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Bibliography on: Homo floresiensis, The Hobbit

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ESP: PubMed Auto Bibliography 20 Jul 2024 at 01:55 Created: 

Homo floresiensis, The Hobbit

Wikipedia: Homo floresiensis ("Flores Man"; nicknamed "hobbit" for its small stature) is an extinct species in the genus Homo. The remains of an individual that would have stood about 3.5 feet (1.1 m) in height were discovered in 2003 at Liang Bua on the island of Flores in Indonesia. Partial skeletons of nine individuals have been recovered, including one complete skull, referred to as "LB1".These remains have been the subject of intense research to determine whether they represent a species distinct from modern humans. This hominin had originally been considered to be remarkable for its survival until relatively recent times, only 12,000 years ago. However, more extensive stratigraphic and chronological work has pushed the dating of the most recent evidence of their existence back to 50,000 years ago. Their skeletal material is now dated to from 100,000 to 60,000 years ago; stone tools recovered alongside the skeletal remains were from archaeological horizons ranging from 190,000 to 50,000 years ago. Fossil teeth and a partial jaw from hominins believed ancestral to H. floresiensis were discovered in 2014 and described in 2016. These remains are from a site on Flores called Mata Menge, about 74 km from Liang Bua. They date to about 700,000 years ago and are even smaller than the later fossils. The form of the fossils has been interpreted as suggesting that they are derived from a population of H. erectus that arrived on Flores about a million years ago (as indicated by the oldest artifacts excavated on the island) and rapidly became dwarfed. The discoverers (archaeologist Mike Morwood and colleagues) proposed that a variety of features, both primitive and derived, identify these individuals as belonging to a new species, H. floresiensis, within the taxonomic tribe of Hominini, which includes all species that are more closely related to humans than to chimpanzees. Based on previous date estimates, the discoverers also proposed that H. floresiensis lived contemporaneously with modern humans on Flores. Two orthopedic researches published in 2007 reported evidence to support species status for H. floresiensis. A study of three tokens of carpal (wrist) bones concluded there were differences from the carpal bones of modern humans and similarities to those of a chimpanzee or an early hominin such as Australopithecus. A study of the bones and joints of the arm, shoulder, and lower limbs also concluded that H. floresiensis was more similar to early humans and other apes than modern humans. In 2009, the publication of a cladistic analysis and a study of comparative body measurements provided further support for the hypothesis that H. floresiensis and Homo sapiens are separate species.

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Citations The Papers (from PubMed®)


RevDate: 2024-02-19
CmpDate: 2024-02-19

Bird EE, Kivell TL, Dunmore CJ, et al (2024)

Trabecular bone structure of the proximal capitate in extant hominids and fossil hominins with implications for midcarpal joint loading and the dart-thrower's motion.

American journal of biological anthropology, 183(3):e24824.

OBJECTIVES: This research examines whether the distribution of trabecular bone in the proximal capitates of extant hominids, as well as several fossil hominin taxa, is associated with the oblique path of the midcarpal joint known as the dart-thrower's motion (DTM).

MATERIALS AND METHODS: We analyzed proximal capitates from extant (Pongo n = 12; Gorilla n = 11; Pan n = 10; fossil and recent Homo sapiens n = 29) and extinct (Australopithecus sediba n = 2; Homo naledi n = 1; Homo floresiensis n = 2; Neandertals n = 3) hominids using a new canonical holistic morphometric analysis, which quantifies and visualizes the distribution of trabecular bone using relative bone volume as a fraction of total volume (rBV/TV).

RESULTS: Homo sapiens and Neandertals had a continuous band of high rBV/TV that extended across the scaphoid, lunate, and hamate subarticular regions, but other fossil hominins and extant great apes did not. A. sediba expressed a distinct combination of human-like and Pan-like rBV/TV distribution. Both H. floresiensis and H. naledi had high rBV/TV on the ulnar-side of the capitate but low rBV/TV on the radial-side.

CONCLUSION: The proximal capitates of H. sapiens and Neandertals share a distinctive distribution of trabecular bone that suggests that these two species of Homo regularly load(ed) their midcarpal joints along the full extent of the oblique path of the DTM. The observed pattern in A. sediba suggests that human-like stress at the capito-scaphoid articular surface was combined with Pan-like wrist postures, whereas the patterns in H. floresiensis and H. naledi suggest their midcarpal joints were loaded differently from that of H. sapiens and Neandertals.

RevDate: 2023-11-16
CmpDate: 2023-02-28

Pestana C, de Sousa AA, Todorov OS, et al (2023)

Evolutionary history of hominin brain size and phylogenetic comparative methods.

Progress in brain research, 275:217-232.

An absolutely and relatively large brain has traditionally been viewed as a distinctive characteristic of the Homo genus, with anatomically modern humans presented at the apex of a long line of progressive increases in encephalization. Many studies continue to focus attention on increasing brain size in the Homo genus, while excluding measures of absolute and relative brain size of more geologically recent, smaller brained, hominins such as Homo floresiensis, and Homo naledi and smaller brained Homo erectus specimens. This review discusses the benefits of using phylogenetic comparative methods to trace the diverse changes in hominin brain evolution and the drawbacks of not doing so.

RevDate: 2022-04-29
CmpDate: 2022-04-29

Scerri EML, Roberts P, Yoshi Maezumi S, et al (2022)

Tropical forests in the deep human past.

Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 377(1849):20200500.

Since Darwin, studies of human evolution have tended to give primacy to open 'savannah' environments as the ecological cradle of our lineage, with dense tropical forests cast as hostile, unfavourable frontiers. These perceptions continue to shape both the geographical context of fieldwork as well as dominant narratives concerning hominin evolution. This paradigm persists despite new, ground-breaking research highlighting the role of tropical forests in the human story. For example, novel research in Africa's rainforests has uncovered archaeological sites dating back into the Pleistocene; genetic studies have revealed very deep human roots in Central and West Africa and in the tropics of Asia and the Pacific; an unprecedented number of coexistent hominin species have now been documented, including Homo erectus, the 'Hobbit' (Homo floresiensis), Homo luzonensis, Denisovans, and Homo sapiens. Some of the earliest members of our own species to reach South Asia, Southeast Asia, Oceania and the tropical Americas have shown an unexpected rapidity in their adaptation to even some of the more 'extreme' tropical settings. This includes the early human manipulation of species and even habitats. This volume builds on these currently disparate threads and, for the first time, draws together a group of interdisciplinary, agenda-setting papers that firmly places a broader spectrum of tropical environments at the heart of the deep human past. This article is part of the theme issue 'Tropical forests in the deep human past'.

RevDate: 2022-03-09
CmpDate: 2022-03-09

Wu XJ, Bae CJ, Friess M, et al (2022)

Evolution of cranial capacity revisited: A view from the late Middle Pleistocene cranium from Xujiayao, China.

Journal of human evolution, 163:103119.

The Late Middle Pleistocene hominin fossils from the Xujiayao site in northern China have been closely studied in light of their morphological variability. However, all previous studies have focused on separated cranial fragments. Here, we report the first reconstruction of a fairly complete posterior cranium, Xujiayao 6 (XJY 6), confidently dated to ∼200-160 ka, which facilitated an assessment of its overall cranial size. XJY 6 was reconstructed from three of the original fragments-the PA1486 (No.7/XJY 6a) occipital bone, PA1490 (No.10/XJY 6b) right parietal bone, and PA1498 (No.17/XJY 15) left temporal bone-which originated from the same young adult individual. The XJY 6 endocranial capacity, estimated by measuring endocranial volume, was estimated using multiple regression formulae derived from ectocranial and endocranial measurements on select samples of Pleistocene hominins and recent modern humans. The results indicate that the larger pooled sample of both Pleistocene and recent modern humans was more robust for the endocranial capacity estimate. Based on the pooled sample using the ectocranial and endocranial measurements, we conservatively estimate the XJY 6 endocranial volume to be ∼1700 cm[3] with a 95% confidence interval of 1555-1781 cm[3]. This is close to Xuchang 1, which dates to 125-105 ka and whose endocranial volume is ∼1800 cm[3]. Thus, XJY 6 provides the earliest evidence of a brain size that falls in the upper range of Neanderthals and modern Homo sapiens. XJY 6, together with Xuchang 1, Homo floresiensis, Homo luzonensis, and Homo naledi, challenge the general pattern that brain size gradually increases over geological time. This study also finds that hominin brain size expansion occurred at different rates across time and space.

RevDate: 2022-05-31
CmpDate: 2022-03-09

Zanolli C, Kaifu Y, Pan L, et al (2022)

Further analyses of the structural organization of Homo luzonensis teeth: Evolutionary implications.

Journal of human evolution, 163:103124.

The species Homo luzonensis has recently been described based on a set of dental and postcranial elements found at Callao Cave (Northern Luzon, Philippines) and dated to at least 50-67 ka. Seven postcanine maxillary teeth are attributed to this taxon, five of them belonging to the same individual (CCH6) and representing the holotype of H. luzonensis, whereas the isolated upper premolar CCH8 and the upper third molar CCH9 are paratypes of the species. The teeth are characterized by their small dimensions associated with primitive features, as also found in Homo floresiensis, another hominin having evolved in an insular environment of Southeast Asia. Postcranial bones of the hands and feet of H. luzonensis and H. floresiensis show Homo habilis-like or australopith-like features, whereas cranial and dental morphology are more consistent with the Asian Homo erectus morphology. Due to this mosaic morphology, the origin and phylogenetic relationships of both H. luzonensis and H. floresiensis are still debated. To test the hypotheses that H. luzonensis derives from H. erectus or from an earlier small-brained hominin, we analyzed the µCT scans of the teeth. We investigated both external and internal tooth structure using morphometric methods including: crown outline shape, tooth crown tissue proportions, enamel-dentine junction shape, and pulp morphology. Homo luzonensis external crown morphology aligns more with H. erectus than with H. habilis/H. rudolfensis. The internal structural organization of H. luzonensis teeth exhibits more affinities with that of H. erectus and H. floresiensis than with Neanderthals and modern humans. Our results suggest that both H. floresiensis and H. luzonensis likely evolved from some H. erectus groups that dispersed in the various islands of this region and became isolated until endemic speciation events occurred at least twice during the Pleistocene in insular environments.

RevDate: 2022-02-04

Cook RW, Vazzana A, Sorrentino R, et al (2021)

The cranial biomechanics and feeding performance of Homo floresiensis.

Interface focus, 11(5):20200083.

Homo floresiensis is a small-bodied hominin from Flores, Indonesia, that exhibits plesiomorphic dentognathic features, including large premolars and a robust mandible, aspects of which have been considered australopith-like. However, relative to australopith species, H. floresiensis exhibits reduced molar size and a cranium with diminutive midfacial dimensions similar to those of later Homo, suggesting a reduction in the frequency of forceful biting behaviours. Our study uses finite-element analysis to examine the feeding biomechanics of the H. floresiensis cranium. We simulate premolar (P[3]) and molar (M[2]) biting in a finite-element model (FEM) of the H. floresiensis holotype cranium (LB1) and compare the mechanical results with FEMs of chimpanzees, modern humans and a sample of australopiths (MH1, Sts 5, OH5). With few exceptions, strain magnitudes in LB1 resemble elevated levels observed in modern Homo. Our analysis of LB1 suggests that H. floresiensis could produce bite forces with high mechanical efficiency, but was subject to tensile jaw joint reaction forces during molar biting, which perhaps constrained maximum postcanine bite force production. The inferred feeding biomechanics of H. floresiensis closely resemble modern humans, suggesting that this pattern may have been present in the last common ancestor of Homo sapiens and H. floresiensis.

RevDate: 2021-11-18
CmpDate: 2021-10-26

Prabhat AM, Miller CK, Prang TC, et al (2021)

Homoplasy in the evolution of modern human-like joint proportions in Australopithecus afarensis.

eLife, 10:.

The evolution of bipedalism and reduced reliance on arboreality in hominins resulted in larger lower limb joints relative to the joints of the upper limb. The pattern and timing of this transition, however, remains unresolved. Here, we find the limb joint proportions of Australopithecus afarensis, Homo erectus, and Homo naledi to resemble those of modern humans, whereas those of A. africanus, Australopithecus sediba, Paranthropus robustus, Paranthropus boisei, Homo habilis, and Homo floresiensis are more ape-like. The homology of limb joint proportions in A. afarensis and modern humans can only be explained by a series of evolutionary reversals irrespective of differing phylogenetic hypotheses. Thus, the independent evolution of modern human-like limb joint proportions in A. afarensis is a more parsimonious explanation. Overall, these results support an emerging perspective in hominin paleobiology that A. afarensis was the most terrestrially adapted australopith despite the importance of arboreality throughout much of early hominin evolution.

RevDate: 2021-03-15
CmpDate: 2021-03-15

Veatch EG, Ringen EJ, Kilgore MB, et al (2021)

Using niche construction theory to generate testable foraging hypotheses at Liang Bua.

Evolutionary anthropology, 30(1):8-16.

Niche construction theory (NCT) has emerged as a promising theoretical tool for interpreting zooarchaeological material. However, its juxtaposition against more established frameworks like optimal foraging theory (OFT) has raised important criticism around the testability of NCT for interpreting hominin foraging behavior. Here, we present an optimization foraging model with NCT features designed to consider the destructive realities of the archaeological record after providing a brief review of OFT and NCT. Our model was designed to consider a foragers decision to exploit an environment given predation risk, mortality, and payoff ratios between different ecologies, like more-open or more-forested environments. We then discuss how the model can be used with zooarchaeological data for inferring environmental exploitation by a primitive hominin, Homo floresiensis, from the island of Flores in Southeast Asia. Our example demonstrates that NCT can be used in combination with OFT principles to generate testable foraging hypotheses suitable for zooarchaeological research.

RevDate: 2021-07-26
CmpDate: 2021-07-26

Young CB (2020)

Static allometry of a small-bodied omnivore: body size and limb scaling of an island fox and inferences for Homo floresiensis.

Journal of human evolution, 149:102899.

Island dwarfing is a paraphyletic adaptation across numerous mammalian genera. From mammoths to foxes, extreme body size reduction is shared by diverse organisms that migrate to an island environment. Because it largely occurs owing to ecological variables, not phylogenetic ones, skeletal characters in a dwarfed taxon compared with its ancestor may appear abnormal. As a result, allometric patterns between body size and morphological traits may differ for an island dwarf compared with its ancestor. The diminutive Late Pleistocene hominin, Homo floresiensis, displays a unique character suite that is outside of the normal range of variation for any extinct or extant hominin species. To better explain these as ecological traits due to island dwarfing, this research looks at how dwarfing on islands influences limb scaling and proportions in an organism in a similar ecological niche as H. floresiensis. Here, I analyze absolute limb lengths and static allometry of limb lengths regressed on predicted body mass of dwarfed island foxes and their nondwarfed relatives. Dwarfed island foxes have significantly smaller intercepts but steeper slopes of all limb elements regressed on predicted body mass than the mainland gray fox. These allometric alterations produce limbs in the island fox that are significantly shorter than predicted for a nondwarfed gray fox of similar body mass. In addition, the humerofemoral, intermembral, and brachial indices are significantly different. These results provide a novel model for understanding skeletal variation of island endemic forms. Unique body size and proportions of H. floresiensis are plausible as ecological adaptations and likely not examples of symplesiomorphies with Australopithecus sp. Caution should be exerted when comparing an island dwarf with a closely related species as deviations from allometric expectations may be common.

RevDate: 2021-04-19
CmpDate: 2021-04-19

Scardia G, Neves WA, Tattersall I, et al (2021)

What kind of hominin first left Africa?.

Evolutionary anthropology, 30(2):122-127.

Recent discoveries of stone tools from Jordan (2.5 Ma) and China (2.1 Ma) document hominin presence in Asia at the beginning of the Pleistocene, well before the conventional Dmanisi datum at 1.8 Ma. Although no fossil hominins documenting this earliest Out of Africa phase have been found, on chronological grounds a pre-Homo erectus hominin must be considered the most likely maker of those artifacts. If so, this sheds new light on at least two disputed subjects in paleoanthropology, namely the remarkable variation among the five Dmanisi skulls, and the ancestry of Homo floresiensis.

RevDate: 2021-05-31
CmpDate: 2021-05-31

Ihara Y, Ikeya K, Nobayashi A, et al (2020)

A demographic test of accidental versus intentional island colonization by Pleistocene humans.

Journal of human evolution, 145:102839.

This study evaluates the hypothesis that some documented cases of long-distance sea crossing by the Late Pleistocene Homo sapiens occurred as a result of accidental drifting, rather than by intentional seafaring. For that purpose, we use an existing computer simulation framework, with some modifications, to investigate the likelihood that a planned or unplanned island colonization by a small group of individuals will persist to establish a viable population. Within the original framework, planned colonization was operationally characterized as being initiated by equal numbers of unrelated young men and women, whereas for unplanned colonization, those who migrate inadvertently were regarded in effect as a random sample of the whole population. Here, we consider a different scenario for unplanned colonization, which we believe is more relevant to sea crossing by the Late Pleistocene humans, that is, we assume that unplanned colonization occurs when members of households on watercrafts with limited voyaging capabilities are drifted away by ocean currents and washed up on a distant island. We also extend the previous analysis by considering a broader range of combinations of fertility and mortality schedules that individuals are assumed to follow. Our simulations suggest the following: (1) colonization of an island by ten or fewer unrelated young men and women can be successful within the feasible range of fertility and mortality levels; (2) in comparison, the likelihood of success for unplanned colonization is considerably smaller for the same range of fertility and mortality levels; and (3) there exists a small range of parameter combinations for which unplanned colonization has a non-negligible prospect of success even without assuming recurrent accidental drifts to the same island, and thus, the accidental colonization scenario cannot be totally excluded. In addition, we find that the minimum founding population required for successful colonization varies substantially depending on the fertility and mortality levels.

RevDate: 2021-03-25
CmpDate: 2020-05-08

Grün R, Pike A, McDermott F, et al (2020)

Dating the skull from Broken Hill, Zambia, and its position in human evolution.

Nature, 580(7803):372-375.

The cranium from Broken Hill (Kabwe) was recovered from cave deposits in 1921, during metal ore mining in what is now Zambia[1]. It is one of the best-preserved skulls of a fossil hominin, and was initially designated as the type specimen of Homo rhodesiensis, but recently it has often been included in the taxon Homo heidelbergensis[2-4]. However, the original site has since been completely quarried away, and-although the cranium is often estimated to be around 500 thousand years old[5-7]-its unsystematic recovery impedes its accurate dating and placement in human evolution. Here we carried out analyses directly on the skull and found a best age estimate of 299 ± 25 thousand years (mean ± 2σ). The result suggests that later Middle Pleistocene Africa contained multiple contemporaneous hominin lineages (that is, Homo sapiens[8,9], H. heidelbergensis/H. rhodesiensis and Homo naledi[10,11]), similar to Eurasia, where Homo neanderthalensis, the Denisovans, Homo floresiensis, Homo luzonensis and perhaps also Homo heidelbergensis and Homo erectus[12] were found contemporaneously. The age estimate also raises further questions about the mode of evolution of H. sapiens in Africa and whether H. heidelbergensis/H. rhodesiensis was a direct ancestor of our species[13,14].

RevDate: 2022-12-07
CmpDate: 2020-04-02

Wall JD, Ratan A, Stawiski E, et al (2019)

Identification of African-Specific Admixture between Modern and Archaic Humans.

American journal of human genetics, 105(6):1254-1261.

Recent work has demonstrated that two archaic human groups (Neanderthals and Denisovans) interbred with modern humans and contributed to the contemporary human gene pool. These findings relied on the availability of high-coverage genomes from both Neanderthals and Denisovans. Here we search for evidence of archaic admixture from a worldwide panel of 1,667 individuals using an approach that does not require the presence of an archaic human reference genome. We find no evidence for archaic admixture in the Andaman Islands, as previously claimed, or on the island of Flores, where Homo floresiensis fossils have been found. However, we do find evidence for at least one archaic admixture event in sub-Saharan Africa, with the strongest signal in Khoesan and Pygmy individuals from Southern and Central Africa. The locations of these putative archaic admixture tracts are weighted against functional regions of the genome, consistent with the long-term effects of purifying selection against introgressed genetic material.

RevDate: 2020-10-01
CmpDate: 2019-12-12

Diniz-Filho JAF, Jardim L, Rangel TF, et al (2019)

Quantitative genetics of body size evolution on islands: an individual-based simulation approach.

Biology letters, 15(10):20190481.

According to the island rule, small-bodied vertebrates will tend to evolve larger body size on islands, whereas the opposite happens to large-bodied species. This controversial pattern has been studied at the macroecological and biogeographical scales, but new developments in quantitative evolutionary genetics now allow studying the island rule from a mechanistic perspective. Here, we develop a simulation approach based on an individual-based model to model body size change on islands as a progressive adaptation to a moving optimum, determined by density-dependent population dynamics. We applied the model to evaluate body size differentiation in the pigmy extinct hominin Homo floresiensis, showing that dwarfing may have occurred in only about 360 generations (95% CI ranging from 150 to 675 generations). This result agrees with reports suggesting rapid dwarfing of large mammals on islands, as well as with the recent discovery that small-sized hominins lived in Flores as early as 700 kyr ago. Our simulations illustrate the power of analysing ecological and evolutionary patterns from an explicit quantitative genetics perspective.

RevDate: 2020-07-14
CmpDate: 2020-07-14

Bailey SE, Brophy JK, Moggi-Cecchi J, et al (2019)

The deciduous dentition of Homo naledi: A comparative study.

Journal of human evolution, 136:102655.

In 2013, 2014 new hominin remains were uncovered in the Dinaledi chamber of the Rising Star cave system in South Africa. In 2015 Berger and colleagues identified these remains as belonging to a new species Homo naledi (Berger et al., 2015). Subsequent comparative studies of the skull, postcrania and permanent dentition have supported this taxonomic affiliation (Harcourt-Smith et al., 2015; Kivell et al., 2015; Irish et al., 2018). The deciduous teeth can offer unique insights into hominin evolution. Due to their early onset and rapid development their morphology is thought to be under stronger genetic control and less influenced by environment than are the permanent teeth. In this study we compared the H. naledi deciduous teeth from the 2013-2014 excavations to samples representing much of the hominin clade including Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, early Homo, Homo antecessor, Homo erectus s.l., Homo floresiensis, Middle Pleistocene Homo, Homo neanderthalensis, early Homo sapiens and recent H. sapiens from Sub-Saharan Africa. By making such a broad morphological comparison, we aimed to contextualize the Dinaledi hominins and to further assess the validity of their taxonomic assignment. Our analysis of the deciduous teeth revealed a unique combination of features that mirror (but also expand) that found in the permanent teeth. This mosaic includes an asymmetrical lower canine with a distal tubercle, an upper first molar with a large hypocone and epicrista associated with a mesial cuspule, a molarized lower first molar resembling Paranthropus, and upper and lower second molars that resemble later Homo in their lack of accessory cusps. The unique combination of deciduous dental characters supports previous studies assigning H. naledi to a new species, although its phylogenetic position vis-à-vis other Homo species remains ambiguous.

RevDate: 2020-06-12
CmpDate: 2020-06-12

Veatch EG, Tocheri MW, Sutikna T, et al (2019)

Temporal shifts in the distribution of murine rodent body size classes at Liang Bua (Flores, Indonesia) reveal new insights into the paleoecology of Homo floresiensis and associated fauna.

Journal of human evolution, 130:45-60.

Liang Bua, the type locality of Homo floresiensis, is a limestone cave located in the western part of the Indonesian island of Flores. The relatively continuous stratigraphic sequence of the site spans the past ∼190 kyr and contains ∼275,000 taxonomically identifiable vertebrate skeletal elements, ∼80% of which belong to murine rodent taxa (i.e., rats). Six described genera are present at Liang Bua (Papagomys, Spelaeomys, Hooijeromys, Komodomys, Paulamys, and Rattus), one of which, Hooijeromys, is newly recorded in the site deposits, being previously known only from Early to Middle Pleistocene sites in central Flores. Measurements of the proximal femur (n = 10,212) and distal humerus (n = 1186) indicate five murine body size classes ranging from small (mouse-sized) to giant (common rabbit-sized) are present. The proportions of these five classes across successive stratigraphic units reveal two major changes in murine body size distribution due to significant shifts in the abundances of more open habitat-adapted medium-sized murines versus more closed habitat-adapted smaller-sized ones. One of these changes suggests a modest increase in available open habitats occurred ∼3 ka, likely the result of anthropogenic changes to the landscape related to farming by modern human populations. The other and more significant change occurred ∼60 ka suggesting a rapid shift from more open habitats to more closed conditions at this time. The abrupt reduction of medium-sized murines, along with the disappearance of H. floresiensis, Stegodon florensis insularis (an extinct proboscidean), Varanus komodoensis (Komodo dragon), Leptoptilos robustus (giant marabou stork), and Trigonoceps sp. (vulture) at Liang Bua ∼60-50 ka, is likely the consequence of these animals preferring and tracking more open habitats to elsewhere on the island. If correct, then the precise timing and nature of the extinction of H. floresiensis and its contemporaries must await new discoveries at Liang Bua or other as yet unexcavated sites on Flores.

RevDate: 2021-07-06
CmpDate: 2019-10-31

Détroit F, Mijares AS, Corny J, et al (2019)

A new species of Homo from the Late Pleistocene of the Philippines.

Nature, 568(7751):181-186.

A hominin third metatarsal discovered in 2007 in Callao Cave (Northern Luzon, the Philippines) and dated to 67 thousand years ago provided the earliest direct evidence of a human presence in the Philippines. Analysis of this foot bone suggested that it belonged to the genus Homo, but to which species was unclear. Here we report the discovery of twelve additional hominin elements that represent at least three individuals that were found in the same stratigraphic layer of Callao Cave as the previously discovered metatarsal. These specimens display a combination of primitive and derived morphological features that is different from the combination of features found in other species in the genus Homo (including Homo floresiensis and Homo sapiens) and warrants their attribution to a new species, which we name Homo luzonensis. The presence of another and previously unknown hominin species east of the Wallace Line during the Late Pleistocene epoch underscores the importance of island Southeast Asia in the evolution of the genus Homo.

RevDate: 2021-10-13

van den Hoek Ostende LW (2018)

Cladistics and insular evolution, an unfortunate marriage? Another tangle in the Deinogalerix analysis of Borrani et al. (2017).

Cladistics : the international journal of the Willi Hennig Society, 34(6):708-713.

Insular taxa, such as the late Miocene giant erinaceid Deinogalerix and the insular hominin Homo floresiensis, display a surprisingly primitive placement in cladistic analyses of their respective groups. This has led to speculations of early dispersal onto islands, not corroborated by biogeographical evidence. Insular evolution is notorious for vast phenotypic changes in at least part of the island fauna. These changes encompass typical apomorphies, but, as they may also include evolutionary reversals, a parsimony-based tree will inevitably yield a more basic position in cladograms. Therefore, we need to be cautious in drawing hasty conclusions from phylogenetic trees including insular taxa. Scrutiny is required to monitor possible evolutionary reversals and scenarios should be tested against biogeography to test their viability.

RevDate: 2019-10-02
CmpDate: 2019-10-02

Sutikna T, Tocheri MW, Faith JT, et al (2018)

The spatio-temporal distribution of archaeological and faunal finds at Liang Bua (Flores, Indonesia) in light of the revised chronology for Homo floresiensis.

Journal of human evolution, 124:52-74.

Liang Bua, the type site of Homo floresiensis, is a limestone cave on the Indonesian island of Flores with sedimentary deposits currently known to range in age from about 190 thousand years (ka) ago to the present. Recent revision of the stratigraphy and chronology of this depositional sequence suggests that skeletal remains of H. floresiensis are between ∼100 and 60 ka old, while cultural evidence of this taxon occurs until ∼50 ka ago. Here we examine the compositions of the faunal communities and stone artifacts, by broad taxonomic groups and raw materials, throughout the ∼190 ka time interval preserved in the sequence. Major shifts are observed in both the faunal and stone artifact assemblages that reflect marked changes in paleoecology and hominin behavior, respectively. Our results suggest that H. floresiensis and Stegodon florensis insularis, along with giant marabou stork (Leptoptilos robustus) and vulture (Trigonoceps sp.), were likely extinct by ∼50 ka ago. Moreover, an abrupt and statistically significant shift in raw material preference due to an increased use of chert occurs ∼46 thousand calibrated radiocarbon ([14]C) years before present (ka cal. BP), a pattern that continues through the subsequent stratigraphic sequence. If an increased preference for chert does, in fact, characterize Homo sapiens assemblages at Liang Bua, as previous studies have suggested (e.g., Moore et al., 2009), then the shift observed here suggests that modern humans arrived on Flores by ∼46 ka cal. BP, which would be the earliest cultural evidence of modern humans in Indonesia.

RevDate: 2020-02-25
CmpDate: 2018-09-13

Tucci S, Vohr SH, McCoy RC, et al (2018)

Evolutionary history and adaptation of a human pygmy population of Flores Island, Indonesia.

Science (New York, N.Y.), 361(6401):511-516.

Flores Island, Indonesia, was inhabited by the small-bodied hominin species Homo floresiensis, which has an unknown evolutionary relationship to modern humans. This island is also home to an extant human pygmy population. Here we describe genome-scale single-nucleotide polymorphism data and whole-genome sequences from a contemporary human pygmy population living on Flores near the cave where H. floresiensis was found. The genomes of Flores pygmies reveal a complex history of admixture with Denisovans and Neanderthals but no evidence for gene flow with other archaic hominins. Modern individuals bear the signatures of recent positive selection encompassing the FADS (fatty acid desaturase) gene cluster, likely related to diet, and polygenic selection acting on standing variation that contributed to their short-stature phenotype. Thus, multiple independent instances of hominin insular dwarfism occurred on Flores.

RevDate: 2018-12-02
CmpDate: 2018-09-27

Ingicco T, van den Bergh GD, Jago-On C, et al (2018)

Earliest known hominin activity in the Philippines by 709 thousand years ago.

Nature, 557(7704):233-237.

Over 60 years ago, stone tools and remains of megafauna were discovered on the Southeast Asian islands of Flores, Sulawesi and Luzon, and a Middle Pleistocene colonization by Homo erectus was initially proposed to have occurred on these islands[1-4]. However, until the discovery of Homo floresiensis in 2003, claims of the presence of archaic hominins on Wallacean islands were hypothetical owing to the absence of in situ fossils and/or stone artefacts that were excavated from well-documented stratigraphic contexts, or because secure numerical dating methods of these sites were lacking. As a consequence, these claims were generally treated with scepticism [5] . Here we describe the results of recent excavations at Kalinga in the Cagayan Valley of northern Luzon in the Philippines that have yielded 57 stone tools associated with an almost-complete disarticulated skeleton of Rhinoceros philippinensis, which shows clear signs of butchery, together with other fossil fauna remains attributed to stegodon, Philippine brown deer, freshwater turtle and monitor lizard. All finds originate from a clay-rich bone bed that was dated to between 777 and 631 thousand years ago using electron-spin resonance methods that were applied to tooth enamel and fluvial quartz. This evidence pushes back the proven period of colonization [6] of the Philippines by hundreds of thousands of years, and furthermore suggests that early overseas dispersal in Island South East Asia by premodern hominins took place several times during the Early and Middle Pleistocene stages[1-4]. The Philippines therefore may have had a central role in southward movements into Wallacea, not only of Pleistocene megafauna [7] , but also of archaic hominins.

RevDate: 2018-08-02
CmpDate: 2018-08-02

Flohr S (2018)

Did the assumed partial skeleton LB1 (aka Homo floresiensis) really have long feet?.

Anthropologischer Anzeiger; Bericht uber die biologisch-anthropologische Literatur, 75(2):169-174.

About 100 hominin bones were found during excavations at the Liang Bua cave on the island of Flores, Indonesia. More than 60 of them were assigned to the partial skeleton LB1 which was designated as the holotype of a new species, Homo floresiensis. Analyses of skeletal proportions of LB1 led to the conclusion that its foot was exceptionally long relative to femur and tibia, respectively. This ratio was considered a unique feature that contributes to the definition of the new species. The published illustrations of the in situ-situation and the published inventory of the bones suggest a high degree of commingling rather than the presence of larger anatomically joining units that was asserted in the publications on the findings. The available information further suggests that hand and foot bones of several individuals were found commingled as well. Here I argue, based on the published data, that certain problems exist regarding the correct anatomical identification of some of the phalangeal bones that contributed to the results on which the conclusions about skeletal proportions in LB1 were based. It is further suggested that the assignment of bones to specific individuals is debatable. Conclusions on the taxonomic status of the Liang Bua hominins and their pattern of bipedalism based on these data therefore need to be substantiated by further studies. Specifically, on the basis of the available information, a large relative foot length should no longer be claimed as a unique feature of the presumed new species H. floresiensis.

RevDate: 2020-10-01

Will M, Pablos A, JT Stock (2017)

Long-term patterns of body mass and stature evolution within the hominin lineage.

Royal Society open science, 4(11):171339.

Body size is a central determinant of a species' biology and adaptive strategy, but the number of reliable estimates of hominin body mass and stature have been insufficient to determine long-term patterns and subtle interactions in these size components within our lineage. Here, we analyse 254 body mass and 204 stature estimates from a total of 311 hominin specimens dating from 4.4 Ma to the Holocene using multi-level chronological and taxonomic analytical categories. The results demonstrate complex temporal patterns of body size variation with phases of relative stasis intermitted by periods of rapid increases. The observed trajectories could result from punctuated increases at speciation events, but also differential proliferation of large-bodied taxa or the extinction of small-bodied populations. Combined taxonomic and temporal analyses show that in relation to australopithecines, early Homo is characterized by significantly larger average body mass and stature but retains considerable diversity, including small body sizes. Within later Homo, stature and body mass evolution follow different trajectories: average modern stature is maintained from ca 1.6 Ma, while consistently higher body masses are not established until the Middle Pleistocene at ca 0.5-0.4 Ma, likely caused by directional selection related to colonizing higher latitudes. Selection against small-bodied individuals (less than 40 kg; less than 140 cm) after 1.4 Ma is associated with a decrease in relative size variability in later Homo species compared with earlier Homo and australopithecines. The isolated small-bodied individuals of Homo naledi (ca 0.3 Ma) and Homo floresiensis (ca 100-60 ka) constitute important exceptions to these general patterns, adding further layers of complexity to the evolution of body size within the genus Homo. At the end of the Late Pleistocene and Holocene, body size in Homo sapiens declines on average, but also extends to lower limits not seen in comparable frequency since early Homo.

RevDate: 2019-10-02
CmpDate: 2019-10-02

VanSickle C, Cofran Z, García-Martínez D, et al (2018)

Homo naledi pelvic remains from the Dinaledi Chamber, South Africa.

Journal of human evolution, 125:122-136.

In the hominin fossil record, pelvic remains are sparse and are difficult to attribute taxonomically when they are not directly associated with craniodental material. Here we describe the pelvic remains from the Dinaledi Chamber in the Rising Star cave system, Cradle of Humankind, South Africa, which has produced hominin fossils of a new species, Homo naledi. Though this species has been attributed to Homo based on cranial and lower limb morphology, the morphology of some of the fragmentary pelvic remains recovered align more closely with specimens attributed to the species Australopithecus afarensis and Australopithecus africanus than they do with those of most (but not all) known species of the genus Homo. As with A. afarensis and A. africanus, H. naledi appears to have had marked lateral iliac flare and either a weakly developed or non-existent acetabulocristal buttress or a distinct, albeit weakly developed, acetabulospinous buttress. At the same time, H. naledi has robust superior pubic and ischiopubic rami and a short ischium with a narrow tuberoacetabular sulcus, similar to those found in modern humans. The fragmentary nature of the Dinaledi pelvic assemblage makes the attribution of sex and developmental age to individual specimens difficult, which in turn diminishes our ability to identify the number of individuals represented in the assemblage. At present, we can only confidently say that the pelvic fossils from Rising Star represent at least four individuals based on the presence of four overlapping right ischial fossils (whereas a minimum of 15 individuals can be identified from the Dinaledi dental assemblage). A primitive, early Australopithecus-like false pelvis combined with a derived Homo-like true pelvis is morphologically consistent with evidence from the lower ribcage and proximal femur of H. naledi. The overall similarity of H. naledi ilia to those of australopiths supports the inference, drawn from the observation of primitive pelvic morphology in the extinct species Homo floresiensis, that there is substantial variation in pelvic form within the genus Homo. In the light of these findings, we urge caution in making taxonomic attributions-even at the genus level-of isolated fossil ossa coxae.

RevDate: 2019-02-05
CmpDate: 2018-04-04

Garvin HM, Elliott MC, Delezene LK, et al (2017)

Body size, brain size, and sexual dimorphism in Homo naledi from the Dinaledi Chamber.

Journal of human evolution, 111:119-138.

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus, together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo, including Homo habilis, Homo rudolfensis, and some early H. erectus, suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus, with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates. H. naledi endocranial volumes (465-560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus, but similar to derived members of genus Homo, the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

RevDate: 2019-01-30
CmpDate: 2018-02-06

Turvey ST, Crees JJ, Hansford J, et al (2017)

Quaternary vertebrate faunas from Sumba, Indonesia: implications for Wallacean biogeography and evolution.

Proceedings. Biological sciences, 284(1861):.

Historical patterns of diversity, biogeography and faunal turnover remain poorly understood for Wallacea, the biologically and geologically complex island region between the Asian and Australian continental shelves. A distinctive Quaternary vertebrate fauna containing the small-bodied hominin Homo floresiensis, pygmy Stegodon proboscideans, varanids and giant murids has been described from Flores, but Quaternary faunas are poorly known from most other Lesser Sunda Islands. We report the discovery of extensive new fossil vertebrate collections from Pleistocene and Holocene deposits on Sumba, a large Wallacean island situated less than 50 km south of Flores. A fossil assemblage recovered from a Pleistocene deposit at Lewapaku in the interior highlands of Sumba, which may be close to 1 million years old, contains a series of skeletal elements of a very small Stegodon referable to S. sumbaensis, a tooth attributable to Varanus komodoensis, and fragmentary remains of unidentified giant murids. Holocene cave deposits at Mahaniwa dated to approximately 2000-3500 BP yielded extensive material of two new genera of endemic large-bodied murids, as well as fossils of an extinct frugivorous varanid. This new baseline for reconstructing Wallacean faunal histories reveals that Sumba's Quaternary vertebrate fauna, although phylogenetically distinctive, was comparable in diversity and composition to the Quaternary fauna of Flores, suggesting that similar assemblages may have characterized Quaternary terrestrial ecosystems on many or all of the larger Lesser Sunda Islands.

RevDate: 2020-09-30

Meijer HJM, Awe Due R, Sutikna T, et al (2017)

Late Pleistocene songbirds of Liang Bua (Flores, Indonesia); the first fossil passerine fauna described from Wallacea.

PeerJ, 5:e3676.

BACKGROUND: Passerines (Aves: Passeriformes) dominate modern terrestrial bird communities yet their fossil record is limited. Liang Bua is a large cave on the Indonesian island of Flores that preserves Late Pleistocene-Holocene deposits (∼190 ka to present day). Birds are the most diverse faunal group at Liang Bua and are present throughout the stratigraphic sequence.

METHODS: We examined avian remains from the Late Pleistocene deposits of Sector XII, a 2 × 2 m area excavated to about 8.5 m depth. Although postcranial passerine remains are typically challenging to identify, we found several humeral characters particularly useful in discriminating between groups, and identified 89 skeletal elements of passerines.

RESULTS: At least eight species from eight families are represented, including the Large-billed Crow (Corvus cf. macrorhynchos), the Australasian Bushlark (Mirafra javanica), a friarbird (Philemon sp.), and the Pechora Pipit (Anthus cf. gustavi).

DISCUSSION: These remains constitute the first sample of fossil passerines described in Wallacea. Two of the taxa no longer occur on Flores today; a large sturnid (cf. Acridotheres) and a grassbird (Megalurus sp.). Palaeoecologically, the songbird assemblage suggests open grassland and tall forests, which is consistent with conditions inferred from the non-passerine fauna at the site. Corvus cf. macrorhynchos, found in the Homo floresiensis-bearing layers, was likely part of a scavenging guild that fed on carcasses of Stegodon florensis insularis alongside vultures (Trigonoceps sp.), giant storks (Leptoptilos robustus), komodo dragons (Varanus komodoensis), and probably H. floresiensis as well.

RevDate: 2019-01-30
CmpDate: 2018-02-01

Diniz-Filho JAF, P Raia (2017)

Island Rule, quantitative genetics and brain-body size evolution in Homo floresiensis.

Proceedings. Biological sciences, 284(1857):.

Colonization of islands often activate a complex chain of adaptive events that, over a relatively short evolutionary time, may drive strong shifts in body size, a pattern known as the Island Rule. It is arguably difficult to perform a direct analysis of the natural selection forces behind such a change in body size. Here, we used quantitative evolutionary genetic models, coupled with simulations and pattern-oriented modelling, to analyse the evolution of brain and body size in Homo floresiensis, a diminutive hominin species that appeared around 700 kya and survived up to relatively recent times (60-90 kya) on Flores Island, Indonesia. The hypothesis of neutral evolution was rejected in 97% of the simulations, and estimated selection gradients are within the range found in living natural populations. We showed that insularity may have triggered slightly different evolutionary trajectories for body and brain size, which means explaining the exceedingly small cranial volume of H. floresiensis requires additional selective forces acting on brain size alone. Our analyses also support previous conclusions that H. floresiensis may be most likely derived from an early Indonesian H. erectus, which is coherent with currently accepted biogeographical scenario for Homo expansion out of Africa.

RevDate: 2018-12-02
CmpDate: 2018-04-04

Argue D, Groves CP, Lee MSY, et al (2017)

The affinities of Homo floresiensis based on phylogenetic analyses of cranial, dental, and postcranial characters.

Journal of human evolution, 107:107-133.

Although the diminutive Homo floresiensis has been known for a decade, its phylogenetic status remains highly contentious. A broad range of potential explanations for the evolution of this species has been explored. One view is that H. floresiensis is derived from Asian Homo erectus that arrived on Flores and subsequently evolved a smaller body size, perhaps to survive the constrained resources they faced in a new island environment. Fossil remains of H. erectus, well known from Java, have not yet been discovered on Flores. The second hypothesis is that H. floresiensis is directly descended from an early Homo lineage with roots in Africa, such as Homo habilis; the third is that it is Homo sapiens with pathology. We use parsimony and Bayesian phylogenetic methods to test these hypotheses. Our phylogenetic data build upon those characters previously presented in support of these hypotheses by broadening the range of traits to include the crania, mandibles, dentition, and postcrania of Homo and Australopithecus. The new data and analyses support the hypothesis that H. floresiensis is an early Homo lineage: H. floresiensis is sister either to H. habilis alone or to a clade consisting of at least H. habilis, H. erectus, Homo ergaster, and H. sapiens. A close phylogenetic relationship between H. floresiensis and H. erectus or H. sapiens can be rejected; furthermore, most of the traits separating H. floresiensis from H. sapiens are not readily attributable to pathology (e.g., Down syndrome). The results suggest H. floresiensis is a long-surviving relict of an early (>1.75 Ma) hominin lineage and a hitherto unknown migration out of Africa, and not a recent derivative of either H. erectus or H. sapiens.

RevDate: 2017-12-19
CmpDate: 2017-08-03

Webber JT, DA Raichlen (2016)

The role of plantigrady and heel-strike in the mechanics and energetics of human walking with implications for the evolution of the human foot.

The Journal of experimental biology, 219(Pt 23):3729-3737.

Human bipedal locomotion is characterized by a habitual heel-strike (HS) plantigrade gait, yet the significance of walking foot-posture is not well understood. To date, researchers have not fully investigated the costs of non-heel-strike (NHS) walking. Therefore, we examined walking speed, walk-to-run transition speed, estimated locomotor costs (lower limb muscle volume activated during walking), impact transient (rapid increase in ground force at touchdown) and effective limb length (ELL) in subjects (n=14) who walked at self-selected speeds using HS and NHS gaits. HS walking increases ELL compared with NHS walking since the center of pressure translates anteriorly from heel touchdown to toe-off. NHS gaits led to decreased absolute walking speeds (P=0.012) and walk-to-run transition speeds (P=0.0025), and increased estimated locomotor energy costs (P<0.0001) compared with HS gaits. These differences lost significance after using the dynamic similarity hypothesis to account for the effects of foot landing posture on ELL. Thus, reduced locomotor costs and increased maximum walking speeds in HS gaits are linked to the increased ELL compared with NHS gaits. However, HS walking significantly increases impact transient values at all speeds (P<0.0001). These trade-offs may be key to understanding the functional benefits of HS walking. Given the current debate over the locomotor mechanics of early hominins and the range of foot landing postures used by nonhuman apes, we suggest the consistent use of HS gaits provides key locomotor advantages to striding bipeds and may have appeared early in hominin evolution.

RevDate: 2018-03-23
CmpDate: 2017-09-07

Marwick B, Clarkson C, O'Connor S, et al (2016)

Early modern human lithic technology from Jerimalai, East Timor.

Journal of human evolution, 101:45-64.

Jerimalai is a rock shelter in East Timor with cultural remains dated to 42,000 years ago, making it one of the oldest known sites of modern human activity in island Southeast Asia. It has special global significance for its record of early pelagic fishing and ancient shell fish hooks. It is also of regional significance for its early occupation and comparatively large assemblage of Pleistocene stone artefacts. Three major findings arise from our study of the stone artefacts. First, there is little change in lithic technology over the 42,000 year sequence, with the most noticeable change being the addition of new artefact types and raw materials in the mid-Holocene. Second, the assemblage is dominated by small chert cores and implements rather than pebble tools and choppers, a pattern we argue pattern, we argue, that is common in island SE Asian sites as opposed to mainland SE Asian sites. Third, the Jerimalai assemblage bears a striking resemblance to the assemblage from Liang Bua, argued by the Liang Bua excavation team to be associated with Homo floresiensis. We argue that the near proximity of these two islands along the Indonesian island chain (c.100 km apart), the long antiquity of modern human occupation in the region (as documented at Jerimalai), and the strong resemblance of distinctive flake stone technologies seen at both sites, raises the intriguing possibility that both the Liang Bua and Jerimalai assemblages were created by modern humans.

RevDate: 2018-03-23
CmpDate: 2017-09-21

Dembo M, Radovčić D, Garvin HM, et al (2016)

The evolutionary relationships and age of Homo naledi: An assessment using dated Bayesian phylogenetic methods.

Journal of human evolution, 97:17-26.

Homo naledi is a recently discovered species of fossil hominin from South Africa. A considerable amount is already known about H. naledi but some important questions remain unanswered. Here we report a study that addressed two of them: "Where does H. naledi fit in the hominin evolutionary tree?" and "How old is it?" We used a large supermatrix of craniodental characters for both early and late hominin species and Bayesian phylogenetic techniques to carry out three analyses. First, we performed a dated Bayesian analysis to generate estimates of the evolutionary relationships of fossil hominins including H. naledi. Then we employed Bayes factor tests to compare the strength of support for hypotheses about the relationships of H. naledi suggested by the best-estimate trees. Lastly, we carried out a resampling analysis to assess the accuracy of the age estimate for H. naledi yielded by the dated Bayesian analysis. The analyses strongly supported the hypothesis that H. naledi forms a clade with the other Homo species and Australopithecus sediba. The analyses were more ambiguous regarding the position of H. naledi within the (Homo, Au. sediba) clade. A number of hypotheses were rejected, but several others were not. Based on the available craniodental data, Homo antecessor, Asian Homo erectus, Homo habilis, Homo floresiensis, Homo sapiens, and Au. sediba could all be the sister taxon of H. naledi. According to the dated Bayesian analysis, the most likely age for H. naledi is 912 ka. This age estimate was supported by the resampling analysis. Our findings have a number of implications. Most notably, they support the assignment of the new specimens to Homo, cast doubt on the claim that H. naledi is simply a variant of H. erectus, and suggest H. naledi is younger than has been previously proposed.

RevDate: 2018-11-13
CmpDate: 2016-07-06

Brumm A, van den Bergh GD, Storey M, et al (2016)

Age and context of the oldest known hominin fossils from Flores.

Nature, 534(7606):249-253.

Recent excavations at the early Middle Pleistocene site of Mata Menge in the So'a Basin of central Flores, Indonesia, have yielded hominin fossils attributed to a population ancestral to Late Pleistocene Homo floresiensis. Here we describe the age and context of the Mata Menge hominin specimens and associated archaeological findings. The fluvial sandstone layer from which the in situ fossils were excavated in 2014 was deposited in a small valley stream around 700 thousand years ago, as indicated by (40)Ar/(39)Ar and fission track dates on stratigraphically bracketing volcanic ash and pyroclastic density current deposits, in combination with coupled uranium-series and electron spin resonance dating of fossil teeth. Palaeoenvironmental data indicate a relatively dry climate in the So'a Basin during the early Middle Pleistocene, while various lines of evidence suggest the hominins inhabited a savannah-like open grassland habitat with a wetland component. The hominin fossils occur alongside the remains of an insular fauna and a simple stone technology that is markedly similar to that associated with Late Pleistocene H. floresiensis.

RevDate: 2018-11-13
CmpDate: 2016-07-06

van den Bergh GD, Kaifu Y, Kurniawan I, et al (2016)

Homo floresiensis-like fossils from the early Middle Pleistocene of Flores.

Nature, 534(7606):245-248.

The evolutionary origin of Homo floresiensis, a diminutive hominin species previously known only by skeletal remains from Liang Bua in western Flores, Indonesia, has been intensively debated. It is a matter of controversy whether this primitive form, dated to the Late Pleistocene, evolved from early Asian Homo erectus and represents a unique and striking case of evolutionary reversal in hominin body and brain size within an insular environment. The alternative hypothesis is that H. floresiensis derived from an older, smaller-brained member of our genus, such as Homo habilis, or perhaps even late Australopithecus, signalling a hitherto undocumented dispersal of hominins from Africa into eastern Asia by two million years ago (2 Ma). Here we describe hominin fossils excavated in 2014 from an early Middle Pleistocene site (Mata Menge) in the So'a Basin of central Flores. These specimens comprise a mandible fragment and six isolated teeth belonging to at least three small-jawed and small-toothed individuals. Dating to ~0.7 Ma, these fossils now constitute the oldest hominin remains from Flores. The Mata Menge mandible and teeth are similar in dimensions and morphological characteristics to those of H. floresiensis from Liang Bua. The exception is the mandibular first molar, which retains a more primitive condition. Notably, the Mata Menge mandible and molar are even smaller in size than those of the two existing H. floresiensis individuals from Liang Bua. The Mata Menge fossils are derived compared with Australopithecus and H. habilis, and so tend to support the view that H. floresiensis is a dwarfed descendent of early Asian H. erectus. Our findings suggest that hominins on Flores had acquired extremely small body size and other morphological traits specific to H. floresiensis at an unexpectedly early time.

RevDate: 2018-12-02
CmpDate: 2016-07-06

Gómez-Robles A (2016)

Palaeoanthropology: The dawn of Homo floresiensis.

Nature, 534(7606):188-189.

RevDate: 2019-02-13
CmpDate: 2017-07-14

Baab KL, Brown P, Falk D, et al (2016)

A Critical Evaluation of the Down Syndrome Diagnosis for LB1, Type Specimen of Homo floresiensis.

PloS one, 11(6):e0155731.

The Liang Bua hominins from Flores, Indonesia, have been the subject of intense scrutiny and debate since their initial description and classification in 2004. These remains have been assigned to a new species, Homo floresiensis, with the partial skeleton LB1 as the type specimen. The Liang Bua hominins are notable for their short stature, small endocranial volume, and many features that appear phylogenetically primitive relative to modern humans, despite their late Pleistocene age. Recently, some workers suggested that the remains represent members of a small-bodied island population of modern Austro-Melanesian humans, with LB1 exhibiting clinical signs of Down syndrome. Many classic Down syndrome signs are soft tissue features that could not be assessed in skeletal remains. Moreover, a definitive diagnosis of Down syndrome can only be made by genetic analysis as the phenotypes associated with Down syndrome are variable. Most features that contribute to the Down syndrome phenotype are not restricted to Down syndrome but are seen in other chromosomal disorders and in the general population. Nevertheless, we re-evaluated the presence of those phenotypic features used to support this classification by comparing LB1 to samples of modern humans diagnosed with Down syndrome and euploid modern humans using comparative morphometric analyses. We present new data regarding neurocranial, brain, and symphyseal shape in Down syndrome, additional estimates of stature for LB1, and analyses of inter- and intralimb proportions. The presence of cranial sinuses is addressed using CT images of LB1. We found minimal congruence between the LB1 phenotype and clinical descriptions of Down syndrome. We present important differences between the phenotypes of LB1 and individuals with Down syndrome, and quantitative data that characterize LB1 as an outlier compared with Down syndrome and non-Down syndrome groups. Homo floresiensis remains a phenotypically unique, valid species with its roots in Plio-Pleistocene Homo taxa.

RevDate: 2016-12-30
CmpDate: 2016-12-13

Balzeau A, P Charlier (2016)

What do cranial bones of LB1 tell us about Homo floresiensis?.

Journal of human evolution, 93:12-24.

Cranial vault thickness (CVT) of Liang Bua 1, the specimen that is proposed to be the holotype of Homo floresiensis, has not yet been described in detail and compared with samples of fossil hominins, anatomically modern humans or microcephalic skulls. In addition, a complete description from a forensic and pathological point of view has not yet been carried out. It is important to evaluate scientifically if features related to CVT bring new information concerning the possible pathological status of LB1, and if it helps to recognize affinities with any hominin species and particularly if the specimen could belong to the species Homo sapiens. Medical examination of the skull based on a micro-CT examination clearly brings to light the presence of a sincipital T (a non-metrical variant of normal anatomy), a scar from an old frontal trauma without any evident functional consequence, and a severe bilateral hyperostosis frontalis interna that may have modified the anterior morphology of the endocranium of LB1. We also show that LB1 displays characteristics, related to the distribution of bone thickness and arrangements of cranial structures, that are plesiomorphic traits for hominins, at least for Homo erectus s.l. relative to Homo neanderthalensis and H. sapiens. All the microcephalic skulls analyzed here share the derived condition of anatomically modern H. sapiens. Cranial vault thickness does not help to clarify the definition of the species H. floresiensis but it also does not support an attribution of LB1 to H. sapiens. We conclude that there is no support for the attribution of LB1 to H. sapiens as there is no evidence of systemic pathology and because it does not have any of the apomorphic traits of our species.

RevDate: 2018-11-13
CmpDate: 2016-05-10

Sutikna T, Tocheri MW, Morwood MJ, et al (2016)

Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia.

Nature, 532(7599):366-369.

Homo floresiensis, a primitive hominin species discovered in Late Pleistocene sediments at Liang Bua (Flores, Indonesia), has generated wide interest and scientific debate. A major reason this taxon is controversial is because the H. floresiensis-bearing deposits, which include associated stone artefacts and remains of other extinct endemic fauna, were dated to between about 95 and 12 thousand calendar years (kyr) ago. These ages suggested that H. floresiensis survived until long after modern humans reached Australia by ~50 kyr ago. Here we report new stratigraphic and chronological evidence from Liang Bua that does not support the ages inferred previously for the H. floresiensis holotype (LB1), ~18 thousand calibrated radiocarbon years before present (kyr cal. BP), or the time of last appearance of this species (about 17 or 13-11 kyr cal. BP). Instead, the skeletal remains of H. floresiensis and the deposits containing them are dated to between about 100 and 60 kyr ago, whereas stone artefacts attributable to this species range from about 190 to 50 kyr in age. Whether H. floresiensis survived after 50 kyr ago--potentially encountering modern humans on Flores or other hominins dispersing through southeast Asia, such as Denisovans--is an open question.

RevDate: 2018-02-13
CmpDate: 2017-06-12

Prang TC (2016)

Conarticular congruence of the hominoid subtalar joint complex with implications for joint function in Plio-Pleistocene hominins.

American journal of physical anthropology, 160(3):446-457.

OBJECTIVE: The purpose of this study is to test the hypothesis that conarticular surfaces areas and curvatures are correlates of mobility at the hominoid talocalcaneal and talonavicular joints.

MATERIALS AND METHODS: Articular surface areas and curvatures of the talonavicular, anterior talocalcaneal, and posterior talocalcaneal joints were quantified using a total of 425 three-dimensional surface models of extant hominoid and fossil hominin tali, calcanei, and naviculars. Quadric surface fitting was used to calculate curvatures, pairwise comparisons were used to evaluate statistical differences between taxa, and regression was used to test for the effects of allometry.

RESULTS: Pairwise comparisons show that the distributions of values for joint curvature indices follow the predicted arboreal-terrestrial morphocline in hominoid primates with no effect of body mass (PGLS p > 0.05). OH 8 (Homo habilis) and LB 1 (Homo floresiensis) can be accommodated within the range of human variation for the talonavicular joint, whereas MH2 (Australopithecus sediba) falls within the ranges of variation for Pan troglodytes and Gorilla gorilla in measures of posterior talocalcaneal joint congruity.

CONCLUSIONS: Joint curvature indices are better discriminators than joint surface area indices, which may reflect a greater contribution of rotation, rather than translation, to joint movement in plantigrade taxa due to discrepancies in conarticular congruence and the "convex-concave" rule. The pattern of joint congruence in Au. sediba contributes to other data on the foot and ankle suggesting that the lateral side of the foot was more mobile than the medial side, which is consistent with suggestions of increased medial weight transfer associated with hyperpronation. Am J Phys Anthropol 160:446-457, 2016. © 2016 Wiley Periodicals, Inc.

RevDate: 2017-02-02
CmpDate: 2017-02-02

Baab K (2016)

The place of Homo floresiensis in human evolution.

Journal of anthropological sciences = Rivista di antropologia : JASS, 94:5-18.

Two main evolutionary scenarios have been proposed to explain the presence of the small-bodied and small-brained Homo floresiensis species on the remote Indonesian island of Flores in the Late Pleistocene. According to these two scenarios, H. floresiensis was a dwarfed descendent of H. erectus or a late-surviving remnant of a older lineage, perhaps descended from H. habilis. Each scenario has interesting and important implications for hominin biogeography, body size evolution, brain evolution and morphological convergences. Careful evaluation reveals that only a small number of characters support each of these scenarios uniquely. H. floresiensis exhibits a cranial shape and many cranial characters that appear to be shared derived traits with H. erectus, but postcranial traits are more primitive and resemble those of early Homo or even australopiths. Mandibular and dental traits show a mix of derived and primitive features. Unfortunately, many traits cannot be used to assess these two hypotheses because their distribution in H. erectus, early Homo (e.g., H. habilis), or both is unknown. H. erectus ancestry implies evolutionary convergence on a postcranial configuration similar to australopiths and early Homo, which could be explained by a return to more climbing behaviors. Body size reduction as well as brain size reduction on a scale only rarely documented in mammals would also accompany the origin of H. floresiensis from a H. erectus ancestor. H. habilis ancestry implies parallel evolution of numerous cranial characters, as well as a few dentognathic traits. A pre-H. erectus ancestry also suggests an early migration to Southeast Asia that is as yet undocumented in mainland Asia, but minimal body and brain size reduction.

RevDate: 2019-01-09
CmpDate: 2016-09-28

Prang TC (2015)

Rearfoot posture of Australopithecus sediba and the evolution of the hominin longitudinal arch.

Scientific reports, 5:17677.

The longitudinal arch is one of the hallmarks of the human foot but its evolutionary history remains controversial due to the fragmentary nature of the fossil record. In modern humans, the presence of a longitudinal arch is reflected in the angular relationships among the major surfaces of the human talus and calcaneus complex, which is also known as the rearfoot. A complete talus and calcaneus of Australopithecus sediba provide the opportunity to evaluate rearfoot posture in an early hominin for the first time. Here I show that A. sediba is indistinguishable from extant African apes in the angular configuration of its rearfoot, which strongly suggests that it lacked a longitudinal arch. Inferences made from isolated fossils support the hypothesis that Australopithecus afarensis possessed an arched foot. However, tali attributed to temporally younger taxa like Australopithecus africanus and Homo floresiensis are more similar to those of A. sediba. The inferred absence of a longitudinal arch in A. sediba would be biomechanically consistent with prior suggestions of increased midtarsal mobility in this taxon. The morphological patterns in talus and calcaneus angular relationships among fossil hominins suggest that there was diversity in traits associated with the longitudinal arch in the Plio-Pleistocene.

RevDate: 2018-11-13
CmpDate: 2016-06-21

Kaifu Y, Kono RT, Sutikna T, et al (2015)

Unique Dental Morphology of Homo floresiensis and Its Evolutionary Implications.

PloS one, 10(11):e0141614.

Homo floresiensis is an extinct, diminutive hominin species discovered in the Late Pleistocene deposits of Liang Bua cave, Flores, eastern Indonesia. The nature and evolutionary origins of H. floresiensis' unique physical characters have been intensively debated. Based on extensive comparisons using linear metric analyses, crown contour analyses, and other trait-by-trait morphological comparisons, we report here that the dental remains from multiple individuals indicate that H. floresiensis had primitive canine-premolar and advanced molar morphologies, a combination of dental traits unknown in any other hominin species. The primitive aspects are comparable to H. erectus from the Early Pleistocene, whereas some of the molar morphologies are more progressive even compared to those of modern humans. This evidence contradicts the earlier claim of an entirely modern human-like dental morphology of H. floresiensis, while at the same time does not support the hypothesis that H. floresiensis originated from a much older H. habilis or Australopithecus-like small-brained hominin species currently unknown in the Asian fossil record. These results are however consistent with the alternative hypothesis that H. floresiensis derived from an earlier Asian Homo erectus population and experienced substantial body and brain size dwarfism in an isolated insular setting. The dentition of H. floresiensis is not a simple, scaled-down version of earlier hominins.

RevDate: 2017-10-13
CmpDate: 2016-05-25

Larick R, RL Ciochon (2015)

Early hominin biogeography in Island Southeast Asia.

Evolutionary anthropology, 24(5):185-213.

Island Southeast Asia covers Eurasia's tropical expanse of continental shelf and active subduction zones. Cutting between island landmasses, Wallace's Line separates Sunda and the Eastern Island Arc (the Arc) into distinct tectonic and faunal provinces. West of the line, on Sunda, Java Island yields many fossils of Homo erectus. East of the line, on the Arc, Flores Island provides one skeleton and isolated remains of Homo floresiensis. Luzon Island in the Philippines has another fossil hominin. Sulawesi preserves early hominin archeology. This insular divergence sets up a unique regional context for early hominin dispersal, isolation, and extinction. The evidence is reviewed across three Pleistocene climate periods. Patterns are discussed in relation to the pulse of global sea-level shifts, as well as regional geo-tectonics, catastrophes, stegodon dispersal, and paleogenomics. Several patterns imply evolutionary processes typical of oceanic islands. Early hominins apparently responded to changing island conditions for a million-and-a-half years, likely becoming extinct during the period in which Homo sapiens colonized the region.

RevDate: 2019-01-08
CmpDate: 2015-05-04

Westaway MC, Durband AC, Groves CP, et al (2015)

Mandibular evidence supports Homo floresiensis as a distinct species.

Proceedings of the National Academy of Sciences of the United States of America, 112(7):E604-5.

RevDate: 2018-12-02
CmpDate: 2015-05-04

Eckhardt RB, Henneberg M, Chavanaves S, et al (2015)

Reply to Westaway et al.: Mandibular misrepresentations fail to support the invalid species Homo floresiensis.

Proceedings of the National Academy of Sciences of the United States of America, 112(7):E606.

RevDate: 2018-11-13
CmpDate: 2014-12-08

Callaway E, Sutikna T, Roberts R, et al (2014)

The discovery of Homo floresiensis: Tales of the hobbit.

Nature, 514(7523):422-426.

RevDate: 2021-10-21
CmpDate: 2014-11-03

Henneberg M, Eckhardt RB, Chavanaves S, et al (2014)

Evolved developmental homeostasis disturbed in LB1 from Flores, Indonesia, denotes Down syndrome and not diagnostic traits of the invalid species Homo floresiensis.

Proceedings of the National Academy of Sciences of the United States of America, 111(33):11967-11972.

Human skeletons from Liang Bua Cave, Flores, Indonesia, are coeval with only Homo sapiens populations worldwide and no other previously known hominins. We report here for the first time to our knowledge the occipitofrontal circumference of specimen LB1. This datum makes it possible to link the 430-mL endocranial volume of LB1 reported by us previously, later confirmed independently by other investigators, not only with other human skeletal samples past and present but also with a large body of clinical data routinely collected on patients with developmental disorders. Our analyses show that the brain size of LB1 is in the range predicted for an individual with Down syndrome (DS) in a normal small-bodied population from the geographic region that includes Flores. Among additional diagnostic signs of DS and other skeletal dysplasiae are abnormally short femora combined with disproportionate flat feet. Liang Bua Cave femora, known only for LB1, match interlimb proportions for DS. Predictions based on corrected LB1 femur lengths show a stature normal for other H. sapiens populations in the region.

RevDate: 2021-10-21
CmpDate: 2014-11-03

Eckhardt RB, Henneberg M, Weller AS, et al (2014)

Rare events in earth history include the LB1 human skeleton from Flores, Indonesia, as a developmental singularity, not a unique taxon.

Proceedings of the National Academy of Sciences of the United States of America, 111(33):11961-11966.

The original centrally defining features of "Homo floresiensis" are based on bones represented only in the single specimen LB1. Initial published values of 380-mL endocranial volume and 1.06-m stature are markedly lower than later attempts to confirm them, and facial asymmetry originally unreported, then denied, has been established by our group and later confirmed independently. Of nearly 200 syndromes in which microcephaly is one sign, more than half include asymmetry as another sign and more than one-fourth also explicitly include short stature. The original diagnosis of the putative new species noted and dismissed just three developmental abnormalities. Subsequent independent attempts at diagnosis (Laron Syndrome, Majewski osteodysplastic primordial dwarfism type II, cretinism) have been hampered a priori by selectively restricted access to specimens, and disparaged a posteriori using data previously unpublished, without acknowledging that all of the independent diagnoses corroborate the patent abnormal singularity of LB1. In this report we establish in detail that even in the absence of a particular syndromic diagnosis, the originally defining features of LB1 do not establish either the uniqueness or normality necessary to meet the formal criteria for a type specimen of a new species. In a companion paper we present a new syndromic diagnosis for LB1.

RevDate: 2021-10-21
CmpDate: 2015-02-12

Parr WC, Soligo C, Smaers J, et al (2014)

Three-dimensional shape variation of talar surface morphology in hominoid primates.

Journal of anatomy, 225(1):42-59.

The hominoid foot is of particular interest to biological anthropologists, as changes in its anatomy through time reflect the adoption of terrestrial locomotion, particularly in species of Australopithecus and Homo. Understanding the osteological morphology associated with changes in whole foot function and the development of the plantar medial longitudinal foot arch are key to understanding the transition through habitual bipedalism in australopithecines to obligate bipedalism and long-distance running in Homo. The talus is ideal for studying relationships between morphology and function in this context, as it is a major contributor to the adduction-abduction, plantar-dorsal flexion and inversion-eversion of the foot, and transmits all forces encountered from the foot to the leg. The talar surface is predominantly covered by articular facets, which have different quantifiable morphological characters, including surface area, surface curvature and orientation. The talus also presents challenges to the investigator, as its globular shape is very difficult to quantify accurately and reproducibly. Here we apply a three-dimensional approach using type 3 landmarks (slid semilandmarks) that are geometrically homologous to determine overall talar shape variations in a range of living and fossil hominoid taxa. Additionally, we use novel approaches to quantify the relative orientations and curvatures of talar articular facets by determining the principal vectors of facet orientation and fitting spheres to articular facets. The resulting metrics are analysed using phylogenetic regressions and principal components analyses. Our results suggest that articular surface curvatures reflect locomotor specialisations with, in particular, orangutans having more highly curved facets in all but the calcaneal facet. Similarly, our approach to quantifying articular facet orientation appears to be effective in discriminating between extant hominoid species, and may therefore provide a sound basis for the study of fossil taxa and evolution of bipedalism in Australopithecus and Homo.

RevDate: 2021-10-21
CmpDate: 2015-01-06

Thomson V, Aplin KP, Cooper A, et al (2014)

Molecular genetic evidence for the place of origin of the Pacific rat, Rattus exulans.

PloS one, 9(3):e91356.

Commensal plants and animals have long been used to track human migrations, with Rattus exulans (the Pacific rat) a common organism for reconstructing Polynesian dispersal in the Pacific. However, with no knowledge of the homeland of R. exulans, the place of origin of this human-commensal relationship is unknown. We conducted a mitochondrial DNA phylogeographic survey of R. exulans diversity across the potential natural range in mainland and Island Southeast Asia in order to establish the origin of this human-commensal dyad. We also conducted allozyme electrophoresis on samples from ISEA to obtain a perspective on patterns of genetic diversity in this critical region. Finally, we compared molecular genetic evidence with knowledge of prehistoric rodent faunas in mainland and ISEA. We find that ISEA populations of R. exulans contain the highest mtDNA lineage diversity including significant haplotype diversity not represented elsewhere in the species range. Within ISEA, the island of Flores in the Lesser Sunda group contains the highest diversity in ISEA (across all loci) and also has a deep fossil record of small mammals that appears to include R. exulans. Therefore, in addition to Flores harboring unusual diversity in the form of Homo floresiensis, dwarfed stegodons and giant rats, this island appears to be the homeland of R. exulans.

RevDate: 2016-11-25
CmpDate: 2014-11-03

Daegling DJ, Patel BA, WL Jungers (2014)

Geometric properties and comparative biomechanics of Homo floresiensis mandibles.

Journal of human evolution, 68:36-46.

The hypodigm of Homo floresiensis from the cave of Liang Bua on Flores Island in the archipelago of Indonesia includes two mandibles (LB1/2 and LB6/1). The morphology of their symphyses and corpora has been described as sharing similarities with both australopiths and early Homo despite their Late Pleistocene age. Although detailed morphological comparisons of these mandibles with those of modern and fossil hominin taxa have been made, a functional analysis in the context of masticatory biomechanics has yet to be performed. Utilizing data on cortical bone geometry from computed tomography scans, we compare the mechanical attributes of the LB1 and LB6 mandibles with samples of modern Homo, Pan, Pongo, and Gorilla, as well as fossil samples of Paranthropus robustus, Australopithecus africanus and South African early Homo. Structural stiffness measures were derived from the geometric data to provide relative measures of mandibular corpus strength under hypothesized masticatory loading regimes. These mechanical variables were evaluated relative to bone area, mandibular length and estimates of body size to assess their functional affinities and to test the hypothesis that the Liang Bua mandibles can be described as scaled-down variants of either early hominins or modern humans. Relative to modern hominoids, the H. floresiensis material appears to be relatively strong in terms of rigidity in torsion and transverse bending, but is relatively weak under parasagittal bending. Thus, they are 'robust' relative to modern humans (and comparable with australopiths) under some loads but not others. Neither LB1 nor LB6 can be described simply as 'miniaturized' versions of modern human jaws since mandible length is more or less equivalent in Homo sapiens and H. floresiensis. The mechanical attributes of the Liang Bua mandibles are consistent with previous inferences that masticatory loads were reduced relative to australopiths but remained elevated relative to modern Homo.

RevDate: 2022-12-07
CmpDate: 2015-04-09

Higham C (2013)

Hunter-gatherers in southeast Asia: from prehistory to the present.

Human biology, 85(1-3):21-43.

Anatomically modern hunter-gatherers expanded from Africa into Southeast Asia at least 50,000 years ago, where they probably encountered and interacted with populations of Homo erectus and Homo floresiensis and the recently discovered Denisovans. Simulation studies suggest that these hunter-gatherers may well have followed a coastal route that ultimately led to the settlement of Sahul, while archaeology confirms that they also crossed significant seas and explored well into the interior. They also adapted to marked environmental changes that alternated between relatively cool and dry conditions and warmer, wetter interludes. During the former, the sea fell by up to 120 m below its present level, which opened up a vast low-lying area known as Sundaland. Three principal alignments can be identified: the first involved the occupation of rock shelters in upland regions, the second has identified settlement on broad riverine floodplains, and the last concentrated on the raised beaches formed from about five millennia ago when the sea level was elevated above its present position. This cultural sequence was dislocated about 4 kya when rice and millet farmers infiltrated the lowlands of Southeast Asia ultimately from the Yangtze River valley. It is suggested that this led to two forms of interaction. In the first, the indigenous hunter-gatherers integrated with intrusive Neolithic communities and, while losing their cultural identity, contributed their genes to the present population of Southeast Asia. In the second, hunter-gatherers withdrew to rainforest refugia and, through selective pressures inherent in such an environment, survived as the small-bodied, dark-skinned humans found to this day in the Philippines, Peninsular Malaysia and Thailand, and the Andaman Islands. Beyond the impact of expansive rice farmers in Melanesia and Australia, hunter-gatherers continued to dominate until they encountered European settlement.

RevDate: 2013-11-18
CmpDate: 2014-06-30

Montgomery SH (2013)

Primate brains, the 'island rule' and the evolution of Homo floresiensis.

Journal of human evolution, 65(6):750-760.

The taxonomic status of the small bodied hominin, Homo floresiensis, remains controversial. One contentious aspect of the debate concerns the small brain size estimated for specimen LB1 (Liang Bua 1). Based on intraspecific mammalian allometric relationships between brain and body size, it has been argued that the brain of LB1 is too small for its body mass and is therefore likely to be pathological. The relevance and general applicability of these scaling rules has, however, been challenged, and it is not known whether highly encephalized primates adapt to insular habitats in a consistent manner. Here, an analysis of brain and body size evolution in seven extant insular primates reveals that although insular primates follow the 'island rule', having consistently reduced body masses compared with their mainland relatives, neither brain mass nor relative brain size follow similar patterns, contrary to expectations that energetic constraints will favour decreased relative brain size. Brain:body scaling relationships previously used to assess the plausibility of dwarfism in H. floresiensis tend to underestimate body masses of insular primates. In contrast, under a number of phylogenetic scenarios, the evolution of brain and body mass in H. floresiensis is consistent with patterns observed in other insular primates.

RevDate: 2021-10-21
CmpDate: 2014-07-01

Haslam M (2013)

'Captivity bias' in animal tool use and its implications for the evolution of hominin technology.

Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 368(1630):20120421.

Animals in captive or laboratory settings may outperform wild animals of the same species in both frequency and diversity of tool use, a phenomenon here termed 'captivity bias'. Although speculative at this stage, a logical conclusion from this concept is that animals whose tool-use behaviour is observed solely under natural conditions may be judged cognitively or physically inferior than if they had also been tested or observed under controlled captive conditions. In turn, this situation creates a potential problem for studies of the behaviour of extinct members of the human family tree-the hominins-as hominin cognitive abilities are often judged on material evidence of tool-use behaviour left in the archaeological record. In this review, potential factors contributing to captivity bias in primates (including increased contact between individuals engaged in tool use, guidance or shaping of tool-use behaviour by other tool-users and increased free time and energy) are identified and assessed for their possible effects on the behaviour of the Late Pleistocene hominin Homo floresiensis. The captivity bias concept provides one way to uncouple hominin tool use from cognition, by considering hominins as subject to the same adaptive influences as other tool-using animals.

RevDate: 2021-10-21
CmpDate: 2014-03-31

Baab KL, McNulty KP, K Harvati (2013)

Homo floresiensis contextualized: a geometric morphometric comparative analysis of fossil and pathological human samples.

PloS one, 8(7):e69119.

The origin of hominins found on the remote Indonesian island of Flores remains highly contentious. These specimens may represent a new hominin species, Homo floresiensis, descended from a local population of Homo erectus or from an earlier (pre-H. erectus) migration of a small-bodied and small-brained hominin out of Africa. Alternatively, some workers suggest that some or all of the specimens recovered from Liang Bua are pathological members of a small-bodied modern human population. Pathological conditions proposed to explain their documented anatomical features include microcephaly, myxoedematous endemic hypothyroidism ("cretinism") and Laron syndrome (primary growth hormone insensitivity). This study evaluates evolutionary and pathological hypotheses through comparative analysis of cranial morphology. Geometric morphometric analyses of landmark data show that the sole Flores cranium (LB1) is clearly distinct from healthy modern humans and from those exhibiting hypothyroidism and Laron syndrome. Modern human microcephalic specimens converge, to some extent, on crania of extinct species of Homo. However in the features that distinguish these two groups, LB1 consistently groups with fossil hominins and is most similar to H. erectus. Our study provides further support for recognizing the Flores hominins as a distinct species, H. floresiensis, whose affinities lie with archaic Homo.

RevDate: 2013-07-08
CmpDate: 2014-06-07

Goulden M (2013)

Hobbits, hunters and hydrology: Images of a "missing link," and its scientific communication.

Public understanding of science (Bristol, England), 22(5):575-589.

Dissemination of the 2004 discovery of Homo floresiensis, aka the "Flores Hobbit," provides the case material for an analysis of science communication models, using coverage of the find in science journals, popular science magazines, and UK newspapers. A distinction is made between "popular science" and "public science," the latter notable for the proactive manner in which it seeks not simply to interpret science for a wider audience, but also expand upon it. The nature of the flows of knowledge between science and nonscience cultures is used to critique both traditional canonical models of science communication, and more recent constructivist accounts like that of Hilgartner's "continuum" model. The paper concludes by suggesting an extended model that builds on the logic of the river metaphor used by both Hilgartner and Gieryn. It is argued that this more complex, multi-directional, "hydrologic" model provides a richer account of science communication.

RevDate: 2013-08-26
CmpDate: 2013-11-13

van Heteren AH (2013)

Is LB1 diseased or modern? A review of the proposed pathologies.

Gene, 528(1):12-20.

The fossil remains of Homo floresiensis have been debated extensively over the past few years. This paper gives a review of the various pathologies ascribed to LB1, the type specimen of H. floresiensis, and associated individuals. This paper will assess the arguments for growth anomalies, microcephaly, Laron syndrome and cretinism. Additionally, some of the analyses done by proponents of the pathology theory will be methodologically evaluated. Subsequently, a brief overview of the alternative hypotheses regarding the origin of H. floresiensis will be given.

RevDate: 2021-10-21
CmpDate: 2013-09-30

Kubo D, Kono RT, Y Kaifu (2013)

Brain size of Homo floresiensis and its evolutionary implications.

Proceedings. Biological sciences, 280(1760):20130338.

The extremely small endocranial volume (ECV) of LB1, the type specimen of Homo floresiensis, poses a challenge in our understanding of human brain evolution. Some researchers hypothesize dramatic dwarfing of relative brain size from Homo erectus presumably without significant decrease in intellectual function, whereas others expect a lesser degree of brain diminution from a more primitive, small-brained form of hominin currently undocumented in eastern Asia. However, inconsistency in the published ECVs for LB1 (380-430 cc), unclear human intraspecific brain-body size scaling and other uncertainties have hampered elaborative modelling of its brain size reduction. In this study, we accurately determine the ECV of LB1 using high-resolution micro-CT scan. The ECV of LB1 thus measured, 426 cc, is larger than the commonly cited figure in previous studies (400 cc). Coupled with brain-body size correlation in Homo sapiens calculated based on a sample from 20 worldwide modern human populations, we construct new models of the brain size reduction in the evolution of H. floresiensis. The results show a more significant contribution of scaling effect than previously claimed.

RevDate: 2013-04-09
CmpDate: 2013-09-30

Montgomery SH, NI Mundy (2013)

Parallel episodes of phyletic dwarfism in callitrichid and cheirogaleid primates.

Journal of evolutionary biology, 26(4):810-819.

The Callitrichidae are the smallest anthropoids, whereas the Cheirogaleidae include the smallest of all primates. Using species-level analyses, we show that these are derived conditions; both neonatal and adult body mass decreased in a gradual, phyletic manner in parallel across callitrichids, and across cheirogaleids. We identify lineages with particularly rapid decreases and highlight the pygmy marmoset, Callithrix pygmaea, as a phenotypic outlier. The life-history traits associated with body-mass reduction in each clade suggest that the convergent evolution of small body size was achieved by changes in different ontogenetic stages. Body-size reduction in callitrichids appears to be almost exclusively due to alterations in prenatal growth rate, whereas body-size reduction in cheirogaleids may have been largely due to reduced duration of growth phases. Finally, we use these results to discuss some of the debates surrounding the evolution of Homo floresiensis and suggest potential parallels between the evolution of H. floresiensis and callitrichids.

RevDate: 2013-03-18
CmpDate: 2013-08-30

Vannucci RC, Barron TF, RL Holloway (2013)

Frontal brain expansion during development using MRI and endocasts: relation to microcephaly and Homo floresiensis.

Anatomical record (Hoboken, N.J. : 2007), 296(4):630-637.

A major hall of hominid brain evolution is an expansion of the frontal lobes. To determine if a similar trajectory occurs during modern human development, the MRI scans of 118 living infants, children, and adolescents were reviewed and three specific measurements obtained: frontal width (FW), maximal cerebral width (MW), and maximal cerebral length (ML). The infantile brain is uniformly wide but relatively short, with near equal FW and MW. The juvenile brain exhibits a wider MW than FW, while FW of the adolescent brain expands to nearly equal MW, concurrent with an increase in ML. The preferential frontal lobe expansion during modern human development parallels that observed during the evolution of Homo. In 17 microcephalic individuals, only 6 (35%) exhibited preferential frontal lobe hypoplasia, presumably a reflection of multiple etiologies that adversely affect differing brain regions. Compared to 79 modern human adult endocasts and 12 modern microcephalic endocasts, LB1 (Homo floresiensis) clustered more consistently with the microcephalic sample than with the normocephalic sample.

RevDate: 2013-02-04
CmpDate: 2013-07-08

Orr CM, Tocheri MW, Burnett SE, et al (2013)

New wrist bones of Homo floresiensis from Liang Bua (Flores, Indonesia).

Journal of human evolution, 64(2):109-129.

The carpals from the Homo floresiensis type specimen (LB1) lack features that compose the shared, derived complex of the radial side of the wrist in Neandertals and modern humans. This paper comprises a description and three-dimensional morphometric analysis of new carpals from at least one other individual at Liang Bua attributed to H. floresiensis: a right capitate and two hamates. The new capitate is smaller than that of LB1 but is nearly identical in morphology. As with capitates from extant apes, species of Australopithecus, and LB1, the newly described capitate displays a deeply-excavated nonarticular area along its radial aspect, a scaphoid facet that extends into a J-hook articulation on the neck, and a more radially-oriented second metacarpal facet; it also lacks an enlarged palmarly-positioned trapezoid facet. Because there is no accommodation for the derived, palmarly blocky trapezoid that characterizes Homo sapiens and Neandertals, this individual most likely had a plesiomorphically wedge-shaped trapezoid (like LB1). Morphometric analyses confirm the close similarity of the new capitate and that of LB1, and are consistent with previous findings of an overall primitive articular geometry. In general, hamate morphology is more conserved across hominins, and the H. floresiensis specimens fall at the far edge of the range of variation for H. sapiens in a number of metrics. However, the hamate of H. floresiensis is exceptionally small and exhibits a relatively long, stout hamulus lacking the oval-shaped cross-section characteristic of human and Neandertal hamuli (variably present in australopiths). Documentation of a second individual with primitive carpal anatomy from Liang Bua, along with further analysis of trapezoid scaling relative to the capitate in LB1, refutes claims that the wrist of the type specimen represents a modern human with pathology. In total, the carpal anatomy of H. floresiensis supports the hypothesis that the lineage leading to the evolution of this species originated prior to the cladogenetic event that gave rise to modern humans and Neandertals.

RevDate: 2012-11-19
CmpDate: 2013-07-05

Oxnard C, Obendorf PJ, Kefford BJ, et al (2012)

More on the Liang Bua finds and modern human cretins.

Homo : internationale Zeitschrift fur die vergleichende Forschung am Menschen, 63(6):407-412.

Brown (2012: LB1 and LB6 Homo floresiensis are not modern human (Homo sapiens) cretins, Journal of Human Evolution) makes errors of fact, omission and interpretation. Brown's comments refer, among others, to (1) delayed growth and development indicated by unfused epiphyses, (2) postcranial limb proportions: limbs to trunk, between limbs, and within limbs, (3) postcranial bone torsions and angles, (4) postcranial robusticity, real and apparent, (5) skull features, and (6) cretinism on Flores. In each of these areas, much information about cretins is incorrect and much information (Oxnard et al., 2010) comparing the Liang Bua remains with cretins is ignored.

RevDate: 2012-09-05
CmpDate: 2013-01-14

Bokma F, van den Brink V, T Stadler (2012)

Unexpectedly many extinct hominins.

Evolution; international journal of organic evolution, 66(9):2969-2974.

Recent studies indicate that Neanderthal and Denisova hominins may have been separate species, while debate continues on the status of Homo floresiensis. The decade-long debate between "splitters," who recognize over 20 hominin species, and "lumpers," who maintain that all these fossils belong to just a few lineages, illustrates that we do not know how many extinct hominin species to expect. Here, we present probability distributions for the number of speciation events and the number of contemporary species along a branch of a phylogeny. With estimates of hominin speciation and extincton rates, we then show that the expected total number of extinct hominin species is 8, but may be as high as 27. We also show that it is highly unlikely that three very recent species disappeared due to natural, background extinction. This may indicate that human-like remains are too easily considered distinct species. Otherwise, the evidence suggesting that Neanderthal and the Denisova hominin represent distinct species implies a recent wave of extinctions, ostensibly driven by the only survivor, H. sapiens.

RevDate: 2012-02-06
CmpDate: 2012-05-31

Brown P (2012)

LB1 and LB6 Homo floresiensis are not modern human (Homo sapiens) cretins.

Journal of human evolution, 62(2):201-224.

Excavations in the late Pleistocene deposits at Liang Bua cave, Flores, have uncovered the skeletal remains of several small-bodied and small-brained hominins in association with stone artefacts and the bones of Stegodon. Due to their combination of plesiomorphic, unique and derived traits, they were ascribed to a new species, Homo floresiensis, which, along with Stegodon, appears to have become extinct ∼17 ka (thousand years ago). However, recently it has been argued that several characteristics of H. floresiensis were consistent with dwarfism and evidence of delayed development in modern human (Homo sapiens) myxoedematous endemic (ME) cretins. This research compares the skeletal and dental morphology in H. floresiensis with the clinical and osteological indicators of cretinism, and the traits that have been argued to be associated with ME cretinism in LB1 and LB6. Contrary to published claims, morphological and statistical comparisons did not identify the distinctive skeletal and dental indicators of cretinism in LB1 or LB6 H. floresiensis. Brain mass, skeletal proportions, epiphyseal union, orofacial morphology, dental development, size of the pituitary fossa and development of the paranasal sinuses, vault bone thickness and dimensions of the hands and feet all distinguish H. floresiensis from modern humans with ME cretinism. The research team responsible for the diagnosis of ME cretinism had not examined the original H. floresiensis skeletal materials, and perhaps, as a result, their research confused taphonomic damage with evidence of disease, and thus contained critical errors of fact and interpretation. Behavioural scenarios attempting to explain the presence of cretinous H. sapiens in the Liang Bua Pleistocene deposits, but not unaffected H. sapiens, are both unnecessary and not supported by the available archaeological and geochronological evidence from Flores.

RevDate: 2011-11-08
CmpDate: 2012-03-02

Kaifu Y, Baba H, Sutikna T, et al (2011)

Craniofacial morphology of Homo floresiensis: description, taxonomic affinities, and evolutionary implication.

Journal of human evolution, 61(6):644-682.

This paper describes in detail the external morphology of LB1/1, the nearly complete and only known cranium of Homo floresiensis. Comparisons were made with a large sample of early groups of the genus Homo to assess primitive, derived, and unique craniofacial traits of LB1 and discuss its evolution. Principal cranial shape differences between H. floresiensis and Homo sapiens are also explored metrically. The LB1 specimen exhibits a marked reductive trend in its facial skeleton, which is comparable to the H. sapiens condition and is probably associated with reduced masticatory stresses. However, LB1 is craniometrically different from H. sapiens showing an extremely small overall cranial size, and the combination of a primitive low and anteriorly narrow vault shape, a relatively prognathic face, a rounded oval foramen that is greatly separated anteriorly from the carotid canal/jugular foramen, and a unique, tall orbital shape. Whereas the neurocranium of LB1 is as small as that of some Homo habilis specimens, it exhibits laterally expanded parietals, a weak suprameatal crest, a moderately flexed occipital, a marked facial reduction, and many other derived features that characterize post-habilis Homo. Other craniofacial characteristics of LB1 include, for example, a relatively narrow frontal squama with flattened right and left sides, a marked frontal keel, posteriorly divergent temporal lines, a posteriorly flexed anteromedial corner of the mandibular fossa, a bulbous lateral end of the supraorbital torus, and a forward protruding maxillary body with a distinct infraorbital sulcus. LB1 is most similar to early Javanese Homo erectus from Sangiran and Trinil in these and other aspects. We conclude that the craniofacial morphology of LB1 is consistent with the hypothesis that H. floresiensis evolved from early Javanese H. erectus with dramatic island dwarfism. However, further field discoveries of early hominin skeletal remains from Flores and detailed analyses of the finds are needed to understand the evolutionary history of this endemic hominin species.

RevDate: 2021-10-20
CmpDate: 2011-10-27

Vannucci RC, Barron TF, RL Holloway (2011)

Craniometric ratios of microcephaly and LB1, Homo floresiensis, using MRI and endocasts.

Proceedings of the National Academy of Sciences of the United States of America, 108(34):14043-14048.

The designation of Homo floresiensis as a new species derived from an ancient population is controversial, because the type specimen, LB1, might represent a pathological microcephalic modern Homo sapiens. Accordingly, two specific craniometric ratios (relative frontal breadth and cerebellar protrusion) were ascertained in 21 microcephalic infants and children by using MRI. Data on 118 age-equivalent control (normocephalic) subjects were collected for comparative purposes. In addition, the same craniometric ratios were determined on the endocasts of 10 microcephalic individuals, 79 normal controls (anatomically modern humans), and 17 Homo erectus specimens. These ratios were then compared with those of two LB1 endocasts. The findings showed that the calculated cerebral/cerebellar ratios of the LB1 endocast [Falk D, et al. (2007) Proc Natl Acad Sci USA 104:2513-2518] fall outside the range of living normocephalic individuals. The ratios derived from two LB1 endocasts also fall largely outside the range of modern normal human and H. erectus endocasts and within the range of microcephalic endocasts. The findings support but do not prove the contention that LB1 represents a pathological microcephalic Homo sapiens rather than a new species, (i.e., H. floresiensis).

RevDate: 2016-11-25
CmpDate: 2011-09-13

Jungers WL, Y Kaifu (2011)

On dental wear, dental work, and oral health in the type specimen (LB1) of Homo floresiensis.

American journal of physical anthropology, 145(2):282-289.

The claim that the lower left first mandibular molar of LB1, the type specimen of Homo floresiensis, displays endodontic work, and a filling is assessed by digital radiography and micro-CT scanning. The M(1) tooth crown is heavily worn and exhibits extensive dentine exposure that is stained white, but there is no trace of endodontic treatment or a dental filling in this Indonesian fossil dated to 17.1-19.0 kya. Dental calculus (commonly observed in foragers) is present on the teeth of LB1, but there are no observable caries. The pattern of dental attrition in the mandibles of both LB1/2 and LB6/1 (moderate to extensive flat wear across the entire arch) is consistent with that seen in Plio-Pleistocene Homo fossils and in modern hunter-gatherers, and is not typical of most agriculturalists. We conclude that the dental-work and farming hypotheses are falsified and therefore irrelevant to the debate over the taxonomy and phylogeny of H. floresiensis.

RevDate: 2021-10-20
CmpDate: 2011-02-18

Oxnard C, Obendorf PJ, BJ Kefford (2010)

Post-cranial skeletons of hypothyroid cretins show a similar anatomical mosaic as Homo floresiensis.

PloS one, 5(9):e13018.

Human remains, some as recent as 15 thousand years, from Liang Bua (LB) on the Indonesian island of Flores have been attributed to a new species, Homo floresiensis. The definition includes a mosaic of features, some like modern humans (hence derived: genus Homo), some like modern apes and australopithecines (hence primitive: not species sapiens), and some unique (hence new species: floresiensis). Conversely, because only modern humans (H. sapiens) are known in this region in the last 40 thousand years, these individuals have also been suggested to be genetic human dwarfs. Such dwarfs resemble small humans and do not show the mosaic combination of the most complete individuals, LB1 and LB6, so this idea has been largely dismissed. We have previously shown that some features of the cranium of hypothyroid cretins are like those of LB1. Here we examine cretin postcrania to see if they show anatomical mosaics like H. floresiensis. We find that hypothyroid cretins share at least 10 postcranial features with Homo floresiensis and unaffected humans not found in apes (or australopithecines when materials permit). They share with H. floresiensis, modern apes and australopithecines at least 11 postcranial features not found in unaffected humans. They share with H. floresiensis, at least 8 features not found in apes, australopithecines or unaffected humans. Sixteen features can be rendered metrically and multivariate analyses demonstrate that H. floresiensis co-locates with cretins, both being markedly separate from humans and chimpanzees (P<0.001: from analysis of similarity (ANOSIM) over all variables, ANOSIM, global R>0.999). We therefore conclude that LB1 and LB6, at least, are, most likely, endemic cretins from a population of unaffected Homo sapiens. This is consistent with recent hypothyroid endemic cretinism throughout Indonesia, including the nearby island of Bali.

RevDate: 2016-10-21
CmpDate: 2010-11-09

Alba DM (2010)

Cognitive inferences in fossil apes (Primates, Hominoidea): does encephalization reflect intelligence?.

Journal of anthropological sciences = Rivista di antropologia : JASS, 88:11-48.

Paleobiological inferences on general cognitive abilities (intelligence) in fossil hominoids strongly rely on relative brain size or encephalization, computed by means of allometric residuals, quotients or constants. Th is has been criticized on the basis that it presumably fails to reflect the higher intelligence of great apes, and absolute brain size has been favored instead. Many problems of encephalization metrics stem from the decrease of allometric slopes towards lower taxonomic level, thus making it difficult to determine at what level encephalization metrics have biological meaning. Here, the hypothesis that encephalization can be used as a good neuroanatomical proxy for intelligence is tested at two different taxonomic levels. A significant correlation is found between intelligence and encephalization only at a lower taxonomic level, i.e. on the basis of a low allometric slope, irrespective of whether species data or independent contrasts are employed. This indicates that higher-level slopes, resulting from encephalization grade shifts between subgroups (including hylobatids vs. great apes), do not reflect functional equivalence, whereas lower-level metrics can be employed as a paleobiological proxy for intelligence. Thus, in accordance to intelligence rankings, lower-level metrics indicate that great apes are more encephalized than both monkeys and hylobatids. Regarding fossil taxa, encephalization increased during hominin evolution (particularly in Homo), but during the Miocene a significant shift towards higher encephalization (and inferred enhanced cognitive abilities) must have been also involved in the emergence of the great-ape-and-human clade (Hominidae). This is confirmed by the modern great-ape-like degree of encephalization displayed by the fossil great ape Hispanopithecus, which contrasts with the rather hylobatid-like degree of the stem hominoid Proconsul. The similarly low encephalization of Oreopithecus might result from secondary reduction under insularity conditions, but the australopith-like degree of encephalization of Homo floresiensis seems incompatible with the cognitive abilities inferred from the stone tools attributed to this taxon.

RevDate: 2023-03-23
CmpDate: 2023-03-23

Mijares AS, Détroit F, Piper P, et al (2010)

New evidence for a 67,000-year-old human presence at Callao Cave, Luzon, Philippines.

Journal of human evolution, 59(1):123-132.

Documentation of early human migrations through Island Southeast Asia and Wallacea en route to Australia has always been problematic due to a lack of well-dated human skeletal remains. The best known modern humans are from Niah Cave in Borneo (40-42ka), and from Tabon Cave on the island of Palawan, southwest Philippines (47+/-11ka). The discovery of Homo floresiensis on the island of Flores in eastern Indonesia has also highlighted the possibilities of identifying new hominin species on islands in the region. Here, we report the discovery of a human third metatarsal from Callao Cave in northern Luzon. Direct dating of the specimen using U-series ablation has provided a minimum age estimate of 66.7+/-1ka, making it the oldest known human fossil in the Philippines. Its morphological features, as well as size and shape characteristics, indicate that the Callao metatarsal definitely belongs to the genus Homo. Morphometric analysis of the Callao metatarsal indicates that it has a gracile structure, close to that observed in other small-bodied Homo sapiens. Interestingly, the Callao metatarsal also falls within the morphological and size ranges of Homo habilis and H. floresiensis. Identifying whether the metatarsal represents the earliest record of H. sapiens so far recorded anywhere east of Wallace's Line requires further archaeological research, but its presence on the isolated island of Luzon over 65,000 years ago further demonstrates the abilities of humans to make open ocean crossings in the Late Pleistocene.

RevDate: 2010-08-23
CmpDate: 2010-12-14

Trueman JW (2010)

A new cladistic analysis of Homo floresiensis.

Journal of human evolution, 59(2):223-6; discussion 227-30.

RevDate: 2010-05-25
CmpDate: 2010-09-15

Aiello LC (2010)

Five years of Homo floresiensis.

American journal of physical anthropology, 142(2):167-179.

Since Homo floresiensis was first described in October 2004 there has been a lively debate over its status. Is it a late surviving species of early Homo or merely a modern individual afflicted with disordered growth and one of the many syndromes resulting in microchephaly? Recently the discovery team has published a series of articles providing detailed descriptions of the hominin material, its geomorphological context, and the associated archaeology and faunal material (Morwood and Jungers: J Hum Evol 57 (2009) 437-648). In addition, other researchers have put forward new hypotheses for possible pathologies including Laron's Syndrome and Myxoedematous Endemic (ME) Cretinism. Here I review this new information and conclude that the evidence supports the hypothesis that Homo floresiensis is a late-surviving species of early Homo with its closest morphological affinities to early African pre-erectus/ergaster hominins. Although this hypothesis requires fundamental paradigm changes in our understanding of human evolution, it provides a more economical explanation for H. floresiensis than do the alternatives. None of the current explanations for microcephaly and disordered growth account for the range of features observed in H. floresiensis. Neither do they provide explanations for why a pathological condition in modern humans would mimic so closely the morphology observed in earlier hominins. This conclusion is based on the current evidence for H. floresiensis and on the particular pathological explanations that have appeared in the literature. There is no doubt that controversy over H. floresiensis will continue until new and conclusive evidence is available to settle the debate one way or another.

RevDate: 2021-10-20
CmpDate: 2010-05-05

Montgomery SH, Capellini I, Barton RA, et al (2010)

Reconstructing the ups and downs of primate brain evolution: implications for adaptive hypotheses and Homo floresiensis.

BMC biology, 8:9.

BACKGROUND: Brain size is a key adaptive trait. It is often assumed that increasing brain size was a general evolutionary trend in primates, yet recent fossil discoveries have documented brain size decreases in some lineages, raising the question of how general a trend there was for brains to increase in mass over evolutionary time. We present the first systematic phylogenetic analysis designed to answer this question.

RESULTS: We performed ancestral state reconstructions of three traits (absolute brain mass, absolute body mass, relative brain mass) using 37 extant and 23 extinct primate species and three approaches to ancestral state reconstruction: parsimony, maximum likelihood and Bayesian Markov-chain Monte Carlo. Both absolute and relative brain mass generally increased over evolutionary time, but body mass did not. Nevertheless both absolute and relative brain mass decreased along several branches. Applying these results to the contentious case of Homo floresiensis, we find a number of scenarios under which the proposed evolution of Homo floresiensis' small brain appears to be consistent with patterns observed along other lineages, dependent on body mass and phylogenetic position.

CONCLUSIONS: Our results confirm that brain expansion began early in primate evolution and show that increases occurred in all major clades. Only in terms of an increase in absolute mass does the human lineage appear particularly striking, with both the rate of proportional change in mass and relative brain size having episodes of greater expansion elsewhere on the primate phylogeny. However, decreases in brain mass also occurred along branches in all major clades, and we conclude that, while selection has acted to enlarge primate brains, in some lineages this trend has been reversed. Further analyses of the phylogenetic position of Homo floresiensis and better body mass estimates are required to confirm the plausibility of the evolution of its small brain mass. We find that for our dataset the Bayesian analysis for ancestral state reconstruction is least affected by inclusion of fossil data suggesting that this approach might be preferable for future studies on other taxa with a poor fossil record.

RevDate: 2010-11-18
CmpDate: 2010-01-27

Morwood MJ, WL Jungers (2009)

Conclusions: implications of the Liang Bua excavations for hominin evolution and biogeography.

Journal of human evolution, 57(5):640-648.

Excavations at Liang Bua, on the Indonesian island of Flores, have yielded a stratified sequence of stone artifacts and faunal remains spanning the last 95k.yr., which includes the skeletal remains of two human species, Homo sapiens in the Holocene and Homo floresiensis in the Pleistocene. This paper summarizes and focuses on some of the evidence for Homo floresiensis in context, as presented in this Special Issue edition of the Journal of Human Evolution and elsewhere. Attempts to dismiss the Pleistocene hominins (and the type specimen LB1 in particular) as pathological pygmy humans are not compatible with detailed analyses of the skull, teeth, brain endocast, and postcranium. We initially concluded that H. floresiensis may have evolved by insular dwarfing of a larger-bodied hominin species over 880k.yr. or more. However, recovery of additional specimens and the numerous primitive morphological traits seen throughout the skeleton suggest instead that it is more likely to be a late representative of a small-bodied lineage that exited Africa before the emergence of Homo erectus sensu lato. Homo floresiensis is clearly not an australopithecine, but does retain many aspects of anatomy (and perhaps behavior) that are probably plesiomorphic for the genus Homo. We also discuss some of the other implications of this tiny, endemic species for early hominin dispersal and evolution (e.g., for the "Out of Africa 1" paradigm and more specifically for colonizing Southeast Asia), and we present options for future research in the region.

RevDate: 2021-10-20
CmpDate: 2010-02-01

Hocknull SA, Piper PJ, van den Bergh GD, et al (2009)

Dragon's paradise lost: palaeobiogeography, evolution and extinction of the largest-ever terrestrial lizards (Varanidae).

PloS one, 4(9):e7241.

BACKGROUND: The largest living lizard species, Varanus komodoensis Ouwens 1912, is vulnerable to extinction, being restricted to a few isolated islands in eastern Indonesia, between Java and Australia, where it is the dominant terrestrial carnivore. Understanding how large-bodied varanids responded to past environmental change underpins long-term management of V. komodoensis populations.

We reconstruct the palaeobiogeography of Neogene giant varanids and identify a new (unnamed) species from the island of Timor. Our data reject the long-held perception that V. komodoensis became a giant because of insular evolution or as a specialist hunter of pygmy Stegodon. Phyletic giantism, coupled with a westward dispersal from mainland Australia, provides the most parsimonious explanation for the palaeodistribution of V. komodoensis and the newly identified species of giant varanid from Timor. Pliocene giant varanid fossils from Australia are morphologically referable to V. komodoensis suggesting an ultimate origin for V. komodoensis on mainland Australia (>3.8 million years ago). Varanus komodoensis body size has remained stable over the last 900,000 years (ka) on Flores, a time marked by major faunal turnovers, extinction of the island's megafauna, the arrival of early hominids by 880 ka, co-existence with Homo floresiensis, and the arrival of modern humans by 10 ka. Within the last 2000 years their populations have contracted severely.

CONCLUSIONS/SIGNIFICANCE: Giant varanids were once a ubiquitous part of Subcontinental Eurasian and Australasian faunas during the Neogene. Extinction played a pivotal role in the reduction of their ranges and diversity throughout the late Quaternary, leaving only V. komodoensis as an isolated long-term survivor. The events over the last two millennia now threaten its future survival.

RevDate: 2010-11-18
CmpDate: 2010-01-27

Morwood MJ, Sutikna T, Saptomo EW, et al (2009)

Preface: research at Liang Bua, Flores, Indonesia.

Journal of human evolution, 57(5):437-449.

Excavations at Liang Bua, Flores, Indonesia, have yielded evidence for an endemic human species, Homo floresiensis, a population that occupied the cave between approximately 95-17ka. This discovery has major implications for early hominin evolution and dispersal in Africa and Asia, attracting worldwide interest. This preface describes the rationale for the excavations in historical, geographical, and wider research contexts, as well as the methods used. It also introduces the other papers on aspects of Liang Bua research that feature in this edition of the Journal of Human Evolution.

RevDate: 2009-09-08
CmpDate: 2009-12-08

Holliday TW, RG Franciscus (2009)

Body size and its consequences: allometry and the lower limb length of Liang Bua 1 (Homo floresiensis).

Journal of human evolution, 57(3):223-228.

Bivariate femoral length allometry in recent humans, Pan, and Gorilla is investigated with special reference to the diminutive Liang Bua (LB) 1 specimen (the holotype of Homo floresiensis) and six early Pleistocene femora referred to the genus Homo. Relative to predicted body mass, Pan and Gorilla femora show strong negative length allometry while recent human femora evince isometry to positive allometry, depending on sample composition and line-fitting technique employed. The allometric trajectories of Pan and Homo show convergence near the small body size range of LB 1, such that LB 1 manifests a low percentage deviation (d(yx) of Smith [1980]) from the Pan allometric trajectory and falls well within the 95% confidence limits around the Pan individuals (but also outside the 95% confidence limits for recent Homo). In contrast, the six early Pleistocene Homo femora, belonging to larger individuals, show much greater d(yx) values from both Pan and Gorilla and fall well above the 95% confidence limits for these taxa. All but one of these Pleistocene Homo specimens falls within the 95% confidence limits of the recent human sample. Similar results are obtained when femoral length is regressed on femoral head diameter in unlogged bivariate space. Regardless of the ultimate taxonomic status of LB 1, these findings are consistent with a prediction made by us (Franciscus and Holliday, 1992) that hominins in the small body size range of A.L. 288-1 ("Lucy"), including members of the genus Homo, will tend to possess short, ape-like lower limbs as a function of body size scaling.

RevDate: 2010-08-31
CmpDate: 2010-01-27

Argue D, Morwood MJ, Sutikna T, et al (2009)

Homo floresiensis: a cladistic analysis.

Journal of human evolution, 57(5):623-639.

The announcement of a new species, Homo floresiensis, a primitive hominin that survived until relatively recent times is an enormous challenge to paradigms of human evolution. Until this announcement, the dominant paradigm stipulated that: 1) only more derived hominins had emerged from Africa, and 2) H. sapiens was the only hominin since the demise of Homo erectus and Homo neanderthalensis. Resistance to H. floresiensis has been intense, and debate centers on two sets of competing hypotheses: 1) that it is a primitive hominin, and 2) that it is a modern human, either a pygmoid form or a pathological individual. Despite a range of analytical techniques having been applied to the question, no resolution has been reached. Here, we use cladistic analysis, a tool that has not, until now, been applied to the problem, to establish the phylogenetic position of the species. Our results produce two equally parsimonious phylogenetic trees. The first suggests that H. floresiensis is an early hominin that emerged after Homo rudolfensis (1.86Ma) but before H. habilis (1.66Ma, or after 1.9Ma if the earlier chronology for H. habilis is retained). The second tree indicates H. floresiensis branched after Homo habilis.

RevDate: 2009-11-16
CmpDate: 2010-01-27

Brown P, T Maeda (2009)

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species.

Journal of human evolution, 57(5):571-596.

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.

RevDate: 2021-10-28
CmpDate: 2009-05-18

Weston EM, AM Lister (2009)

Insular dwarfism in hippos and a model for brain size reduction in Homo floresiensis.

Nature, 459(7243):85-88.

Body size reduction in mammals is usually associated with only moderate brain size reduction, because the brain and sensory organs complete their growth before the rest of the body during ontogeny. On this basis, 'phyletic dwarfs' are predicted to have a greater relative brain size than 'phyletic giants'. However, this trend has been questioned in the special case of dwarfism of mammals on islands. Here we show that the endocranial capacities of extinct dwarf species of hippopotamus from Madagascar are up to 30% smaller than those of a mainland African ancestor scaled to equivalent body mass. These results show that brain size reduction is much greater than predicted from an intraspecific 'late ontogenetic' model of dwarfism in which brain size scales to body size with an exponent of 0.35. The nature of the proportional change or grade shift observed here indicates that selective pressures on brain size are potentially independent of those on body size. This study demonstrates empirically that it is mechanistically possible for dwarf mammals on islands to evolve significantly smaller brains than would be predicted from a model of dwarfing based on the intraspecific scaling of the mainland ancestor. Our findings challenge current understanding of brain-body allometric relationships in mammals and suggest that the process of dwarfism could in principle explain small brain size, a factor relevant to the interpretation of the small-brained hominin found on the Island of Flores, Indonesia.

RevDate: 2021-10-20
CmpDate: 2009-05-18

Jungers WL, Harcourt-Smith WE, Wunderlich RE, et al (2009)

The foot of Homo floresiensis.

Nature, 459(7243):81-84.

Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

RevDate: 2021-10-20
CmpDate: 2009-05-18

Lieberman DE (2009)

Palaeoanthropology: Homo floresiensis from head to toe.

Nature, 459(7243):41-42.

RevDate: 2022-12-07
CmpDate: 2009-09-02

Wall JD, Lohmueller KE, V Plagnol (2009)

Detecting ancient admixture and estimating demographic parameters in multiple human populations.

Molecular biology and evolution, 26(8):1823-1827.

We analyze patterns of genetic variation in extant human polymorphism data from the National Institute of Environmental Health Sciences single nucleotide polymorphism project to estimate human demographic parameters. We update our previous work by considering a larger data set (more genes and more populations) and by explicitly estimating the amount of putative admixture between modern humans and archaic human groups (e.g., Neandertals, Homo erectus, and Homo floresiensis). We find evidence for this ancient admixture in European, East Asian, and West African samples, suggesting that admixture between diverged hominin groups may be a general feature of recent human evolution.

RevDate: 2010-11-18
CmpDate: 2010-01-27

Moore MW, Sutikna T, Jatmiko , et al (2009)

Continuities in stone flaking technology at Liang Bua, Flores, Indonesia.

Journal of human evolution, 57(5):503-526.

This study examines trends in stone tool reduction technology at Liang Bua, Flores, Indonesia, where excavations have revealed a stratified artifact sequence spanning 95k.yr. The reduction sequence practiced throughout the Pleistocene was straightforward and unchanging. Large flakes were produced off-site and carried into the cave where they were reduced centripetally and bifacially by four techniques: freehand, burination, truncation, and bipolar. The locus of technological complexity at Liang Bua was not in knapping products, but in the way techniques were integrated. This reduction sequence persisted across the Pleistocene/Holocene boundary with a minor shift favoring unifacial flaking after 11ka. Other stone-related changes occurred at the same time, including the first appearance of edge-glossed flakes, a change in raw material selection, and more frequent fire-induced damage to stone artifacts. Later in the Holocene, technological complexity was generated by "adding-on" rectangular-sectioned stone adzes to the reduction sequence. The Pleistocene pattern is directly associated with Homo floresiensis skeletal remains and the Holocene changes correlate with the appearance of Homo sapiens. The one reduction sequence continues across this hominin replacement.

RevDate: 2010-10-20
CmpDate: 2009-11-17

Kaifu Y, Baba H, Kurniawan I, et al (2009)

Brief communication: "Pathological" deformation in the skull of LB1, the type specimen of Homo floresiensis.

American journal of physical anthropology, 140(1):177-185.

If the holotype of Homo floresiensis, LB1, suffered from a severe developmental pathology, this could undermine its status as the holotype of a new species. One of the proposed pathological indicators that still remains untested is asymmetric distortion in the skull of LB1 (Jacob et al.: Proc Natl Acad Sci USA 103 (2006) 13421-13426). Here, we present evidence that LB1 exhibits antemortem craniofacial deformities that are consistent with posterior deformational (positional) plagiocephaly. This is a relatively common condition in modern people with no serious associated health problems and does not represent a severe developmental abnormality in LB1.

RevDate: 2010-11-18
CmpDate: 2010-01-27

Westaway KE, Roberts RG, Sutikna T, et al (2009)

The evolving landscape and climate of western Flores: an environmental context for the archaeological site of Liang Bua.

Journal of human evolution, 57(5):450-464.

The rapidly changing landscape of the eastern Indonesian archipelago has evolved at a pace dictated by its tropical climate and its geological and tectonic history. This has produced accelerated karstification, flights of alluvial terraces, and complex, multi-level cave systems. These cave systems sometimes contain a wealth of archaeological evidence, such as the almost complete skeleton of Homo floresiensis found at the site of Liang Bua in western Flores, but this information can only be understood in the context of the geomorphic history of the cave, and the more general geological, tectonic, and environmental histories of the river valley and region. Thus, a reconstruction of the landscape history of the Wae Racang valley using speleothems, geological structure, tectonic uplift, karst, cave, and terrace development, provides the necessary evidence to determine the formation, age, evolution, and influences on the site. This evidence suggests that Liang Bua was formed as two subterranean chambers approximately 600ka, but could not be occupied until approximately 190ka when the Wae Racang wandered to the southern side of the valley, exposing the chamber and depositing alluvial deposits containing artifacts. During the next approximately 190k.yr., the chambers coalesced and evolved into a multi-level and interconnected cave that was subjected to channel erosion and pooling events by the development of sinkholes. The domed morphology of the front chamber accumulated deep sediments containing well stratified archaeological and faunal remains, but ponded water in the chamber further prevented hominin use of the cave until approximately 100ka. These chambers were periodically influenced by river inundation and volcanic activity, whereas the area outside the cave was greatly influenced by glacial phases, which changed humid forest environments into grassland environments. This combined evidence has important implications for the archaeological interpretation of the site.

RevDate: 2016-11-25
CmpDate: 2009-11-17

Falk D, Hildebolt C, Smith K, et al (2009)

The type specimen (LB1) of Homo floresiensis did not have Laron syndrome.

American journal of physical anthropology, 140(1):52-63.

The type specimen (LB1) of Homo floresiensis has been hypothesized to be a pathological human afflicted with Laron Syndrome (LS), a type of primary growth hormone insensitivity (Hershkovitz et al.: Am J Phys Anthropol 134 [2007] 198-208). Comparing measurements, photographs and three-dimensional, computed-tomography reconstructions of LB1 with data and diagnoses from the literature on LS, we critically evaluate numerous skull and postcranial traits that Hershkovitz et al. identified as being shared by LB1 and patients with LS. The statements regarding most of these traits are new to the clinical literature and lack quantitative support. LB1 and patients with LS differ markedly in the size and shape of the cranium; thickness and pneumatization of cranial bones; morphology of the face, mandible, teeth, and chin; form of the shoulder, wrist, and pelvis; and general body proportions including relative foot size. Claims that patients with LS are similar to LB1 in displaying protracted scapulae, short clavicles, low degrees of humeral torsion, flaring ilia, and curved tibiae are not supported by data or corroborating images. Some points of similarity (e.g., femoral neck-shaft angle, femoral bicondylar angle, and estimated stature) can be found in other hominins, and cannot be considered diagnostic. From our review and analysis, we conclude that LB1 did not suffer from LS.

RevDate: 2010-11-18
CmpDate: 2010-01-27

Falk D, Hildebolt C, Smith K, et al (2009)

LB1's virtual endocast, microcephaly, and hominin brain evolution.

Journal of human evolution, 57(5):597-607.

Earlier observations of the virtual endocast of LB1, the type specimen for Homo floresiensis, are reviewed, extended, and interpreted. Seven derived features of LB1's cerebral cortex are detailed: a caudally-positioned occipital lobe, lack of a rostrally-located lunate sulcus, a caudally-expanded temporal lobe, advanced morphology of the lateral prefrontal cortex, shape of the rostral prefrontal cortex, enlarged gyri in the frontopolar region, and an expanded orbitofrontal cortex. These features indicate that LB1's brain was globally reorganized despite its ape-sized cranial capacity (417cm(3)). Neurological reorganization may thus form the basis for the cognitive abilities attributed to H. floresiensis. Because of its tiny cranial capacity, some workers think that LB1 represents a Homo sapiens individual that was afflicted with microcephaly, or some other pathology, rather than a new species of hominin. We respond to concerns about our earlier study of microcephalics compared with normal individuals, and reaffirm that LB1 did not suffer from this pathology. The intense controversy about LB1 reflects an older continuing dispute about the relative evolutionary importance of brain size versus neurological reorganization. LB1 may help resolve this debate and illuminate constraints that governed hominin brain evolution.

RevDate: 2013-11-21
CmpDate: 2010-01-27

Roberts RG, Westaway KE, Zhao JX, et al (2009)

Geochronology of cave deposits at Liang Bua and of adjacent river terraces in the Wae Racang valley, western Flores, Indonesia: a synthesis of age estimates for the type locality of Homo floresiensis.

Journal of human evolution, 57(5):484-502.

A robust timeframe for the extant cave deposits at Liang Bua, and for the river terraces in the adjoining Wae Racang valley, is essential to constrain the period of existence and time of extinction of Homo floresiensis and other biota that have been excavated at this hominin type locality. Reliable age control is also required for the variety of artifacts excavated from these deposits, and to assist in environmental reconstructions for this river valley and for the region more broadly. In this paper, we summarize the available geochronological information for Liang Bua and its immediate environs, obtained using seven numerical-age methods: radiocarbon, thermoluminescence, optically- and infrared-stimulated luminescence (collectively known as optical dating), uranium-series, electron spin resonance, and coupled electron spin resonance/uranium-series. We synthesize the large number of numerical age determinations reported previously and present additional age estimates germane to questions of hominin evolution and extinction.

RevDate: 2021-12-03
CmpDate: 2009-05-29

Perry GH, NJ Dominy (2009)

Evolution of the human pygmy phenotype.

Trends in ecology & evolution, 24(4):218-225.

Small human body size, or the 'pygmy' phenotype, is characteristic of certain African, Southeast Asian and South American populations. The convergent evolution of this phenotype, and its strong association with tropical rainforests, have motivated adaptive hypotheses that stress the advantages of small size for coping with food limitation, warm, humid conditions and dense forest undergrowth. Most recently, a life-history model has been used to suggest that the human pygmy phenotype is a consequence of early growth cessation that evolved to facilitate early reproductive onset amid conditions of high adult mortality. As we discuss here, these adaptive scenarios are not mutually exclusive and should be evaluated in consort. Findings from this area of research are expected to inform interpretations of diversity in the hominin fossil record, including the purported small-bodied species Homo floresiensis.

RevDate: 2021-10-20
CmpDate: 2009-03-04

Gallagher A (2008)

Size variation in small-bodied humans from palau, micronesia.

PloS one, 3(12):e3939.

BACKGROUND: Recent discoveries on Palau are claimed to represent the remains of small-bodied humans that may display evidence insular size reduction. This claim has yet to be statistically validated

Published postcranial specimens (n = 16) from Palau were assessed relative to recent small-bodied comparative samples. Resampling statistical approaches were employed to test specific hypotheses relating to body size in the Palau sample. Results confirm that the Palau postcranial sample is indisputably small-bodied.

CONCLUSIONS/SIGNIFICANCE: A single, homogenous body size morph is represented in early prehistoric postcrania from Palau. Small body size in early Palauans is an ancestral characteristic and was likely not a consequence of in-situ size reduction. Specimens from Palau have little bearing upon hypothesised insular size reduction in the ancestral lineage of Homo floresiensis.

RevDate: 2009-11-16
CmpDate: 2010-01-27

Baab KL, KP McNulty (2009)

Size, shape, and asymmetry in fossil hominins: the status of the LB1 cranium based on 3D morphometric analyses.

Journal of human evolution, 57(5):608-622.

The unique set of morphological characteristics of the Liang Bua hominins (Homo floresiensis) has been attributed to explanations as diverse as insular dwarfism and pathological microcephaly. This study examined the relationship between cranial size and shape across a range of hominin and African ape species to test whether or not cranial morphology of LB1 is consistent with the basic pattern of static allometry present in these various taxa. Correlations between size and 3D cranial shape were explored using principal components analysis in shape space and in Procrustes form space. Additionally, patterns of static allometry within both modern humans and Plio-Pleistocene hominins were used to simulate the expected cranial shapes of each group at the size of LB1. These hypothetical specimens were compared to LB1 both visually and statistically. Results of most analyses indicated that LB1 best fits predictions for a small specimen of fossil Homo but not for a small modern human. This was especially true for analyses of neurocranial landmarks. Results from the whole cranium were less clear about the specific affinities of LB1, but, importantly, demonstrated that aspects of facial morphology associated with smaller size converge on modern human morphology. This suggests that facial similarities between LB1 and anatomically modern humans may not be indicative of a close relationship. Landmark data collected from this study were also used to test the degree of cranial asymmetry in LB1. These comparisons indicated that the cranium is fairly asymmetrical, but within the range of asymmetry exhibited by modern humans and all extant African ape species. Compared to other fossil specimens, the degree of asymmetry in LB1 is moderate and readily explained by the taphonomic processes to which all fossils are subject. Taken together, these findings suggest that H. floresiensis was most likely the diminutive descendant of a species of archaic Homo, although the details of this evolutionary history remain obscure.

RevDate: 2009-11-16
CmpDate: 2010-01-27

Jungers WL, Larson SG, Harcourt-Smith W, et al (2009)

Descriptions of the lower limb skeleton of Homo floresiensis.

Journal of human evolution, 57(5):538-554.

Bones of the lower extremity have been recovered for up to nine different individuals of Homo floresiensis - LB1, LB4, LB6, LB8, LB9, LB10, LB11, LB13, and LB14. LB1 is represented by a bony pelvis (damaged but now repaired), femora, tibiae, fibulae, patellae, and numerous foot bones. LB4/2 is an immature right tibia lacking epiphyses. LB6 includes a fragmentary metatarsal and two pedal phalanges. LB8 is a nearly complete right tibia (shorter than that of LB1). LB9 is a fragment of a hominin femoral diaphysis. LB10 is a proximal hallucal phalanx. LB11 includes pelvic fragments and a fragmentary metatarsal. LB13 is a patellar fragment, and LB14 is a fragment of an acetabulum. All skeletal remains recovered from Liang Bua were extremely fragile, and some were badly damaged when they were removed temporarily from Jakarta. At present, virtually all fossil materials have been returned, stabilized, and hardened. These skeletal remains are described and illustrated photographically. The lower limb skeleton exhibits a uniquely mosaic pattern, with many primitive-like morphologies; we have been unable to find this combination of ancient and derived (more human-like) features in either healthy or pathological modern humans, regardless of body size. Bilateral asymmetries are slight in the postcranium, and muscle markings are clearly delineated on all bones. The long bones are robust, and the thickness of their cortices is well within the ranges seen in healthy modern humans. LB1 is most probably a female based on the shape of her greater sciatic notch, and the marked degree of lateral iliac flaring recalls that seen in australopithecines such as "Lucy" (AL 288-1). The metatarsus has a human-like robusticity formula, but the proximal pedal phalanges are relatively long and robust (and slightly curved). The hallux is fully adducted, but we suspect that a medial longitudinal arch was absent.

RevDate: 2010-11-18
CmpDate: 2010-01-27

van den Bergh GD, Meijer HJ, Due Awe R, et al (2009)

The Liang Bua faunal remains: a 95k.yr. sequence from Flores, East Indonesia.

Journal of human evolution, 57(5):527-537.

Excavations at Liang Bua, a limestone cave on the island of Flores, East Indonesia, have yielded a well-dated archaeological and faunal sequence spanning the last 95k.yr., major climatic fluctuations, and two human species -H. floresiensis from 95 to 17k.yr.(1), and modern humans from 11k.yr. to the present. The faunal assemblage comprises well-preserved mammal, bird, reptile and mollusc remains, including examples of island gigantism in small mammals and the dwarfing of large taxa. Together with evidence from Early-Middle Pleistocene sites in the Soa Basin, it confirms the long-term isolation, impoverishment, and phylogenetic continuity of the Flores faunal community. The accumulation of Stegodon and Komodo dragon remains at the site in the Pleistocene is attributed to Homo floresiensis, while predatory birds, including an extinct species of owl, were largely responsible for the accumulation of the small vertebrates. The disappearance from the sequence of the two large-bodied, endemic mammals, Stegodon florensis insularis and Homo floresiensis, was associated with a volcanic eruption at 17 ka and precedes the earliest evidence for modern humans, who initiated use of mollusc and shell working, and began to introduce a range of exotic animals to the island. Faunal introductions during the Holocene included the Sulawesi warty pig (Sus celebensis) at about 7ka, followed by the Eurasian pig (Sus scrofa), Long-tailed macaque, Javanese porcupine, and Masked palm civet at about 4ka, and cattle, deer, and horse - possibly by the Portuguese within historic times. The Holocene sequence at the site also documents local faunal extinctions - a result of accelerating human population growth, habitat loss, and over-exploitation.

RevDate: 2009-11-16
CmpDate: 2010-01-27

Larson SG, Jungers WL, Tocheri MW, et al (2009)

Descriptions of the upper limb skeleton of Homo floresiensis.

Journal of human evolution, 57(5):555-570.

Several bones of the upper extremity were recovered during excavations of Late Pleistocene deposits at Liang Bua, Flores, and these have been attributed to Homo floresiensis. At present, these upper limb remains have been assigned to six different individuals - LB1, LB2, LB3, LB4, LB5, and LB6. Several of these bones are complete or nearly so, but some are quite fragmentary. All skeletal remains recovered from Liang Bua were extremely fragile, but have now been stabilized and hardened in the laboratory in Jakarta. They are now curated in museum-quality containers at the National Research and Development Centre for Archaeology in Jakarta, Indonesia. These skeletal remains are described and illustrated photographically. The upper limb presents a unique mosaic of derived (human-like) and primitive morphologies, the combination of which is never found in either healthy or pathological modern humans.

RevDate: 2021-10-20
CmpDate: 2008-12-18

Fitzpatrick SM, Nelson GC, G Clark (2008)

Small scattered fragments do not a dwarf make: biological and archaeological data indicate that prehistoric inhabitants of Palau were normal sized.

PloS one, 3(8):e3015.

UNLABELLED: Current archaeological evidence from Palau in western Micronesia indicates that the archipelago was settled around 3000-3300 BP by normal sized populations; contrary to recent claims, they did not succumb to insular dwarfism.

BACKGROUND: Previous and ongoing archaeological research of both human burial and occupation sites throughout the Palauan archipelago during the last 50 years has produced a robust data set to test hypotheses regarding initial colonization and subsequent adaptations over the past three millennia.

PRINCIPAL FINDINGS: Close examination of human burials at the early (ca. 3000 BP) and stratified site of Chelechol ra Orrak indicates that these were normal sized individuals. This is contrary to the recent claim of contemporaneous "small-bodied" individuals found at two cave sites by Berger et al. (2008). As we argue, their analyses are flawed on a number of different analytical levels. First, their sample size is too small and fragmentary to adequately address the variation inherent in modern humans within and outside of Palau. Second, the size and stature of all other prehistoric (both older and contemporaneous) skeletal assemblages found in Palau fall within the normal parameters of modern human variation in the region, indicating this was not a case of insular dwarfism or a separate migratory group. Third, measurements taken on several skeletal elements by Berger et al. may appear to be from smaller-bodied individuals, but the sizes of these people compares well with samples from Chelechol ra Orrak. Last, archaeological, linguistic, and historical evidence demonstrates a great deal of cultural continuity in Palau through time as expected if the same population was inhabiting the archipelago.

CONCLUSIONS: Prehistoric Palauan populations were normal sized and exhibit traits that fall within the normal variation for Homo sapiens-they do not support the claims by Berger et al. (2008) that there were smaller-bodied populations living in Palau or that insular dwarfism took place such as may be the case for Homo floresiensis.

RevDate: 2022-03-21
CmpDate: 2008-06-26

Tocheri MW, Orr CM, Jacofsky MC, et al (2008)

The evolutionary history of the hominin hand since the last common ancestor of Pan and Homo.

Journal of anatomy, 212(4):544-562.

Molecular evidence indicates that the last common ancestor of the genus Pan and the hominin clade existed between 8 and 4 million years ago (Ma). The current fossil record indicates the Pan-Homo last common ancestor existed at least 5 Ma and most likely between 6 and 7 Ma. Together, the molecular and fossil evidence has important consequences for interpreting the evolutionary history of the hand within the tribe Hominini (hominins). Firstly, parsimony supports the hypothesis that the hand of the last common ancestor most likely resembled that of an extant great ape overall (Pan, Gorilla, and Pongo), and that of an African ape in particular. Second, it provides a context for interpreting the derived changes to the hand that have evolved in various hominins. For example, the Australopithecus afarensis hand is likely derived in comparison with that of the Pan-Homo last common ancestor in having shorter fingers relative to thumb length and more proximo-distally oriented joints between its capitate, second metacarpal, and trapezium. This evidence suggests that these derived features evolved prior to the intensification of stone tool-related hominin behaviors beginning around 2.5 Ma. However, a majority of primitive features most likely present in the Pan-Homo last common ancestor are retained in the hands of Australopithecus, Paranthropus/early Homo, and Homo floresiensis. This evidence suggests that further derived changes to the hands of other hominins such as modern humans and Neandertals did not evolve until after 2.5 Ma and possibly even later than 1.5 Ma, which is currently the earliest evidence of Acheulian technology. The derived hands of modern humans and Neandertals may indicate a morphological commitment to tool-related manipulative behaviors beyond that observed in other hominins, including those (e.g. H. floresiensis) which may be descended from earlier tool-making species.

RevDate: 2021-10-20
CmpDate: 2008-04-22

Gordon AD, Nevell L, B Wood (2008)

The Homo floresiensis cranium (LB1): size, scaling, and early Homo affinities.

Proceedings of the National Academy of Sciences of the United States of America, 105(12):4650-4655.

The skeletal remains of a diminutive small-brained hominin found in Late Pleistocene cave deposits on the island of Flores, Indonesia were assigned to a new species, Homo floresiensis [Brown P, et al. (2004) A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia. Nature 431: 1055-1061]. A dramatically different interpretation is that this material belongs not to a novel hominin taxon but to a population of small-bodied modern humans affected, or unaffected, by microcephaly. The debate has primarily focused on the size and shape of the endocranial cavity of the type specimen, LB1, with less attention being paid to the morphological evidence provided by the rest of the LB1 cranium and postcranium, and no study thus far has addressed the problem of how scaling would affect shape comparisons between a diminutive cranium like LB1 and the much larger crania of modern humans. We show that whether or not the effects of its small cranial size are accounted for, the external cranial morphology of the LB1 cranium cannot be accommodated within a large global sample of normal modern human crania. Instead, the shape of LB1, which is shown by multivariate analysis to differ significantly from that of modern humans, is similar to that of Homo erectus sensu lato, and, to a lesser extent, Homo habilis. Our results are consistent with hypotheses that suggest the Liang Bua specimens represent a diminutive population closely related to either early H. erectus s. l. from East Africa and/or Dmanisi or to H. habilis.

RevDate: 2021-10-20
CmpDate: 2008-06-23

Obendorf PJ, Oxnard CE, BJ Kefford (2008)

Are the small human-like fossils found on Flores human endemic cretins?.

Proceedings. Biological sciences, 275(1640):1287-1296.

Fossils from Liang Bua (LB) on Flores, Indonesia, including a nearly complete skeleton (LB1) dated to 18kyr BP, were assigned to a new species, Homo floresiensis. We hypothesize that these individuals are myxoedematous endemic (ME) cretins, part of an inland population of (mostly unaffected) Homo sapiens. ME cretins are born without a functioning thyroid; their congenital hypothyroidism leads to severe dwarfism and reduced brain size, but less severe mental retardation and motor disability than neurological endemic cretins. We show that the fossils display many signs of congenital hypothyroidism, including enlarged pituitary fossa, and that distinctive primitive features of LB1 such as the double rooted lower premolar and the primitive wrist morphology are consistent with the hypothesis. We find that the null hypothesis (that LB1 is not a cretin) is rejected by the pituitary fossa size of LB1, and by multivariate analyses of cranial measures. We show that critical environmental factors were potentially present on Flores, how remains of cretins but not of unaffected individuals could be preserved in caves, and that extant oral traditions may provide a record of cretinism.

RevDate: 2013-11-21
CmpDate: 2008-06-25

Vaughan CL, MB Blaszczyk (2008)

Dynamic similarity predicts gait parameters for Homo floresiensis and the Laetoli hominins.

American journal of human biology : the official journal of the Human Biology Council, 20(3):312-316.

Late in 2004, the skeletal remains of a pygmy-sized hominin recovered from a cave on the Indonesian island of Flores were first documented, with the authors concluding that the "postcranial anatomy [was] consistent with human-like obligate bipedalism" (Brown et al. [2004]: Nature 431:1055-1061). We have assumed that Homo floresiensis, who was estimated to be 18,000-years-old, walked with a gait pattern that was dynamically similar to modern man. The dynamic similarity hypothesis was also applied to the Australopithecines that left their footprints at Laetoli 4-million-years ago. According to this hypothesis, dimensionless gait parameters can be used in combination with known leg length or step length to calculate velocity of bipedal locomotion. We have gathered data on 20 extant modern humans to calculate the standard estimates of error when predicting gait parameters. We predict that the Homo floresiensis specimen walked at a velocity of 1.11 +/- 0.14 m/s. For the Laetoli footprints, the velocity for Track 1 was estimated to be 1.03 +/- 0.12 m/s and for Track 2 to be 1.14 +/- 0.12 m/s. These latter values for Australopithecines are greater than prior analyses, but are in good agreement with more recent work based on evolutionary robotics. Since modern man walks at 1.44 +/- 0.14 m/s, our results suggest that, despite their diminutive size, these ancient hominins were capable of ranging across a wide geographical area.


ESP Quick Facts

ESP Origins

In the early 1990's, Robert Robbins was a faculty member at Johns Hopkins, where he directed the informatics core of GDB — the human gene-mapping database of the international human genome project. To share papers with colleagues around the world, he set up a small paper-sharing section on his personal web page. This small project evolved into The Electronic Scholarly Publishing Project.

ESP Support

In 1995, Robbins became the VP/IT of the Fred Hutchinson Cancer Research Center in Seattle, WA. Soon after arriving in Seattle, Robbins secured funding, through the ELSI component of the US Human Genome Project, to create the original ESP.ORG web site, with the formal goal of providing free, world-wide access to the literature of classical genetics.

ESP Rationale

Although the methods of molecular biology can seem almost magical to the uninitiated, the original techniques of classical genetics are readily appreciated by one and all: cross individuals that differ in some inherited trait, collect all of the progeny, score their attributes, and propose mechanisms to explain the patterns of inheritance observed.

ESP Goal

In reading the early works of classical genetics, one is drawn, almost inexorably, into ever more complex models, until molecular explanations begin to seem both necessary and natural. At that point, the tools for understanding genome research are at hand. Assisting readers reach this point was the original goal of The Electronic Scholarly Publishing Project.

ESP Usage

Usage of the site grew rapidly and has remained high. Faculty began to use the site for their assigned readings. Other on-line publishers, ranging from The New York Times to Nature referenced ESP materials in their own publications. Nobel laureates (e.g., Joshua Lederberg) regularly used the site and even wrote to suggest changes and improvements.

ESP Content

When the site began, no journals were making their early content available in digital format. As a result, ESP was obliged to digitize classic literature before it could be made available. For many important papers — such as Mendel's original paper or the first genetic map — ESP had to produce entirely new typeset versions of the works, if they were to be available in a high-quality format.

ESP Help

Early support from the DOE component of the Human Genome Project was critically important for getting the ESP project on a firm foundation. Since that funding ended (nearly 20 years ago), the project has been operated as a purely volunteer effort. Anyone wishing to assist in these efforts should send an email to Robbins.

ESP Plans

With the development of methods for adding typeset side notes to PDF files, the ESP project now plans to add annotated versions of some classical papers to its holdings. We also plan to add new reference and pedagogical material. We have already started providing regularly updated, comprehensive bibliographies to the ESP.ORG site.


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One of the most intriguing, and philosophically suggestive, recent scientific findings has been the discovery that the human lineage included several branches at the species level in which each species had developed culture (tool making, mastery of fire, burial of the dead). Before the Chicxulub impact that ended the realm of the dinosaurs, sentience and culture had not occurred in any lineage, despite several hundred million years of evolution. However, in the mammalian radiation that occurred afterwards, several primate lineages occurred. In just the last few million years, one of those lineages diverged into several sentient, culture-developing species. This book explores how only one of those species (ours) survived, while the others went extinct. Recommended. R. Robbins

Electronic Scholarly Publishing
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Papers in Classical Genetics

The ESP began as an effort to share a handful of key papers from the early days of classical genetics. Now the collection has grown to include hundreds of papers, in full-text format.

Digital Books

Along with papers on classical genetics, ESP offers a collection of full-text digital books, including many works by Darwin and even a collection of poetry — Chicago Poems by Carl Sandburg.


ESP now offers a large collection of user-selected side-by-side timelines (e.g., all science vs. all other categories, or arts and culture vs. world history), designed to provide a comparative context for appreciating world events.


Biographical information about many key scientists (e.g., Walter Sutton).

Selected Bibliographies

Bibliographies on several topics of potential interest to the ESP community are automatically maintained and generated on the ESP site.

ESP Picks from Around the Web (updated 07 JUL 2018 )