Pleistocene Hominins as a Resource for Carnivores: A c. 500,000-Year-Old Human Femur Bearing Tooth-Marks in North Africa (Thomas Quarry I,Morocco)

In many Middle Pleistocene sites, the co-occurrence of hominins with carnivores, who both contributed to faunal accumulations, suggests competition for resources as well as for living spaces. Despite this, there is very little evidence of direct interaction between them to-date. Recently, a human femoral diaphysis has been recognized in South-West of Casablanca (Morocco), in the locality called Thomas Quarry I. This site is famous for its Middle Pleistocene fossil hominins considered representatives of Homo rhodesiensis. The bone was discovered in Unit 4 of the Grotte à Hominidés (GH), dated to c. 500 ky and was associated with Acheulean artefacts and a rich mammalian fauna. Anatomically, it fits well within the group of known early Middle Pleistocene Homo, but its chief point of interest is that the diaphyseal ends display numerous tooth marks showing that it had been consumed shortly after death by a large carnivore, probably a hyena. This bone represents the first evidence of consumption of human remains by carnivores in the cave. Whether predated or scavenged, this chewed femur indicates that humans were a resource for carnivores, underlining their close relationships during the Middle Pleistocene in Atlantic Morocco.     

Validation of bone surface modification models for inferring fossil hominin and carnivore feeding interactions, with reapplication to FLK 22, Olduvai Gorge, Tanzania

Resolving the issue of how Early Stone Age hominins acquired large mammal carcasses requires information on their feeding interactions with large carnivores. This ecological information and its behavioral and evolutionary implications are revealed most directly from the tooth, cut, and percussion marks on bone surfaces generated by hominin and carnivore feeding activities. This paper employs a bootstrap method, a form of random resampling with replacement, to refine published neotaphonomic models that use the assemblage-wide proportions of long bones bearing feeding traces to infer the sequences in which Plio-Pleistocene hominins and carnivores accessed flesh, marrow, and/or grease from carcasses. Results validate the sensitivity of the models for inferring hominin feeding ecology, which have been questioned on grounds shown here to be unfounded. The bootstrapped feeding trace models are applied to the late Pliocene larger mammal fossil assemblage from FLK 22 (Zinjanthropus site), Olduvai Gorge, Tanzania. High frequencies of tooth and percussion marking on long bone midshaft fragments from FLK 22 are most consistent with those feeding trace models that simulate hominin scavenging from carcasses defleshed by carnivores, while cut mark data indicate that hominins more often had access to upper forelimb flesh than upper hind limb flesh. Together, the bone surface modification data indicate that hominins typically gained secondary access to partially defleshed carnivore kills, but they also allow for the possibility of some carcasses being processed only by carnivores and only by hominins.     

Duinefontein 2: an Acheulean site in the Western Cape Province of South Africa.

Excavations at Duinefontein (DFT) 2 near Cape Town, South Africa have recovered numerous stone artefacts and animal bones on an ancient surface sealed within iron-stained eolian sands. U-series analysis of an overlying calcrete places the sands before 150 ka ago, while the large mammal taxa imply an age between 400 and 200 ka ago. The artefacts include a classic Acheulean handaxe and probable biface shaping flakes that support this age estimate. The principal mammalian species are long-horned buffalo, black wildebeest, greater kudu, Cape zebra, and grysbok/steenbok, which imply a grass-and-bush mosaic instead of the historic small-leafed shrubland.Hippopotamus and reedbuck indicate that water stood nearby, probably in dune swales. The large mammal bones are mostly vertebrae and other axial elements, often in near-anatomical order. Both proximal and distal appendicular elements are rare. Bones with carnivore damage are common, but ones with stone tool marks are scarce. The sum suggests a water-edge attritional death site where people played a minimal role and carcasses were disarticulated mainly by carnivore feeding and by trampling. Stone tool marks tend to be equally rare at other Acheulean attritional death sites, and the implication may be that Acheulean people rarely obtained large mammals, whether by hunting or scavenging. Human scavengers at DFT2 would not have encountered a disproportionate number of distal (versus proximal) limb elements, and it follows that the tendency for distal elements to dominate many archeological assemblages need not reflect scavenging versus hunting. Even if DFT2 was not itself a locus of intense human activity, it provides a useful baseline for evaluating bone damage, skeletal part representation, and other variables at sites where people were deeply involved.     

Taphonomic analysis of the Middle Stone Age faunal assemblage from Pinnacle Point Cave 13B, Western Cape, South Africa

A detailed taphonomic analysis is provided for the mammalian and tortoise faunal assemblages from Pinnacle Point Cave 13B (PP13B). It is the first of several reports on the fauna from this site, and must necessarily precede analyses focused on higher level interpretations of Middle Stone Age (MSA) butchery, transport, and hunting behavior. The taphonomic work shows that the faunal assemblage is well preserved and there are discernable differences in the taphonomic pathways to which the fauna was subjected at PP13B between the Middle and Late Pleistocene, between the front and back of the cave, and between body size classes. The largest mammals (size classes 2-5, body weight >24?kg) were mainly accumulated by MSA hominins. Size class 1 ungulates also exhibit a degree of hominin modification consistent with some hominin accumulation of fresh carcasses, but this is more variable through time and includes an observable degree of independent carnivore contribution. Basic taxonomic comparisons reveal a low representation of small mammals, tortoises, and marine mammals at PP13B relative to larger (>4.5?kg) terrestrial mammals. This is a different pattern from other MSA sites along the southwestern coast of South Africa, where small mammals and tortoises are abundant. A microscopic study of the bone surfaces confirms that MSA hominins exploited these small faunal components opportunistically, while focusing most heavily on large terrestrial ungulates. All faunal components show evidence of carnivore scavenging of hominin food debris and a high degree of density mediated destruction. Raptors are at no point implicated as major accumulators of any fauna. The study demonstrates that the full spectrum of MSA faunal exploitation can only be understood when the large mammal, small mammal, and tortoise components of fossil assemblages have all been subjected to comprehensive taphonomic analyses.     

Sr/Ca and early hominin diets revisited: new data from modern and fossil tooth enamel

A previous study of strontium/calcium (Sr/Ca) ratios in Paranthropus suggested that it consumed more animal foods than was previously believed. However, that study looked at Sr/Ca in fossil bone, which is known to be highly susceptible to diagenesis. Enamel, in contrast, is resistant to post-mortem alteration making it a more appropriate material for Sr/Ca analysis of Plio-Pleistocene fossils. Yet, we know virtually nothing about Sr/Ca in the enamel of modern African mammals, much less fossil taxa. To address this gap, we studied Sr/Ca in tooth enamel from modern mammals in the greater Kruger National Park, South Africa, as well as fossil fauna from the Sterkfontein Valley. Grazing herbivores have the highest Sr/Ca, followed by browsers and carnivores in both modern and fossil fauna. This similarity in ecological Sr/Ca patterning between modern and fossil fauna shows that diagenesis has not obscured the primary dietary signals. Australopithecus has significantly higher Sr/Ca than Paranthropus, and higher Sr/Ca than fossil papionins, browsers, and carnivores. Paranthropus has lower Sr/Ca than grazers, but its Sr/Ca is higher or equal to that of fossil papionins, browsers, and carnivores. Thus, Sr/Ca for both hominins is relatively high, and provides no direct evidence for omnivory in either taxon. The consumption of underground resources or insects are among the possible explanations for the highly elevated Sr/Ca in Australopithecus.     

藏北尼阿木底遗址发现的似阿舍利石器——兼论晚更新世人类向青藏高原的扩张

2013年夏,在藏北那曲地区申扎县羌塘高原旧石器调查中,从规模恢弘的尼阿木底遗址地表暴露的、数以万计的石制品中,分别采集到似阿舍利类型"手斧"9件和"薄刃斧"2件。本文对尼阿木底遗址所见的这类器物进行了细致的观察与分析,认为它们只是旧石器时代晚期勒瓦娄哇技术产品的石核,或偶尔为之的、与手斧和薄刃斧形似的石制品,并非真正意义上的手斧与薄刃斧。尼阿木底遗址不存在从选料、剥片、加工出成品,到使用、损坏和废弃等各个阶段的、明确的阿舍利类型石器工业产品生产体系的工艺链条,换言之,在尼阿木底生活的古人脑海中,并不存在一个加工阿舍利类型手斧和薄刃斧等工具的"概念模板"(Mental template)。联系到青藏高原其他地区以前所报道的同类材料,我们认为在青藏高原腹地目前所见的旧石器遗址中,还没有真正意义上的阿舍利石器工业类型的遗存。晚更新世末期时,来自于印巴次大陆方向的早期占领者,沿着喜马拉雅山脉、冈底斯山脉和昆仑山脉三条东西走向的巨大山系之间的通道,自高原西南方向开始向高原腹地扩张,其间阿舍利石器工业技术在非洲和欧亚大陆早已消弭,当时的青藏高原腹地并不具备重新产生该类石器工业技术的土壤。     

The Pleistocene fauna (other than Primates) from Asbole, lower Awash Valley, Ethiopia, and its environmental and biochronological implications

The Asbole area in the Lower Awash Valley yielded a diverse fauna of large and small mammals, associated with an Acheulean industry. The most notable forms are a potentially new species of herpestid, a large collection of Kolpochoerus majus, and the earliest known Bos in Africa. Biochronologically, this fauna belongs to the earliest Middle Pleistocene, and is roughly contemporaneous with the Bodo site further south. Paleoenvironmentally, the fauna suggests a mosaic of landscapes among which humid environments, grasslands and forests, are predominant.     

周口店第一地点西剖面2009-2010年发掘报告

周口店第一地点再次发掘的主要目的在于提升遗址西剖面的保护状态与水平,同时也为东亚直立人演化方面某些焦点问题的研究与解决提供机遇与可能。2009-2010年,在西剖面最顶部堆积(即第三层)的野外发掘中共计出土大中型动物化石材料约800件。埋藏学研究表明,西剖面第三层中葛氏斑鹿(Cervus(Sika)grayi)等动物属种的骨骼富集并非古人类生存行为的结果;石制品以及古人类化石材料的缺失也在一定程度上印证了这一结论。结合发掘过程中揭露出来的一系列洞穴坍塌事件的证据,我们初步认为,至少在第三层堆积形成时期,第一地点西剖面位置处的洞穴形态并不适合于古人类的栖居活动;葛氏斑鹿等骨骼材料则是当时在遗址附近活动的各种动物因意外因素陷落于其中而得以聚集的。     

The Vallonnet cave on the northern Mediterranean border: A record of one of the oldest human presences in Europe

Prehistoric sites testifying to human presence older than one million years in Europe are rare, and in the current state of knowledge, the oldest of them have been dated to around 1.4–1.5 Ma. The Vallonnet cave at Roquebrune-Cap-Martin in the Alpes-Maritimes, on the Mediterranean border, is one of the oldest sites in France to have yielded evidence of human activity: a lithic assemblage of about a hundred pieces and traces of butchery on bones of an Epivillafranchian fauna. The archaeological levels of this small cave were recently dated between 1.1 and 1.2 Ma by U-Pb correlated with paleomagnetic data. The site was occupied alternately by large carnivores that used it as a den or a lair, and by hominins that stayed there briefly in bivouac. The lithic remains are mainly percussion tools, shaped pebbles, flakes and cores, whose raw materials are local, or even semi-local, and on the whole not very diversified with mainly limestone, and to a lesser extent sandstone, quartzite, flint and quartz. This assemblage is attributed to a Mode 1 technology (Oldowayen), among which macro-tools (hammerstones, shaped and fractured pebbles) are found alongside rarer elements resulting from debitage operating chains aimed at producing sharp-edged flakes, very rarely retouched. The bipolar-on-anvil flaking technique could be identified from the characteristics of some artifacts. Several refitting flakes on shaped pebbles or percussion tools attest to knapping and percussion activities in the cave. Hominins consumed the remains of large herbivore carcasses, as attested by the presence of cutting and fracturing marks on some bones. The presence of a freshwater source in the immediate vicinity, and the knapping and butchering activities here therefore document the subsistence behavior of Lower Pleistocene human groups, certainly in competition with the carnivores present.     

河南栾川发现直立人化石及其演化意义

洛阳考古研究院2012年5月至10月在栾川孙家洞作抢救性发掘,于原生层位发现古人类化石,同时伴有大量哺乳动物化石,以及少量人工石制品。根据伴生动物群的初步鉴定,栾川古人类的生存时代为中更新世。6件古人类化石代表3个个体:1个成年人和2个未成年人,未成年人的牙齿生长发育阶段分别与6～7岁和11～12岁的现代青少年相当。栾川孙家洞古人类牙齿具有一定的原始性,明显区别于现代人,其形态大小位于周口店直立人牙齿的变异范围内,可归入直立人的形态范畴;同时具有东亚古人类及现代蒙古人群的地区性特征。基于幼年个体上下颌骨的牙齿发育,推测栾川古人类第一臼齿萌出年龄可能接近6岁,提示具有接近于现代人的牙齿生长模式和生活史特点。栾川中更新世古人类化石的发现为中国古人类连续演化假说提供了新证据。     

Hominin occupations at the Dmanisi site, Georgia, Southern Caucasus: raw materials and technical behaviours of Europe's first hominins

Dmanisi is the oldest site outside of Africa that records unquestioned hominin occupations as well as the dispersal of hominins in Europe and Asia. The site has yielded large numbers of artefacts from several periods of hominin occupation. This analysis of Dmanisi stone tool technology includes a review of all the pieces recovered during the last 15 years of excavations. This lithic assemblage gives insights into the hominin behaviour at 1.7-1.8 Ma in Eurasia. Dmanisi hominins exploited local rocks derived from either nearby riverbeds or outcrops, and petrographic study provides data on patterns of stone procurement. Recent geological surveys and technological studies of the artefacts illustrate the roles of hominins in composing the assemblage. Dmanisi hominins selected two types of blanks, including cobbles and angular blocks, of basalt, andesite, and tuffs. Many complete cobbles, pebbles, and rolled blocks in basalt were unmodified, and geological analyses and surveys indicate that hominins brought manuports back to the site, suggesting a complex procurement strategy. Cores, flakes and debris show that all stages of flaking activity took place at the site. Numerous unifacial cores suggest that knapping was not very elaborate. Centripetal knapping is observed on some flake-cores. Knapping was influenced by the blank shape and natural angles. Most flaked objects were either cores or chopper-cores. Flakes predominate while flake tools are rare. The Dmanisi lithic assemblage is comparable to Oldowan sites in Africa in terms of reduction sequence, organisation of the removals, platform types, and the lack of retouched flakes. Dmanisi artefacts and may have been produced by the original hominins in Europe and Asia.     

Understanding Ancient Hominin Dispersals Using Artefactual Data: A Phylogeographic Analysis of Acheulean Handaxes

Background

Reconstructing the dispersal patterns of extinct hominins remains a challenging but essential goal. One means of supplementing fossil evidence is to utilize archaeological evidence in the form of stone tools. Based on broad dating patterns, it has long been thought that the appearance of Acheulean handaxe technologies outside of Africa was the result of hominin dispersals, yet independent tests of this hypothesis remain rare. Cultural transmission theory leads to a prediction of a strong African versus non-African phylogeographic pattern in handaxe datasets, if the African Acheulean hypothesis is to be supported.

Methodology/Principal Findings

Here, this prediction is tested using an intercontinental dataset of Acheulean handaxes and a biological phylogenetic method (maximum parsimony). The analyses produce a tree consistent with the phylogeographic prediction. Moreover, a bootstrap analysis provides evidence that this pattern is robust, and the maximum parsimony tree is also shown to be statistically different from a tree constrained by stone raw materials.

Conclusions/Significance

These results demonstrate that nested analyses of behavioural data, utilizing methods drawn from biology, have the potential to shed light on ancient hominin dispersals. This is an encouraging prospect for human palaeobiology since sample sizes for lithic artefacts are many orders of magnitude higher than those of fossil data. These analyses also suggest that the sustained occurrence of Acheulean handaxe technologies in regions such as Europe and the Indian subcontinent resulted from dispersals by African hominin populations.     

Taphonomy of the fossil hominid bones from the Acheulean site of Castel di Guido near Rome, Italy

Castel di Guido near Rome is one of the few open air Middle Pleistocene European sites that has yielded hominid skeletal remains associated with fossil fauna and Acheulean implements. The fossil hominid bones include two femoral shafts, respectively designated Castel di Guido-1 (CdG-1) and CdG-2, an occipital fragment (CdG-3), a right maxilla lacking teeth (CdG-4), a portion of right parietal (CdG-5), a right temporal (CdG-6), and a fragment of left parietal vault (CdG-7). CdG-1 through CdG-4 were collected in 1979-1982 on the surface, together with fossil fauna, where ploughing incised fossiliferous tuffaceous sands. Excavations conducted in the same area from 1980 to 1990 led to the discovery of CdG-5, CdG-6 and CdG-7 within the tuffaceous sands, which were shown to overlay a bone-bearing paleosurface, with abundant evidence of hominid activities. The Castel di Guido hominid assemblage poses intriguing taphonomic questions. The analysis of the physical evidence offered by the bone surfaces, reported in the present study, indicates that the hominid skeletal remains were heavily fragmented before fossilization and exposed to carnivores and rodents, as well as to trampling and/or friction in abrasive sediment. Although definitive conclusions cannot be reached on the basis of the available evidence, it is possible that clusters of incisions localized on specific regions of the Castel di Guido fossil hominid bones might reflect deliberate human manipulations.     

Landscapes and their relation to hominin habitats: case studies from Australopithecus sites in eastern and southern Africa

We examine the links between geomorphological processes, specific landscape features, surface water drainage, and the creation of suitable habitats for hominins. The existence of mosaic (i.e., heterogeneous) habitats within hominin site landscape reconstructions is typically explained using models of the riverine and gallery forest settings, or the pan or lake setting. We propose a different model: the Tectonic Landscape Model (TLM), where tectonic faulting and volcanism disrupts existing pan or river settings at small-scales (∼10-25 km). Our model encompasses the interpretation of the landscape features, the role of tectonics in creating these landscapes, and the implications for hominins. In particular, the model explains the underlying mechanism for the creation and maintenance of heterogeneous habitats in regions of active tectonics. We illustrate how areas with faulting and disturbed drainage patterns would have been attractive habitats for hominins, such as Australopithecus, and other fauna. Wetland areas are an important characteristic of surface water disturbance by fault activity; therefore we examine the tectonically-controlled Okavango Delta (Botswana) and the Nylsvley wetland (South Africa) as modern examples of how tectonics in a riverine setting significantly enhance the faunal and floral biodiversity. While tectonic landscapes may not have been the only type of attractive habitats to hominins, we propose a suite of landscape, faunal, and floral indicators, which when recovered together suggest that site environments may have been influenced by tectonic and/or volcanic activity while hominins were present. For the fossil sites, we interpret the faulting and landscapes around australopithecine-bearing sites of the Middle Awash (Ethiopia) and Makapansgat, Taung, and Sterkfontein (South Africa) to illustrate these relationships between landscape features and surface water bodies. Exploitation of tectonically active landscapes may explain why the paleoenvironmental signals, anatomy, diets, as well as the fauna associated with Australopithecus appear largely heterogeneous through time and space. This hypothesis is discussed in light of potential preservation and time-averaging effects which may affect patterns visible in the fossil record. The model, however, offers insight into the landscape processes of how such habitats are formed. The landscape features and range of habitat conditions, specifically the wetter, down-dropped plains and drier, uplifted flanks persist in close proximity for as long as the fault motion continues. The Tectonic Landscape Model provides an alternative explanation of why mixed habitats may be represented at certain sites over longer timescales.     

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.     

Retourner à Moulin Quignon

In March 1863, Jacques Boucher de Perthes found in the alluvia of Moulin Quignon (Abbeville, France) a human jaw in association with lithic artifacts and fossil bones of mammals. Quickly challenged, this discovery will appear as a fraud whose prehistorian was the victim. Since that time, the collection and the Moulin Quignon site are synonymous with an archaeological forgery. Forgotten for almost 150 years, these collections became in 2012 a research topic in the context of a program devoted to oldest settlements and Acheulean. The objective was to identify and study the whole 1863–1864 documentation set (human remains, artifacts, sediments, fauna bones, malacological remains, archives, papers, site) to allow further analysis of the whole subject.     

A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania

Neotaphonomic studies have determined the patterns of bone damage created by larger mammalian carnivores when consuming mammalian carcasses. Typically, mammalian carnivores gnaw and break bones to various degrees in order to access marrow, grease, and brain tissue. In contrast, crocodiles attempt to swallow whole parts of mammal carcasses, inflicting in the process tooth marks and other feeding traces on some of the bones they are unable to ingest. Although crocodiles are major predators of larger mammals along the margins of protected tropical rivers and lakes, their feeding traces on bone have received little systematic attention in neotaphonomic research. We present diagnostic characteristics of Crocodylus niloticus damage to uningested mammal bones resulting from a series of controlled observations of captive crocodile feeding. The resulting bone assemblages are composed of primarily complete elements from articulating units, some of which bear an extremely high density of shallow to deep, transversely to obliquely oriented tooth scores over often large areas of the bone, along with shallow to deep pits and punctures. Some of the tooth marks (bisected pits and punctures, hook scores) have a distinctive morphology we have not observed to be produced by mammalian carnivores. The assemblages are also characterized by the retention of both low- and high-density bone portions, an absence of gross gnawing, and minimal fragmentation. Together, the damage characteristics associated with feeding by crocodiles are highly distinctive from those produced by mammalian carnivores. Modern surface bone assemblages along the Grumeti River in Tanzania's Serengeti National Park contain a mixture of specimens bearing damage characteristic of crocodiles and mammalian carnivores. Comparison of Plio-Pleistocene fossil bones from Olduvai Gorge, Tanzania, to bones damaged by captive and free-ranging Nile crocodiles reveals direct evidence of fossil crocodilian feeding from larger mammal bones associated with Oldowan stone artifacts.     

Brain expansion in early hominins predicts carnivore extinctions in East Africa

While the anthropogenic impact on ecosystems today is evident, it remains unclear if the detrimental effect of hominins on co‐occurring biodiversity is a recent phenomenon or has also been the pattern for earlier hominin species. We test this using the East African carnivore fossil record. We analyse the diversity of carnivores over the last four million years and investigate whether any decline is related to an increase in hominin cognitive capacity, vegetation changes or climatic changes. We find that extinction rates in large carnivores correlate with increased hominin brain size and with vegetation changes, but not with precipitation or temperature changes. While temporal analyses cannot distinguish between the effects of vegetation changes and hominins, we show through spatial analyses of contemporary carnivores in Africa that only hominin causation is plausible. Our results suggest that substantial anthropogenic influence on biodiversity started millions of years earlier than currently assumed.     

Temporal Change in Functional Richness and Evenness in the Eastern African Plio-Pleistocene Carnivoran Guild

We analyze functional richness and functional evenness of the carnivoran guild in eastern Africa from 3.5 Ma to 1.5 Ma, and compare them to the present day. The data consist of characters of the craniodental apparatus of 76 species of fossil and extant carnivorans, divided into four 0.5 Ma time slices from 3.5 to 1.5 Ma, together with the modern fauna. Focus is on large (>21.5 kg) carnivores. Results show that the large carnivore guild has lost nearly 99% of its functional richness since 3.5 Ma, in a process starting prior to 2 Ma. Measurement of functional evenness shows the modern large carnivore guild to be unique in being randomly distributed in morphospace while in all past time slices there is significant clustering of species. The results are analyzed in the light of known changes to climate and environment in eastern Africa. We conclude that climate change is unlikely to explain all of the changes found and suggest that the evolution of early hominins into carnivore niche space, especially the evolution of derived dietary strategies after 2 Ma, played a significant part in the reduction of carnivore functional richness.