Internal Disorder: Post-Mortem Burrows within the Tests of the Holasteroid Echinoid Echinocorys Leske (Upper Cretaceous to Paleocene)

A test of the Lower Paleocene holasteroid echinoid Echinocorys ex. gr. scutata Leske from Göynük, Turkey, preserves a Planolites beverleyensis (Billings) burrow close to the internal surface. Identification of such a specimen is fortuitous; only where the test has broken cleanly away from the internal mould can such burrows be exposed, if present at all. A second specimen, from the Upper Cretaceous of Norfolk, England, bears a sinuous boring identified as Planolites montanus Richter.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Faunal remains from Porc-Epic: paleoecological and zooarchaeological investigations from a Middle Stone Age site in southeastern Ethiopia

The Middle Stone Age (MSA) sediments of Porc-Epic Cave near Dire Dawa, Ethiopia, have produced a large assemblage of archaeofaunal remains dating to the later periods of the Late Pleistocene. As one of the few MSA faunal assemblages from East Africa subject to detailed analysis, it provides rare insight into the foraging behavior of the early modern humans in this region. The MSA fauna include taxa that were not historically present in the area, with the occurrence of reduncines and alcelaphines indicating widespread grasslands and proximity to a perennial source of water. The faunal analysis examined several lines of evidence, including breakage patterns, surface modification, and skeletal-element representation. By incorporating comparisons with multiple actualistic data sets, the analysis of the assemblage demonstrates that, while the influence of carnivores and other biotic agents was evident, the faunal remains were accumulated primarily by hominid activity. The skeletal-element representation at Porc-Epic includes an abundance of high utility bones, in contrast to the predominance of "head-and-foot" elements documented at other MSA and Middle Paleolithic (MP) sites. The MSA occupants of Porc-Epic effectively exploited a wide range of prey with more emphasis on small- and medium-sized mammals.     

Lower Paleolithic hominin ecology at the fringe of the desert: Faunal remains from Bizat Ruhama and Nahal Hesi, Northern Negev, Israel

The Southern Levant is a pivotal area for the study of hominin paleoecology during the Lower Paleolithic, because of its location on the out-of-Africa dispersal route and its significant ecological diversity. Important information has been gained by archaeofaunal studies, which usually reveal that exploitation of diverse Mediterranean environments with woodlands, marshes and lake margins, represents a dominant subsistence strategy for Lower Paleolithic hominins. Here, we present new taxonomic and taphonomic data from two sites in the southern coastal plain of the Southern Levant, at the fringe of the Negev Desert: Bizat Ruhama (Early Pleistocene) and Nahal Hesi (Middle Pleistocene). The sites preserve anthropogenic faunas, with the former signaling a marrow-exploitation strategy, perhaps related to scavenging from carnivore kills, and the latter showing evidence for primary access to fleshed ungulate carcasses. The species composition of these Northern Negev sites is unique for the Levantine Lower Paleolithic in that these sites lack typical woodland and riparian species, probably indicating an open, relatively uniform environment with patchy water sources and trees, much like this semiarid region today. Bizat Ruhama and Nahal Hesi are among the only Levantine Lower Paleolithic faunas associated with such a setting, thereby widening the known spectrum of environments exploited by hominins in the region. It is suggested that the two sites, coupled with the nearby Late Pleistocene evidence, reflect a largely stable semiarid environment on the northwestern fringe of the Negev Desert throughout much of the Pleistocene.     

Ecological implications of the relative rarity of fossil hominins at Laetoli

Hominins are a very rare component of the large-mammal fauna at Laetoli. Although no equivalent data are available for Hadar, the much higher count and relative abundance of hominins suggests that they may have been more common at the latter site. The apparent relative rarity of hominins at Laetoli may have significant implications for understanding the ecology of Australopithecus afarensis. However, it is essential to first assess the extent to which taphonomic variables might have been a contributing factor. Using data from fossil ruminants, we show that the survivability of skeletal elements at Laetoli relates to the extent to which they can resist carnivore scavenging and their likelihood of being entirely buried by volcanic ashes and tuffaceous sediments. The rarity of hominins at Laetoli is probably due in part to the influence of these two taphonomic factors. However, these factors cannot account entirely for the difference in hominin relative abundance between these two sites, and ecological differences were probably a contributing factor. The highest population densities of chimpanzees today occur in forest and closed woodland, with reduced densities in open woodland. If similar levels of population-density variation characterized A. afarensis, the differences between Hadar and Laetoli may relate to the quality/optimality of the habitats. Hadar was, in general, much more densely wooded and mesic than Laetoli, with permanent and substantial bodies of water. In contrast, Laetoli was predominantly a woodland-shrubland-grassland mosaic supported only by ephemeral streams and ponds. The apparent greater relative abundance of hominins at Hadar compared with Laetoli suggests that, like chimpanzees, A. afarensis may have been more successful in more densely wooded habitats. Compared with Hadar, Laetoli probably represented a less optimal habitat for the foraging and dietary behavior of A. afarensis, and this is reflected in their inferred lower abundance, density, and biomass.     

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.     

Quaternary fossil fish from the Kibish Formation, Omo Valley, Ethiopia

The late Quaternary Kibish Formation of the Omo Valley, southwestern Ethiopia, preserves environments reflecting a history of fluctuations in the level of nearby Lake Turkana over the past 200,000 years. The Kibish Formation has yielded a diverse mammalian fauna (as well as birds and crocodiles), stone tools, and the oldest anatomically modern Homo sapiens. Fish, the most common vertebrate fossils in this unit, are reported in this study. Catfish (especially clariids and Synodontis) and Nile perch (Lates niloticus) predominate, but the gymnarchid Gymnarchus, a cyprinid (Barbus), tigerfish (Hydrocynus), pufferfish (Tetraodon), and other catfish are also present. In total, nine teleost genera are found in the Kibish Formation, representing a subset of the 37 genera that constitute the modern Omo-Turkana ichthyofauna. Several taxa present in the modern fauna, including Polypterus and members of the family Cichlidae, are not found in the Kibish deposits. Most specimens are preserved as disarticulated or broken skeletal elements, but some preservation of articulated elements (e.g., sets of vertebrae, crania with lower jaws or cleithra) also occurs. Many of the catfish and Nile perch specimens are larger than the largest reported from the modern river or lake. Faunas of Kibish Members I and III closely resemble one another; the fauna from Member IV contains only the three most common taxa (Clarias, Synodontis, Lates), though this may result from insufficient sampling. Barbed bone points have been collected from the upper part of the formation, indicating a long association between the human inhabitants and the fish fauna of the Omo Valley.     

Application of trophic transfer efficiency and age structure in the trophic analysis of fossil assemblages

We evaluate onshore-offshore trends in age-frequency distributions and trophic transfer efficiencies using 11 modern death assemblages off the Texas coast. Trophic transfer efficiencies within trophic levels offer little insight over that achieved by a size-frequency distribution. Production/biomass ratios will always be 1 in the fossil record. Within trophic-level estimates of paleogrowth efficiency, the ratio of paleoproduction to paleoingestion (Pⁱ_glt/Iⁱ_lt where i indicates the i^th trophic level and lt indicates the time-averaged value) follow the expected ecological trend precisely in that paleogrowth efficiency is consistently higher in primary consumers than in predators in all 11 death assemblages. Paleoutilization efficiency, the ratio of predator paleoingestion to prey paleoproduction, I²_lt^°/P¹_glt^°, may provide information on the degree of bias in the preservation of primary (1 °) and secondary (2 °) consumer trophic groups. I²_lt^°/P¹_glt^° fell below 0.1 in most cold-seep and bay assemblages, indicating a large surplus of primary consumers. In sharp contrast, I²_lt^°/P¹_glt^°     

Shell Beds as paleoecological puzzles: A case study from the Upper Permian of the Paraná Basin, Brazil

Summary Microstratigraphic, sedimentological, and taphonomic features of the Ferraz Shell Bed, from the Upper Permian (Kazanian-Tatarian?) Corumbataí Formation of Rio Claro Region (the Paraná Basin, Brazil), indicate that the bed consists of four distinct microstratigraphic units. They include, from bottom to top, a lag concentration (Unit 1), a partly reworked storm deposit (Unit 2), a rapidly deposited sandstone unit with three thin horizons recording episodes of reworking (Unit 3), and a shell-rich horizon generated by reworking/winnowing that was subsequently buried by storm-induced obrution deposit (Unit 4). The bioclasts of the Ferraz Shell Bed represent exclusively bivalve mollusks.Pinzonellaillusa andTerraia aequilateralis are the dominant species. Taphonomic analysis indicates that mollusks are heavily time-averaged (except for some parts of Unit 3). Moreover, different species are time-averaged to a different degree (disharmonious time-averaging). The units differ statistically from one another in their taxonomic and ecological composition, in their taphonomic pattern, and in the size-frequency distributions of the two most common species. Other Permian shell beds of the Paraná Basin are simílar to the Ferraz Shell Bed in their faunal composition (they typically contain similar sets of 5 to 10 bivalve species) and in their taphonomic, sedimentologic, and microstratigraphic characteristics. However, rare shell beds that include 2–3 species only and are dominated by articulated shells preserved in life position also occur. Diversity levels in the Permian benthic associations of the Paraná Basin were very low, with the point diversity of 2–3 species and with the within-habitat and basin-wide (alpha and gamma) diversities of 10 species, at most. The Paraná Basin benthic communities may have thus been analogous to low-diversity bivalve-dominated associations of the present-day Baltic Sea. The ‘Ferraz-type’ shell beds of the Paraná Basin represent genetically complex and highly heterogeneous sources of paleontological data. They are cumulative records of spectra of benthic ecosystems time-averaged over long periods of time (10²–10⁴ years judging from actualistic research). Detailed biostratinomic reconstructions of shell beds can not only offer useful insights into their depositional histories, but may also allow paleoecologists to optimize their sampling designs, and consequently, refine paleoecological and paleoenvironmental interpretations.     

Piero Leonardi (1908–1998)

Continental Trace Fossils, Stephen T. Hasiotis (with contributions from 11 others), 2002, SEPM Short Course Notes, no. 51, ii+132 pp., ISBN 1-56576-092-1 (paperback). US$62-00 (SEPM member price US$48-00). Reviewed by Stephen K. Donovan, Department of Palaeontology, Nationaal Natuurhistorisch Museum, Darwinweg 2, NL-2300 RA Leiden, The Netherlands     

Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda

This study provides a taphonomic analysis of the largest known sample of bone fragments collected from chimpanzee hunts. The entire sample consists of 455 bone fragments from 57 chimpanzee hunting episodes of 65 prey individuals at Ngogo, Kibale National Park, Uganda. It has low taxonomic diversity, consisting overwhelmingly of primates, especially red colobus monkeys. The age distribution of the prey remains is skewed towards pre-adults. Cranial bones are the dominant element, followed by long bones. Axial postcranial elements have low survivorship, with a complete absence of pre-caudal vertebrae. Bone is damaged in distinct ways, such as: destruction of long bone ends, typically with intact but chewed shafts; fragmentation and compression cracking of crania; and preservation of only the iliac blades of the innominates. Tooth marks are present but uncommon (4.4% of total NISP). These analyses enable us to: 1) describe and characterize consistent patterns of bone damage inflicted by chimpanzees across a much larger prey sample than has been previously studied; 2) make a preliminary comparison of the generalized chimpanzee taphonomic signature to that of leopard and eagle consumption of primates, as well as modern human consumption of small mammals; and 3) assess the utility of such samples for recognition of early hominin small mammal carnivory. We present a model that may be useful for detecting a pre-technological hominin carnivory and suggest some fossil locales at which close inspection of cercopithecoid remains for the above patterns might reveal traces of hominin hunting, though we caution that a pre-technological hominin hunted "assemblage" is not likely to be archaeologically visible.     

The palaeoecology of the dinosaurs of the Wessex Formation (Wealden Group, Early Cretaceous), Isle of Wight, Southern England

The Wessex Formation on the Isle of Wight yields an Early Cretaceous dinosaur fauna. Sedimentological evidence shows that this represents a mosaic of fluvial, floodplain and lacustrine environments within a relatively narrow east-west oriented valley. The vegetational cover on the alluvial plain had a savannah-or chaparral-like aspect, probably of low productivity. The relative scarcity of small aquatic vertebrates, absence of coals, abundance of oxidixed sediments and the presence of immature calcretes indicate seasonal water supply. The dinosaur taxa compising the Wessex Formation faunal assemblage represent a single palaeocommunity which inhabited the local alluvial plain, although some species may have been transient. The fauna had a relatively low diversity and this is attributed to the low productivity of the local vegetation. Iguanodontids and Hypsilophodon were the dominant elements in the fauna. In contrast to Late Jurassic dinosaur faunas, sauropods are less abundant in the Wessex Formation, although they remain taxonomically diverse. It is concluded that climatic changes which took place in the Late Jurassic or Early Cretaceous resulted in the appearance of low productivity vegetation and that this was incapable of supporting large sauropod populations.     

Paleolimnology of Neusiedlersee,Austria

Neusiedlersee is a shallow, alkaline lake (Table 1), which came into existance by tectonic subsidence about 12000–14000 yr ago. At present, half of it is covered by Phragmites australis which developed after the lake fell dry for the last time in 1868.Due to its astatic character, with about 100–200 dry periods since the lake came into existence, most of the sediment of the open lake overlying a thick Tertiary layer presents a mixture of terrestrial and lacustrine components. Because of disturbance by wind, this lacks stratification. Preliminary investigations, however, demonstrated that within the fringing Phragmites belt — especially landward — such a stratification does still exist. In order to learn more about the history of the lake 18 deep freeze cores of between 135 and 190 cms in depth, were collected from the Phragmites belt and investigated for chemical data (see Gunatilaka, this volume) and the succession of ostracods. Most of the cores contain Tertiary material (foraminifera and ostracods) at their base, followed by a cold water fauna as indicated by the presence of Cytherissa lacustris. Indicators of high alkalinity (i.e. the remains of male Limnocythere inopinata and high salinity (Heterocypris salina) were found in only a few cores.     

Biogenic and taphonomic processes affecting the development of shell assemblages: an actuopaleontological case study from mussel banks on North Sea tidal flats

Summary From spring 1997 to winter 2000, the composition and structure of epibenthic bivalve clusters and patches on a backbarrier tidal flat of the East Frisian Wadden Sea were continoously analyzed. The studied bivalve aggregates developed from intense and approximately simultaneous spatfalls of Mytilus edulis and Cerastoderma edule in 1996, after a severe winter (1995/96) with 64 days of driftice coverage. Spatfall occurred on a bare sandy tidal flat, whereas adjacent shell layers situated at the southern margin of the spatfall site were not colonized. Usually, such shell layers are the most common secondary hard substrates in the backbarrier area favorable for initial settlement of Mytilus larvae. The shell layers predominantly originate from erosion of former Mya arenaria and C. edule populations. So far, the studied bivalve aggregates are composed of approximately equal proportions of living specimens of both species, M. edulis and C. edule. The incorporated cockles were prevented from burrowing into the rediment by the coherent byssal mesh. In contrast to the results of previous studies, the great bulk of epibenthic living C. edule did survive for several years within the clusters. Therefore, the common terms ‘Mytilus banks’, ‘Mytilus patches’ or ‘Mytilus clumps’ may be too general and may erroneously imply a dominance of (or mono-specific composition by) M. edulis. In the fossil record, i.e. in diagenetically consolidated shell layers, the enforced epibenthic life-mode of C. edule and the remarkable species composition as well as the temporal development of the aggregates would not be recognizable.     

禄丰古猿化石产地沉积环境与埋藏学的初步研究

本文根据沉积物成因、古气候和古生物特征,把禄丰古猿化石产地新第三纪地层划分为晚中新世石灰坝组和早上新世庙山坡组。据岩性和岩相特征,可划分出五个不同的沉积阶段,每个阶段代表不同的沉积环境。古猿和其他脊椎动物化石主要埋在石灰坝组(四段)湖泊沼泽化沉积环境,次之为庙山坡组(五段)河流相沉积环境。