A human mandible (BH-1) from the Pleistocene deposits of Mala Balanica cave (Sićevo Gorge, Niš, Serbia)

Neandertals and their immediate predecessors are commonly considered to be the only humans inhabiting Europe in the Middle and early Late Pleistocene. Most Middle Pleistocene western European specimens show evidence of a developing Neandertal morphology, supporting the notion that these traits evolved at the extreme West of the continent due, at least partially, to the isolation produced by glacial events. The recent discovery of a mandible, BH-1, from Mala Balanica (Serbia), with primitive character states comparable with Early Pleistocene mandibular specimens, is associated with a minimum radiometric date of 113 + 72 − 43 ka. Given the fragmented nature of the hemi-mandible and the fact that primitive character states preclude assignment to a species, the taxonomic status of the specimen is best described as an archaic Homo sp. The combination of primitive traits and a possible Late Pleistocene date suggests that a more primitive morphology, one that does not show Neandertal traits, could have persisted in the region. Different hominin morphologies could have survived and coexisted in the Balkans, the “hotspot of biodiversity.” This first hominin specimen to come from a secure stratigraphic context in the Central Balkans indicates a potentially important role for the region in understanding human evolution in Europe that will only be resolved with more concentrated research efforts in the area.     

The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW,NW Germany)

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100 m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave.     

A comparative study of frontal bone morphology among Pleistocene hominin fossil groups

Features of the frontal bone that are conventionally used to distinguish among fossil hominin groups were quantitatively examined. Fifty-five fossil crania dating from the early to the late Pleistocene were analyzed. Using a modified pantograph, outlines of the frontal bone were collected along the midsagittal and two parasagittal planes. The profile from nasion to bregma, as well as two profiles above the medial and lateral sections of the orbit, respectively, extending from the orbital margin to the coronal suture were traced. The outlines were measured using Elliptical Fourier Function Analysis (EFFA), which enabled a quantification of aspects of the frontal bone that have historically been described primarily in nonmetric or linear terms. Four measurements were obtained: 1) overall morphology as expressed in the Fourier harmonic amplitudes; 2) maximum projection of the supraorbital torus at three points along the browridge (glabella and the medial and lateral aspects of the torus above the orbit); 3) maximum distance of the frontal squama from the frontal chord, capturing forehead curvature; and 4) nasion-bregma chord length. The results indicate that the midsagittal profile is significantly different among all Pleistocene groups in analyses that include both size and shape, as well as size-adjusted data. Homo erectus is significantly different from the late Pleistocene groups (Neandertals and early modern H. sapiens) in glabellar projection. Anatomically modern humans are significantly different from all other groups in both raw and size-standardized analyses of all three outlines that captured overall morphology, as well as forehead curvature and lateral supraorbital torus prominence, and middle Pleistocene Homo are significantly different in both medial and lateral overall parasagittal form. However, for the majority of analyses there were no significant differences among the Pleistocene archaic groups in supraorbital torus projection, frontal squama curvature, nasion-bregma chord length, or overall frontal bone morphology.     

New Radiometric Ages for the BH-1 Hominin from Balanica (Serbia): Implications for Understanding the Role of the Balkans in Middle Pleistocene Human Evolution

Newly obtained ages, based on electron spin resonance combined with uranium series isotopic analysis, and infrared/post-infrared luminescence dating, provide a minimum age that lies between 397 and 525 ka for the hominin mandible BH-1 from Mala Balanica cave, Serbia. This confirms it as the easternmost hominin specimen in Europe dated to the Middle Pleistocene. Inferences drawn from the morphology of the mandible BH-1 place it outside currently observed variation of European Homo heidelbergensis. The lack of derived Neandertal traits in BH-1 and its contemporary specimens in Southeast Europe, such as Kocabaş, Vasogliano and Ceprano, coupled with Middle Pleistocene synapomorphies, suggests different evolutionary forces acting in the east of the continent where isolation did not play such an important role during glaciations.     

Fossil human remains from Bolomor Cave (Valencia, Spain)

Systematic excavations carried out since 1989 at Bolomor Cave have led to the recovery of four Pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. All of these specimens date to the late Middle and early Late Pleistocene (MIS 7-5e). The fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-Homo sapiens) members of the genus Homo. Among the dental remains, the lack of a midtrigonid crest in the M₁ represents a departure from the morphology reported for the majority of Neandertal specimens, while the large dimensions and pronounced shoveling of the marginal ridges in the C¹ are similar to other European Middle and late Pleistocene fossils. The parietal bone is very thick, with dimensions that generally fall above Neandertal fossils and resemble more closely the Middle Pleistocene Atapuerca (SH) adult specimens. Based on the presence of archaic features, all the fossils from Bolomor are attributed to the Neandertal evolutionary lineage.     

A critical review of the German Paleolithic hominin record

We review the hominin fossil record from western Central Europe in light of the recent major revisions of the geochronological context. The mandible from Mauer (Homo heidelbergensis), dated to circa 500,000 years ago, continues to represent the earliest German hominin and may coincide with the occupation of Europe north of the high alpine mountain chains. Only limited new evidence is available for the Middle Pleistocene, mostly in the form of skull fragments, a pattern that may relate to taphonomic processes. These finds and their ages suggest the gradual evolution of a suite of Neandertal features during this period. Despite new finds of classic Neandertals, there is no clear proof for Neandertal burial from Germany. Alternatively, cut marks on a skull fragment from the Neandertal type site suggest special treatment of that individual. New Accelerator Mass Spectrometry (AMS) radiocarbon dates of previous finds leave little reliably dated evidence for anatomically modern humans (AMH) in Europe before 30,000 BP; the remains from Hahn?fersand, Binshof-Speyer, Paderborn-Sande, and Vogelherd are now of Holocene age. Thus, a correlation of AMH with the Aurignacian remains to be proven, and the general idea of a long coexistence of Neandertals and AMH in Europe may be questioned. In western Central Europe, evidence of Gravettian human fossils is also very limited, although a new double grave from lower Austria may be relevant. The only dated burial from the German Upper Paleolithic (from Mittlere Klause) falls into a time period (circa 18,600 BP) represented by only a few occupation sites in western Central Europe. A number of human remains at Magdalenian sites appear to result from variable (secondary) burial practices. In contrast, the Final Paleolithic (circa 12,000-9600 cal. BC) yields an increase of hominin finds, including multiple burials (Bonn-Oberkassel, Neuwied-Irlich), similar to the situation in western and southern Europe.     

Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave, Atapuerca, Spain

The Gran Dolina cave site is famous for having delivered some of the oldest hominin remains of Western Europe (Homo antecessor, ca. 960 ka). Moreover, the evidence of lithic industries throughout the long vertical section suggests occupation on the part of hominins from the latest early Pleistocene (levels TD3/4, TD5, and TD6) to the late middle Pleistocene (level TD10). The Gran Dolina Sondeo Sur (TDS) has furnished a great number of small-vertebrate remains; among them some 40,000 bones are attributed to amphibians and squamates. Although they do not differ specifically from the extant herpetofauna of the Iberian Peninsula, the overlap of their current distribution areas (= mutual climatic range method) in Spain can provide mean annual temperatures (MAT), the mean temperatures of the coldest (MTC) and warmest (MTW) months, and mean annual precipitation (MAP) estimations for each sub-level, and their change can be studied throughout the sequence. Results from the squamate and amphibian study indicate that during hominin occupation the MAT (10-13 °C) was always slightly warmer than at present in the vicinity of the Gran Dolina Cave, and the MAP (800-1000 mm) was greater than today in the Burgos area. Climatic differences between “glacial” and “interglacial” phases are poorly marked. Summer temperatures (MTW) show stronger oscillations than winter temperatures (MTC), but seasonality remains almost unchanged throughout the sequence. These results are compared with those for large mammals, small mammals, and pollen analysis, giving a scenario for the palaeoclimatic conditions that occurred during the early to middle Pleistocene in Atapuerca, and hence a scenario for the hominins that once lived in the Sierra de Atapuerca.     

A fossil <Emphasis Type="Italic">Potadoma</Emphasis> (Gastropoda: Pachychilidae) from Pleistocene central Kalahari fluvio-lacustrine sediments

A shell of the gastropod Potadoma was discovered in fluvio-lacustrine sediments of the Kalahari (Botswana). A late Pleistocene age of the sediments of ca. 46,000 cal. years BP was established by radiocarbon dating. It is the first record of the freshwater genus Potadoma from southern Africa. Modern counterparts as well as other Pleistocene species of Potadoma had been reported elsewhere from (palaeo-) habitats located at least 2,000 km further in the north, e.g., in the Congo Basin. So far it can be only speculated about possible causes for such a disjunct distribution. The discovery of Potadoma and five more fossil gastropods from the same sediments was used to outline palaeolimnological features. Considering the geomorphological setting, the assemblage of fossil gastropods indicates that the nowadays mainly dry Boteti River was permanently flowing through the western Makgadikgadi Pans ca. 46,000 cal. years BP. The existence of such a riverine palaeoenvironment in the pans demonstrates that the so-called Lake Palaeo-Makgadikgadi was comparatively small during that time, not spanning the area of investigation.     

The medial pterygoid tubercle in the Atapuerca Early and Middle Pleistocene mandibles: Evolutionary implications

Numerous studies have attempted to identify the presence of uniquely derived (autoapomorphic) Neandertal features. Here, we deal with the medial pterygoid tubercle (MTP), which is usually present on the internal face of the ascending ramus of Neandertal specimens. Our study stems from the identification of a hypertrophied tubercle in ATD6‐96, an Early Pleistocene mandible recovered from the TD6 level of the Atapuerca‐Gran Dolina site and attributed to Homo antecessor. Our review of the literature and study of numerous original fossil specimens and high quality replicas confirm that the MTP occurs at a high frequency in Neandertals (ca. 89%) and is also present in over half (ca. 55%) of the Middle Pleistocene Sima de los Huesos (SH) hominins. In contrast, it is generally absent or minimally developed in other extinct hominins, but can be found in variable frequencies (<ca. 25%) in Pleistocene and recent H. sapiens samples. The presence of this feature in ATD6‐96 joins other traits shared by H. antecessor, the SH hominins and Neandertals. Since the TD6 hominins have been attributed either to MIS 21 or to MIS 25, it seems that a suite of assumed derived Neandertal features appeared in the Early Pleistocene, and they should be interpreted as synapomorphies shared among different taxa. We suggest that H. antecessor, the SH hominins and Neandertals shared a common ancestor in which these features appeared during the Early Pleistocene. The presence of the MTP in taxa other than H. neanderthalensis precludes this feature from being a Neandertal autapomorphy. Am J Phys Anthropol 156:102–109, 2015 © 2014 Wiley Periodicals, Inc.     

安徽东至华龙洞遗址洞穴演化与古人类活动

安徽东至华龙洞因发现距今约30万年的古人类头骨化石和大量动物化石及石制品受到学术界广泛关注。本文对华龙洞遗址的地质、地貌、沉积物特点及洞穴演化过程与古人类活动的关系进行分析。华龙洞地处扬子陆块区西北缘,周边呈现低山—丘陵—湖泊平原地貌景观。与华龙洞遗址密切关联的岩溶洞穴,发育在上寒武统微晶灰岩和白云质灰岩岩系内,中更新以来的地壳运动和岩溶发育是其形成的主要营力。华龙洞遗址是一处坍塌的洞穴,其发育大致经历发育初期（中更新世早期甚至更早）—稳定发育期（中更新世中期）—坍塌埋藏期（中更新世中晚期）等三个阶段,岩溶发育和洞外溪谷的侵蚀使得原始洞穴和堆积物一起在重力作用下坍塌。洞穴坍塌沉积物主要包括围岩岩块与碎屑、各种岩溶沉积物和文化遗物,胶结坚硬,不规则地埋藏于裂隙和巨石之间。古人类在遗址的活动时间处在距今约30万年前的稳定发育期,石制品和骨骼表面痕迹证据表明,华龙洞古人类具备依据不同原料的特点采取砸击法与锤击法并用的技术策略;石片边缘的使用痕迹和动物骨骼表面痕迹显示,古人类在遗址可能进行过肢解动物的行为。本研究对揭示长江下游中更新世中晚期古人类演化与适应生存行为具有重要意义。     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

A fossil hominid frontal from Velika Pećina (Croatia) and a consideration of Upper Pleistocene hominids from Yugoslavia

Fossil hominid remains dating to the Upper Pleistocene in Yugoslavia are reviewed. Particular emphasis is placed on the assessment of a hominid frontal from the site of Velika Pe?ina in northwestern Croatia. This specimen represents the earliest absolutely-dated hominid associated with the Upper Paleolithic in Europe. Also the hominid material from the site of Veternica is discussed, and data are presented on the new remains from ?andalja. It is concluded that no Neandertal remains, except for Krapina, have been found in Yugoslavia.     

Form and seismic stratigraphy of Halimeda banks in part of the northern Great Barrier Reef Province

Twenty-six percent of the total shelf area in the Northern Region of the Great Barrier Reef Province between latitudes 14°27 and 15°02S is occupied by algal (Halimeda) lithofacies. Sea-bed relief over this bankforming algal lithofacies, which dominates the outer shelf, is generally complex and variable over relatively short distances, but lateral continuity of morphological features near Petricola and Stewart shoals has been demonstrated by parallel profiling. The most prominent subbttom reflector is a pre-Holocene erosion surface, regarded as the Holocene/Pleistocene unconformity. In this area, the maximum thickness of Holocene Halimeda banks above the main subbottom reflector is 19 m. Seismic response suggests three main sequences in the Halimeda bank depostis, which probably relate to changes in environmental conditions, and the recognition of similar seismic characteristics in the deposits beneath the Holocene/Pleistocene unconformity indicates that Halimeda banks may have been a feature of the outer shelf of the Great Barrier Reef in Plesitocene times.     

Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik‐Tash child

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.     

Neandertal versus Modern Human Dietary Responses to Climatic Fluctuations

The Neandertal lineage developed successfully throughout western Eurasia and effectively survived the harsh and severely changing environments of the alternating glacial/interglacial cycles from the middle of the Pleistocene until Marine Isotope Stage 3. Yet, towards the end of this stage, at the time of deteriorating climatic conditions that eventually led to the Last Glacial Maximum, and soon after modern humans entered western Eurasia, the Neandertals disappeared. Western Eurasia was by then exclusively occupied by modern humans. We use occlusal molar microwear texture analysis to examine aspects of diet in western Eurasian Paleolithic hominins in relation to fluctuations in food supplies that resulted from the oscillating climatic conditions of the Pleistocene. There is demonstrable evidence for differences in behavior that distinguish Upper Paleolithic humans from members of the Neandertal lineage. Specifically, whereas the Neandertals altered their diets in response to changing paleoecological conditions, the diets of Upper Paleolithic humans seem to have been less affected by slight changes in vegetation/climatic conditions but were linked to changes in their technological complexes. The results of this study also indicate differences in resource exploitation strategies between these two hominin groups. We argue that these differences in subsistence strategies, if they had already been established at the time of the first contact between these two hominin taxa, may have given modern humans an advantage over the Neandertals, and may have contributed to the persistence of our species despite habitat-related changes in food availabilities associated with climate fluctuations.     

Search for possible latitudinal ecotypes inDumontia contorta (Rhodophyta)

Effects of daylength and temperature on the formation of erect fronds (macrothalli) from crusts (microthalli) ofDumontia contorta (S. G. Gmel.) Rupr. from three localities in Nova Scotia and one locality in Southern Iceland were investigated and compared to such effects shown by strains from three different East Atlantic localities (Isle of Man; Zeeland, S. W. Netherlands; and Roscoff, Brittany, France). Although these strains showed small differences in their temperature-daylength responses, these could not be interpreted as latitudinal adaptations, and consequently no latitudinal ecotypes could be found forDumontia contorta in the N. Atlantic Ocean. Upright fronds are formed at a broad temperature range of about 4°–18°C and at daylengths 13 h. Only in the southernmost part of its distribution area can high autumnal temperatures be expected to block the reappearance of upright fronds after passage of the critical daylength in September. In the larger part of the distribution area even summer temperatures are not high enough to block formation of uprights and here apparently only short daylengths initiate the reappearance of young upright fronds in autumn. The consequences of these aspects of the life history regulation for the geographic distribution are discussed.Paper presented at the Seaweed Biogeography Workshop of the International Working Group on Seaweed Biogeography, held from 3–7 April, 1984 at the Department of Marine Biology, University of Groningen, The Netherlands. Convenor: C. van den Hoek.     

The past and present occurrence of <Emphasis Type="Italic">Elatine</Emphasis> and implications for palaeoenvironmental reconstructions

The past and present occurrence of Elatine hydropiper L. (eight-stamened waterwort), E. hexandra (Lapierre) DC. (six-stamened waterwort) and E. triandra Schkuhr (three-stamened waterwort) in the Netherlands is discussed. It has proved possible to distinguish the slightly curved seeds of E. hexandra and E. triandra in subfossil material on morphological grounds. E. hexandra is the most common species at present, but subfossil finds are confined to Late-glacial and Pre-boreal sediments of one lake in the Pleistocene area of the Netherlands. Living plants of E. triandra have only been found in the Netherlands in 1838–1839, but there are 17 records from five archaeological sites, all located in the western Dutch estuarine area. Several of these sites also yielded E. hydropiper, archaeobotanically the most common species. The occurrence of E. triandra and E. hydropiper in the Netherlands seems to have been favoured by high summer temperatures. The ecological amplitude of this combination of species gives firm clues for the reconstruction of the environment, which must have been a freshwater tidal area. Since this type of environment is strongly threatened on a worldwide scale, the presence of these species in the past may also provide interesting information for present nature development projects in the Dutch estuarine area.