Remains of both groups, once buried, suffered the diagenetic processes of the host rock. Consequently, the fossil assemblage from Telén would represent a condensed assemblage corresponding to two distinct time spans, i.e., the accumulation of microvertebrates took place in a short time interval whereas that of macromammals occurred over a longer period, coincident with the development of both soils.     

Systematics and taphonomy of an Early Kimmeridgian belemnite fauna from the Mediterranean Tethys (Monte Nerone, Central Apennines, Italy)

About three hundred belemnite rostra were collected from Lower Kimmeridgian beds of a structural high sequence cropping out at Mt. Nerone (central Apennines, Pesaro Province, Italy). The belemnite fauna is composed mainly of new species. Nine species were recognised, ascribed to five genera, which include Hibolithes semisulcatusMünster, 1830; H. pignattii nov. sp.; Acutibelu0s sp. cf. acuariformisRiegraf, 1981; Belemnopsis neronensis nov. sp., Duvalia matteuccii nov. sp., D. nicosiai nov. sp., D. pallinii nov. sp., D. raymondi nov. sp. and Rhopaloteuthis massimoi nov. sp.; moreover a single specimen is treated in open nomenclature as Belemnopseidae incertae sedis. The stratigraphic and palaeobiogeographic significance of the new fauna is discussed. The taphonomy of the belemnite-rich level is described, with reference to borings found on the belemnite rostra.La fauna a belemniti risulta composta quasi esclusivamente da specie nuove. Sono state riconosciute le seguenti specie: Hibolithes semisulcatusMünster, 1830; H. pignattii nov. sp.; Acutibelus sp. cf. acuariformisRiegraf, 1981, Belemnopsis neronensis nov. sp., Duvalia matteuccii nov. sp., D. nicosiai nov. sp., D. pallinii nov. sp., D. raymondi nov. sp. and Rhopaloteuthis massimoi nov. sp. Inoltre un esemplare è stato discusso con nomenclatura aperta come Belemnopseidae incertae sedis. Alcuni nuovi dati stratigrafici e paleobiogeografici sono stati analizzati ed integrati con i dati della letteratura. E’ stata anche esaminata la tafonomia del livello a belemniti e le perforazioni presenti sulla superficie dei rostri.     

Population paleoecology and comparative taphonomy of two edrioasteroid (echinodermata) pavements: Upper ordovician of kentucky and Ohio

Two strophomenid brachiopod shell pavements bearing abundant edrioasteroid epizoans occur stratigraphically within the top 4.5 m of the Corryville Formation (Upper Ordovician, Maysvillian) in Boone County, Kentucky, and Hamilton County, Ohio. Both assemblages are dominated by Isorophus cincinnatiensis and contain Carneyella pilea and Streptaster vorticellatus as lesser constituents. Pedicle exterior valves of Rafinesquina “alternata”; are the preferred substratum for all species at both sites, but the Kentucky population occurs mainly on unabraded, articulated shells and the Ohio population occupies abraded, disarticulated shells. The Kentucky population includes a greater size range of Isorophus (at least three cohorts) than the Ohio population (at least two cohorts), suggesting a greater time span of colonization for the Kentucky site. Host shells from Kentucky have anterior‐posterior axes strongly aligned NE‐SW whereas the Ohio shells are unoriented. Isorophus in Kentucky show a preference for the anterior margin of the Rafinesquina pedicle exterior valve, whereas Isorophus in Ohio show no preferred location on host shells. The Kentucky edrioasteroids are inferred to have attached to mainly live Rafinesquina in brachial‐valve‐up position. Host shells were inverted and oriented by a NE‐SW storm surge, then buried rapidly. Ohio edrioasteroids occupied mainly dead, already inverted host shells not oriented by storm surge possibly because of their more downslope (distal) location. Comparative taphonomic relationships between the two coeval populations are consistent with a model of proximal to distal distribution along a gentle shallow subtidal N‐S paleoslope subject to episodic storm activity.     

The fossil record and taphonomy of butterflies and moths (Insecta,Lepidoptera): implications for evolutionary diversity and divergence-time estimates

Background

It is conventionally accepted that the lepidopteran fossil record is significantly incomplete when compared to the fossil records of other, very diverse, extant insect orders. Such an assumption, however, has been based on cumulative diversity data rather than using alternative statistical approaches from actual specimen counts.

Results

We reviewed documented specimens of the lepidopteran fossil record, currently consisting of 4,593 known specimens that are comprised of 4,262 body fossils and 331 trace fossils. The temporal distribution of the lepidopteran fossil record shows significant bias towards the late Paleocene to middle Eocene time interval. Lepidopteran fossils also record major shifts in preservational style and number of represented localities at the Mesozoic stage and Cenozoic epoch level of temporal resolution. Only 985 of the total known fossil specimens (21.4%) were assigned to 23 of the 40 extant lepidopteran superfamilies. Absolute numbers and proportions of preservation types for identified fossils varied significantly across superfamilies. The secular increase of lepidopteran family-level diversity through geologic time significantly deviates from the general pattern of other hyperdiverse, ordinal-level lineages.

Conclusion

Our statistical analyses of the lepidopteran fossil record show extreme biases in preservation type, age, and taxonomic composition. We highlight the scarcity of identified lepidopteran fossils and provide a correspondence between the latest lepidopteran divergence-time estimates and relevant fossil occurrences at the superfamily level. These findings provide caution in interpreting the lepidopteran fossil record through the modeling of evolutionary diversification and in determination of divergence time estimates.

Electronic supplementary material

The online version of this article (doi:10.1186/s12862-015-0290-8) contains supplementary material, which is available to authorized users.     

Comparative taphonomy of three bivalve species from a mass shell accumulation in the intertidal regime of North Sea tidal flats

This study comprises a comparative taphonomic analysis of three endobenthic bivalves (Mya arenaria, Cerastoderma edule, and Macoma balthica) derived from a mass accumulation of mainly vertically packed shells from the “Wurster Watt” in Lower Saxony’s Wadden Sea, German Bight. Bulk samples from two transects were analyzed with respect to taxonomic composition, left/right valve presence, counts and weight percentages of taxa, and size-frequency distributions. Taphonomic features including abrasion, fragmentation, encrustation and bioerosion were subjected to a semi-quantitative analysis. Taphonomic results show significant differences with respect to bivalve taxa as well as between transects. Mya arenaria, a large endobenthic bivalve, shows the greatest amount of fragmentation and is often encrusted by balanids and bryozoans. The smaller and more robust Cerastoderma edule has the greatest values for bioerosion especially by the polychaete Polydora ciliata. Macoma balthica, which has the thinnest valves, shows the highest values for abrasion, but low values for the rest of the measured taphonomic features. Taphonomic differences between the taxa, samples and transects are related to (1) the origin of the bivalves (from living populations or exposed colonization horizons), (2) the different size and morphology of the valves (themselves related to mode of life), (3) the taphonomic trajectories of the different bivalves, as well as (4) the varying depositional environment of the two transects.     

Experimental taphonomy in museums: Preparation protocols for skeletons and fossil vertebrates under the scanning electron microscopy

Natural history institutions use different standards and methods to prepare and preserve skeletons of both modern and fossil vertebrates. These standards and treatments may be considered as taphonomic agents of museum collections. Modifications may occur during preparation, affecting the integrity of bone tissues and skeletons or their molecular and geochemical composition. The present evaluation uses a Scanning Electron Microscope (SEM) to analyze the most common preparation methods and standards. The aim of this paper is to evaluate preparation methods to determine those that provide the best SEM observations and those that best preserve osseous tissues and least damage fossil surfaces. In addition, the study of these alterations, produced under known conditions, provides criteria, which aids recognition of modifications that may occur in nature.     

Linking taphonomy to community-level abundance: Insights into compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps)

Although actualistic live/dead comparisons lead to robust estimates of fidelity of modern death assemblages, quantitative evaluation of fidelity of fossil assemblage remains uncertain. In this paper, effects of storm reworking on compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps, Austria) are evaluated. An exploratory approach is based on comparison of reworked and non-reworked assemblages in ordination analyses. Non-reworked assemblages of one or more communities provide a baseline for evaluation of fidelity of reworked assemblages. In siliciclastic-rich intervals of the Kössen Formation, shell concentrations are represented by (1) packstones with small, shallow infaunal bivalves, (2) floatstones and pavements with large semi-infaunal bivalves, and (3) bioclastic marlstones. In carbonate-rich intervals, bioclastic floatstones with bivalves and brachiopods occur. Analyzing all shell concentrations, eight sample groups sharing similar species composition are discriminated. Limited effect of storm reworking on composition of shell concentrations is indicated by (1) a general persistence of six sample groups when only non-reworked assemblages are analyzed, (2) similarity in composition between reworked and non-reworked assemblages within sample groups, and (3) compositional segregation between non-reworked assemblages of distinctive sample groups, mostly without any reworked assemblages of intermediate composition.Depth-related variations in dead-shell production, shell destruction and body size governed preservation and distribution of the shell concentrations along onshore-offshore gradient in the Kössen Basin. First, at times when environmental conditions were unfavorable for shell producers, coupled with high background shell destruction rates, limestone beds formed during storm events were shell-poor. Second, less common shell concentrations in upper than in lower parts of siliciclastic intervals can be related to higher environmental stress in shallower habitats. Third, the difference between shell concentrations dominated by small and large bivalves is driven by between-habitat differences in body size and is not due to a differential sorting of small and large shells. Combining community analysis based on species abundances with taphonomic analysis can thus be helpful in tracking fidelity of fossil assemblages.     

Submarine foraging behavior of alcids in an artificial environment

We used an artificial environment at Sea World, Inc, San Diego, to study underwater foraging behavior of alcids. Larger birds dove longer and had greater wingbeat frequencies. The pigeon guillemot Cepphus columba was the only species to use both feet and wings for propulsion; all others used just wings. Aggressive interactions underwater were common. Competition among alcids in the wild may occur primarily underwater, and artificial environments may be the best means to study such interactions.     

Mechanical advantages of the Neanderthal thumb in flexion: a test of an hypothesis

It has been proposed that the pollical phalangeal length proportions of the Neanderthals provided them with a greater mechanical advantage relative to recent humans for their pollical flexor muscles in power grips across the interphalangeal (IP) joint at the expense of the mechanical advantage of those pollical flexor muscles in precision grips at the finger tip. To test these related hypotheses, we compared the pollical load arm dimensions (phalanx lengths) to power arm dimensions (dorsopalmar articular heights) for the European and Near Eastern Neanderthals and for European and Amerindian samples of recent humans. It was found, initially, that the proximal articular height of the pollical distal phalanx is a poor predictor of the power arm at the IP articulation, even though the proximal articular height of the pollical proximal phalanx was an adequate indicator of the power arm size at the metacarpophalangeal (MCP) joint. In addition, differences in distal pollical ulnar deviation at the IP joint appeared to make little difference in the mechanical advantage comparisons. More importantly, the relative shortness of Neanderthal proximal pollical phalanges and the relative lengthening of their distal pollical phalanges was confirmed, and it was determined that, despite some minor differences in articular dimensions between Neanderthals and recent humans, these pollical phalangeal length contrasts translated into significant differences in mechanical advantages for the flexor muscles across the MCP and IP articulations.     

Fossil analogues of present-day Cladocora caespitosa coral banks: Sedimentary setting, dwelling community, and taphonomy (Late Pliocene, W Mediterranean)

 Cladocora caespitosa is a common zooxanthellate, ahermatypic, constructional scleractinian coral in shallow waters of the present-day Mediterranean. Extensive coral banks in Upper Pliocene shallow marine deposits of the Almería-Níjar Basin (SE Spain) contain the same species. These banks occur on debris-flow conglomerates deposited in a fan delta, or on bioclastic accumulations interpreted as storm deposits. Direct relationships of coral beds with coastal facies indicate that C. caespitosa colonized shallow settings near the paleocoast, probably not deeper than 20–30 m. Low turbulence allowed corals to colonize substrates, which remained stable for long periods. Activity of organisms in the coral community, storms, and detritic discharges from the fan delta were the most significant mechanisms disturbing the coral development. The hard substrata provided by coral banks promoted colonization by cemented and epibyssate organisms. Coral banks marked maximum flooding surfaces at the end of transgressive systems tracks. They were suddenly buried by sediment input into the basin. Taphonomic signatures measured on components of the coral bank communities indicate a low turbulence environment, probably a bay. The low hydraulic energy further inhibited post-burial reworking, thus promoting the in situ preservation of a great part of the organisms inhabiting the bioconstructions. Accepted: 2 December 1997     

A physio-anthropological approach in evaluation of human adaptability to living environment: in the case of artificial light environment

Attention has been focused on human adaptability to human-adopted artificially modern and comfortable environments which may not always match the adaptability by the physiological state of humans. This discrepancy was found to induce a slight tension in the human body on evaluation with a physio-anthropological approach. Although a standard methodological approach has yet to be established, the present study attempted to evaluate human adaptability to artificial light environment, based on evaluations from three major possible perspectives: a) central nervous system, b) autonomic nervous system and c) biological rhythm. In order to detect the slight tension, human volunteers were exposed to an artificial light environment, and the appropriate physiological parameters were then examined from photic signal pathways in a brain. The findings revealed that a higher color temperature of fluorescent lamps induced a slight tension, although many other factors remained to be elucidated.     

Carnivore alteration of human bone from a late prehistoric site in Illinois

The nature and pattern of destruction of human bone by scavenging animals, probably canids, is described for 30 Oneota skeletons from a west-central Illinois cemetery dated at ca. A.D. 1300. The most frequently damaged parts of the skeleton include bones covered by relatively little soft tissue, such as the major joints of the limbs, and those in the facial, abdominal, and gluteal regions.     

Adaptations of the IGF system during malignancy: human skeletal muscle versus the systemic environment.

Presented in this study are data derived from a unique cohort of patients both with and without cancer, for whom we not only have serum samples, allowing us to investigate systemic factors impacting on skeletal muscle maintenance, but also primary skeletal muscle cultures giving us a model to mimic the in vivo muscle milieu. Possible local effects of autocrine/paracrine and endocrine IGF system components impacting on myoblast growth and differentiation could therefore be assessed. We report for the first time that the decrease in myoblast stem cell numbers seen with normal aging is lost in cancer patients. We further report that serum IGF-I, IGF-II and IGFBP-3 all show positive correlations with myoblast retrieval in control patients, but that with the exception of IGFBP-3 these correlations are lost in malignancy. Indeed IGF-II switches to a negative correlation with myotube formation in malignancy. Furthermore we provide initial evidence to suggest that there is an apparent altered regulation of local IGFBP-3 production during malignancy which may enable satellite cell proliferation, stem cell infiltration or both. Finally we show the importance of investigations not only monitoring the systemic impact of serum factors on skeletal muscle responses but also critically assessing the role that locally produced muscle IGFBP-3 may have on the systemic environment.