Risk and opportunity for humans coexisting with large carnivores

Models of Plio-Pleistocene hominid behavioral ecology often emphasize competition with large carnivores. This paper describes competition between modern humans and large carnivores in rural Uganda, including active, confrontational scavenging of carnivore kills by humans and carnivore attacks on humans. Information gathered from Ugandan Game Department archives (1923-1994) reveals that twentieth-century agropastoralists regularly tried to scavenge from leopard (Panthera pardus) and lion (Panthera leo) kills, and that these large carnivores have preyed on hundreds of humans in Uganda over the past several decades. Men were most often targets of carnivore attack, particularly while engaged in hunting-related activities. However attacks on men were less often lethal than attacks on women and children. Analyses show that lion attacks were more dangerous than leopard attacks. These data support recent contentions that hominids armed with even simple weapons can succeed in active, confrontational scavenging by chasing carnivores from kills. Hominids sharing East African habitats with large carnivores may have been regularly subject to attack.     

Does riparian habitat condition influence mammalian carnivore abundance in Mediterranean ecosystems?

The severe loss or degradation of riparian habitats has led to their impoverishment and impaired function, which may have severe consequences on both the riparian habitats themselves and their associated biota, including mammalian carnivores. We selected 70 riparian habitat reaches to evaluate the condition of the riparian habitats in southern Portugal and their use by carnivores. These sites were assessed for riparian condition using the stream visual assessment protocol (SVAP) and surveyed for carnivore presence along the riparian zones and across the surrounding matrix landscape, both in the wet (winter) and the dry (summer) season. Results show that carnivore surveys adjacent to riparian habitats consistently had significantly higher species richness than the matrix habitats, in both sampling seasons. Carnivore relative abundance and relative abundance of stone marten, common genet and Egyptian mongoose also showed higher values in riparian habitats, with significant differences in at least one season. The Eurasian badger, on the other hand, showed higher relative abundance values in the landscape matrix, though differences were not significant. The SVAP index ranked about 83% riparian reaches as poor or fair condition, and species richness was significantly higher in fair condition reaches during the wet season. These results reflect the importance of riparian habitats in Mediterranean ecosystems for mammalian carnivores. However, the generalized poor condition of these habitats suggests that direct measures for riparian restoration could be appropriate. The preservation or improvement of riparian habitats would certainly benefit the mammalian carnivore populations and consequently their conservation.     

Relative value of riparian woodlands in landscapes with different forest cover for medium-sized Iberian carnivores

This study analyses the role of riparian woodland in the conservation of five common carnivore species as compared with other non-riparian habitats according to woodland cover on the landscape scale (>60, 20–35 and <15% on 20 × 20-km surface area). I hypothesised that the importance of riparian woodlands in carnivore conservation on the regional scale would be greater in landscapes with low forest cover than in those with intermediate or high forest cover. To test this hypothesis, in each landscape type I sampled five riparian forests and 10–15 non-riparian habitats and recorded species richness and frequency of occurrence (number of sampled sites with species presence/total sampled sites) in both habitat types. The presence of (or use by) species at each particular sampling site was recorded using sign-surveys (search for scats and badger dens). The relative importance of riparian and non-riparian habitats in each landscape type was analysed by comparing species richness and frequency of occurrence for each species. Comparison of species richness and frequency of occurrence between landscape types indicate that riparian woodlands are important habitats in all landscape contexts. However, in accordance with the initial hypothesis, riparian woodlands are essential for carnivore conservation in the most deforested areas. Any agricultural or development policy should take into account the need to maintain and preserve riparian woodlands in landscapes, especially in intensively cultivated landscapes.     

Alternative uses of primates and carnivores in the reconstruction of early hominid behavior

A common approach to the reconstruction of early hominid behavior is the comparative study of living species. Some practitioners of this approach have strongly emphasized the use of carnivores while others have argued for the exclusive use of nonhuman primates. This paper takes the position that the interrelationship of primate and carnivore studies with respect to the reconstruction of early hominid behavior offers four possibilities. One is that the primate evidence is more useful and another is that the carnivore evidence is more useful. A third possibility is that primate and carnivore studies may supplement each other, i.e., taken separately, each body of evidence points toward the same conclusion. The fourth and most interesting posibility is that primate and carnivore studies may complement each other, i.e., taken together, they illuminate different aspects of a single hominid behavior pattern. It is argued that all four of these possibilities are applicable to the reconstruction of early hominid behavior, depending on the particular behavior pattern or research problem under consideration. This contention is supported with a series of examples. It is concluded that arguments favoring primates or carnivores should give way to discussions of how best to coordinate the data from these two valuable sources of inference about early hominid behavior.     

Disentangling Early Stone Age palimpsests: determining the functional independence of hominid- and carnivore-derived portions of archaeofaunas

Determining the extent to which hominid- and carnivore-derived components of fossil bone palimpsests formed independently of each other can provide valuable information to paleoanthropologists interested in reconstructing the foraging adaptations of hominids. Because stone tool cutmarks, hammerstone percussion marks, and carnivore tooth marks are usually only imparted on bone during nutrient extraction from a carcass, these bone surface modifications are particularly amenable to the types of analyses that might meet this goal. This study compares the percentage of limb bone specimens that preserve evidence of both hominid- and carnivore-imparted bone damage from actualistic control samples and several Plio-Pleistocene archaeofaunas, including new data from Swartkrans Member 3 (South Africa). We argue that this procedure, which elucidates the degree of hominid-carnivore independence in assemblage formation, will allow researchers to extract for focused analyses high integrity components (hominid and carnivore) from presumably low integrity sites. Comparisons suggest that the hominid- and carnivore-derived components from sites in Olduvai Gorge Bed II (Tanzania), the ST Site Complex at Peninj (Tanzania), and Swartkrans Member 3 formed largely independent of each other, while data from the FLK 22 Zinjanthropus (FLK Zinj) site (Olduvai Gorge Bed I) indicate significant interdependence in assemblage formation. This contrast suggests that some Early Stone Age assemblages (e.g., the Olduvai Gorge Bed II sites, the Peninj ST Site Complex, and Swartkrans Member 3) are probably more useful than others (e.g., FLK Zinj) for assessing the maximal carcass-acquiring abilities of early hominids; in such assemblages as those in the former set, sole hominid-contribution is more confidently discerned and isolated for analysis than in assemblages such as FLK Zinj.     

Modeling the formation of Early Stone Age artifact concentrations

The nature of stone artifact concentrations at early Plio-Pleistocene sites in East Africa is evaluated with regard to hominid transport behaviors responsible for their formation. These archaeological occurrences indicate ranging behaviors involving deliberate and repeated transport of flaked stone artifacts. The stone transported to archaeological sites within the time range of Homo habilis indicates planned transport of tools or material for tool manufacture to an extent far beyond transport behaviors reported among living apes, even stone hammer-using chimpanzees. Analysis of technological evidence in a lithic assemblage at a Plio-Pleistocene site at Koobi Fora (c. 1·5 ya) indicates on-site manufacturing activities and transport of flaked stone both to and from the site locale. Possible explanations for transport of stone artifacts are discussed in view of hominid strategies of environmental exploitation and resource utilization. A model is proposed for planned, habitual transport of artifacts by hominids positively correlated with distance of planned foraging range. In this model, larger-scale sites tended to develop at locales favorably located near abundant resources, where stone imports were high but export was relaxed due to the proximity of resources to be processed.     

Spanish late Pliocene and early Pleistocene hominid, palaeolithic and faunal finds from Orce (Granada) and Cueva Victoria (Murcia)

In SE Spain, recent excavations in the Orce basin and at Cueva Victoria indicate presence of both hominids and hominid activity from the Plio-Pleistocene boundary and early Lower Pleistocene.     

Brief communication: cutmarks on a plio-pleistocene hominid from Sterkfontein, South Africa

Cutmarks inflicted by a stone tool were observed on the right maxilla of Stw 53, an early hominid partial skull from Sterkfontein "Member 5" (South Africa). The morphology of the marks, their anatomical placement, and the lack of random striae on the specimen all support an interpretation of this linear damage as cutmarks. The location of the marks on the lateral aspect of the zygomatic process of the maxilla is consistent with that expected from slicing through the masseter muscle, presumably to remove the mandible from the cranium. Although radioisotopic dates are not available and relative faunal dating of the deposit from which Stw 53 derives is problematic, the morphology of the hominid skull suggests a Plio-Pleistocene age for the specimen. This therefore constitutes the earliest unambiguous evidence that hominids disarticulated the remains of one another.     

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.     

Computed tomography and enamel thickness of maxillary molars of Plio-Pleistocene hominids from Sterkfontein, Swartkrans, and Kromdraai (South Africa): An exploratory study.

This paper is one in a series which explores the possibility of using the non-destructive CT technique to identify patterns in tooth enamel distribution and structure of hominid molars from Plio-Pleistocene sites in South Africa, notably Swartkrans, Sterkfontein, and Kromdraai. Whereas previous investigators have emphasised gross differences in absolute and relative or average enamel thickness between hominid taxa, the present study highlights differences in enamel thickness over functionally significant regions of the crown. Differences in the distribution of enamel in A. robustus, A. africanus, and Homo sp. are identified through the use of bivariate and multivariate analyses, and are interpreted in terms of dietary regimes.     

Role of carnivores in the accumulation of the Sterkfontein Member 4 hominid assemblage: a taphonomic reassessment of the complete hominid fossil sample (1936-1999)

New taphonomic data on the Sterkfontein Member 4 (South Africa) fossil hominid assemblage are presented. The previous estimate of hominid individuals represented in the deposit (45) is increased to 87. New minimum numbers of hominid skeletal elements are provided, and incidences of bone surface damage inflicted by prehistoric biological agents are summarized. The hominid sample from Member 4 is composed predominately of gnathic remains and has a paucity of postcrania. This dearth of postcrania limits, to some extent, inferences about the formation of the Sterkfontein assemblage. However, carnivore tooth marks on some fossil specimens and an overall broad similarity in patterns of skeletal part representation between Sterkfontein and primate bone assemblages created by extant carnivores suggest that carnivores did have some involvement in the accumulation of the fossil hominid assemblage. Thus, this study provides support for the “carnivore‐collecting hypothesis” of Brain (Brain [ 1981 ] The Hunters or the Hunted? Chicago: University of Chicago Press), which implicates large carnivores as prominent collecting agents of hominid body parts in Sterkfontein Member 4. Evidence of bone surface damage is, however, too scant to make confident inferences about specific carnivore taxon/taxa involved in hominid bone collection at the site. Am J Phys Anthropol, 2004. © 2004 Wiley‐Liss, Inc.     

Toward an Ecological Model of African Plio-Pleistocene Hominid Adaptations

What was the environmental complex in which the Plio-Pleistocene hominids were evolving? What situations selected f o r increasing variation in hominid morphology? A n ap-preciation of ecological dynamics is important to develop answers to those questions. The circumstances that accompany periodically more severe semiarid successions appear to have promoted a shqt in early australopithecine morphology toward hyper-robust forms. Successional dynamics (including ameliorated conditions) may have resulted in repeated sympat y of potentially competing lines. Through the process of character divergence, some earliergracile populations appear to have emerged more Homo-like by the end of the Pliocene. Palaeodemographic analysis combined with detailed ecological modeling offers new possibilities f o r explorato y retrodictions concerned with common and divergent hominid adaptations. [australopithecines. palaeoecology, early Homo]     

Foramen magnum position variation in Pan troglodytes, Plio-Pleistocene hominids, and recent Homo sapiens: implications for recognizing the earliest hominids

The anteroposterior position of the foramen magnum distinguishes living Homo sapiens from apes, and has been used as evidence for the hominid status of numerable fossils in the history of human paleontology. During the past decade, foramen magnum position has been cited as evidence of the hominid status of Ardipithecus and Sahelanthropus. Specifically, the basion of Ardpithecus is reported to be inline with the bicarotid chord, while the basion of Sahelanthropus is reported to both touch the biporion chord and intersect the bicarotid chord. In order to assess the effectiveness of anteroposterior foramen magnum position in distinguishing hominids from nonhominid apes, this study examined whether or not the positions of biporion and bicarotid relative to basion sufficiently distinguished Pan troglodytes from recent Homo sapiens and Plio-Pleistocene hominids. The distances from basion to the biporion chord (BSBIP) and from basion to the bicarotid chord (BSBIC) were measured on samples of chimpanzee (n = 69) and recent human (n = 42) crania and a sample of Plio-Pleistocene hominid fossils (n = 8). The data were used to test the hypothesis that BSBIP and BSBIC measurements do not sufficiently distinguish P. troglodytes from hominids. While basion to biporion (BSBIP) does not effectively distinguish P. troglodytes from Plio-Pleistocene hominids and humans when used univariately, basion to bicarotid (BSBIC), when used univariately or bivariately with BSBIP, can be used to test whether or not an unknown specimen is a hominid. These results are used to evaluate the hominid status of Ardipithecus and Sahelanthropus.     

Taphonomic perspectives on hominid site use and foraging strategies during Bed II times at Olduvai Gorge, Tanzania

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1–2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3–5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3–5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.     

Long-term responses in population dynamics and diversity of small mammals in riparian and upland habitats within an agricultural landscape

Riparian zones in agricultural landscapes provide linear non-crop habitats for a variety of plant and mammal species, and hence are an important component of biodiversity. To date, variable responses of abundance, species richness, and species diversity of small mammals have been recorded in riparian and upland habitats. To address this variability, we provide a detailed analysis of seasonal changes in abundance and diversity of terrestrial small-mammal communities over a 7-year period within an agricultural landscape in south-central British Columbia, Canada. We tested the hypotheses (H) that abundance, species richness, and species diversity of communities of small mammals (H₁), and demographic parameters of reproduction, recruitment, and survival of the major species: deer mouse (Peromyscus maniculatus) and montane vole (Microtus montanus) (H₂), would be higher in riparian than upland habitats. Mean total abundance of small mammals was higher in summer and winter, and species richness higher in summer, in riparian than hedgerow habitats. Winter population data supported the total and species abundance patterns for small mammals, but species richness was similar, and diversity lower, in riparian than hedgerow sites during winter periods. Deer mice were the dominant species in terms of abundance and reproductive output for pregnancies and recruitment, but not survival, in riparian sites. Montane voles were similar in abundance and demographic parameters in the two habitats. House mice (Mus musculus) preferred hedgerows and wandering shrews (Sorex vagrans) riparian sites. Demographic parameters for deer mice and montane voles indicated that both riparian and hedgerow sites were “source” rather than “sink” habitats, and likely contribute to maintenance of mammal diversity in agricultural landscapes.     

Vulnerability to xylem cavitation and the distribution of Sonoran Desert vegetation

We studied 15 riparian and upland Sonoran desert species to evaluate how the limitation of xylem pressure (Ψ(x)) by cavitation corresponded with plant distribution along a moisture gradient. Riparian species were obligate riparian trees (Fraxinus velutina, Populus fremontii, and Salix gooddingii), native shrubs (Baccharis spp.), and an exotic shrub (Tamarix ramosissima). Upland species were evergreen (Juniperus monosperma, Larrea tridentata), drought-deciduous (Ambrosia dumosa, Encelia farinosa, Fouquieria splendens, Cercidium microphyllum), and winter-deciduous (Acacia spp., Prosopis velutina) trees and shrubs. For each species, we measured the "vulnerability curve" of stem xylem, which shows the decrease in hydraulic conductance from cavitation as a function of Ψ(x) and the Ψ(crit) representing the pressure at complete loss of transport. We also measured minimum in situ Ψ(x)(Ψ(xmin)) during the summer drought. Species in desert upland sites were uniformly less vulnerable to cavitation and exhibited lower Ψ(xmin) than riparian species. Values of Ψ(crit) were correlated with minimum Ψ(x). Safety margins (Ψ(xmin)-Ψ(crit)) tended to increase with decreasing Ψ(xmin) and were small enough that the relatively vulnerable riparian species could not have conducted water at the Ψ(x) experienced in upland habitats (-4 to -10 MPa). Maintenance of positive safety margins in riparian and upland habitats was associated with minimal to no increase in stem cavitation during the summer drought. The absence of less vulnerable species from the riparian zone may have resulted in part from a weak but significant trade-off between decreasing vulnerability to cavitation and conducting efficiency. These data suggest that cavitation vulnerability limits plant distribution by defining maximum drought tolerance across habitats and influencing competitive ability of drought tolerant species in mesic habitats.     

Invasive American Mink: Linking Pathogen Risk Between Domestic and Endangered Carnivores

Infectious diseases, in particular canine distemper virus (CDV), are an important threat to the viability of wild carnivore populations. CDV is thought to be transmitted by direct contact between individuals; therefore, the study of species interactions plays a pivotal role in understanding CDV transmission dynamics. However, CDV often appears to move between populations that are ecologically isolated, possibly through bridge hosts that interact with both species. This study investigated how an introduced species could alter multihost interactions and act as a bridge host in a novel carnivore assemblage of domestic dogs (Canis familiaris), invasive American mink (Neovison vison), and threatened river otters (Lontra provocax) in southern Chile. We found that rural dogs interact with mink near farms whereas in riparian habitats, minks and river otters shared the same latrines with both species visiting sites frequently within time intervals well within CDV environmental persistence. No interactions were observed between dogs and otters at either location. Both dog and mink populations were serologically positive for CDV, making the pathogen transfer risk to otters a conservation concern. Altogether, introduced mink in this ecosystem have the potential to act as bridge hosts between domestic dogs and endangered carnivores.     

Hominid carpal,metacarpal, and phalangeal bones recovered from the Hadar formation: 1974–1977 collections

The fossil hominid hand bone collection from the Pliocene Hadar Formation, Ethiopia, is described anatomically. These hand bones, all from A.L. (Afar Locality) 333 and 333w, constitute the largest sample of hominid manus remains thus far recovered from the Plio-Pleistocene of Africa.     

Gladysvale: First early hominid site discovered in South Africa since 1948

We report here the discovery of fossil hominid teeth at Gladysvale, near Johannesburg in the southern Transvaal. This find makes the site the seventh in South Africa to yield australopithecine remains and the first new early hominid-bearing locality to be found in this region since 1948. Apart from the hominid specimens, our excavations at Gladysvale have added appreciably to the abundant Plio-Pleistocene fauna previously recorded from the cave deposit. The fauna indicates that savanna conditions prevailed during deposition of at least part of the fill. Preliminary faunal dating gives an age of deposition of between c1.7 and c2.5 mya. © 1993 Wiley-Liss, Inc.     

Evidence of cave use by savanna chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) at Fongoli,Senegal: implications for thermoregulatory behavior

Much attention has been paid to how humans both adapt and acclimate to heat stress, primarily due to the relevance of these issues to hominid evolution in open Plio-Pleistocene environments. However, little is known about the responses of human’s closest living relative, the chimpanzee (Pan troglodytes), to similar environmental stressors. In southeastern Senegal, one of the hottest and driest habitats that chimpanzees (P. t. verus) live in today, apes rely on behavioral mechanisms of dealing with thermal stress. Chimpanzees’ use of caves was based primarily on indirect evidence (feeding traces, feces, and hairs) gathered from one cave from January to December 2004, but data from observational records collected from May 2001 through March 2006 supplement these data. The hypothesis that chimpanzees’ use of caves is a response to heat was tested by collecting data on temperatures within the largest cave and in different habitats used by chimpanzees, such as gallery forest and woodland. Results indicate that chimpanzees primarily use caves as shelters during the hottest times of year and that caves are consistently and significantly cooler than open habitats. Insight into the way that chimpanzees in Senegal cope with extreme temperatures may help us to better understand the behavior of early hominids in such an environment.