Dental data challenge the ubiquitous presence of Homo in the Cradle of Humankind.

The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel-dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

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

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.

Cochlear morphology of Indonesian Homo erectus from Sangiran.

Homo erectus s.l. is key for deciphering the origin and subsequent evolution of genus Homo. However, the characterization of this species is hindered by the existence of multiple variants in both mainland and insular Asia, as a result of divergent chronogeographical evolutionary trends, genetic isolation, and interbreeding with other human species. Previous research has shown that cochlear morphology embeds taxonomic and phylogenetic information that may help infer the phylogenetic relationships among hominin species. Here we describe the cochlear morphology of two Indonesian H. erectus individuals (Sangiran 2 and 4), and compare it with a sample of australopiths, Middle to Late Pleistocene humans, and extant humans by means of linear measurements and both principal components and canonical variates analyses performed on shape ratios. Our results indicate that H. erectus displays a mosaic morphology that combines plesiomorphic (australopithlike) features (such as a chimplike round cochlear cross section and low cochlear thickness), with derived characters of later humans (a voluminous and long cochlea, possibly related to hearing abilities)-consistent with the more basal position of H. erectus. Our results also denote substantial variation between the two studied individuals, particularly in the length and radius of the first turn, as well as cross-sectional shape. Given the small size of the available sample, it is not possible to discern whether such differences merely reflect intraspecific variation among roughly coeval H. erectus individuals or whether they might result from greater age differences between them than currently considered. However, our results demonstrate that most characters found in later humans were already present in Indonesian H. erectus, with the exception of Neanderthals, which display an autapomorphic condition relative to other Homo species.

Dietary versatility of Early Pleistocene hominins.

New geochemical data from the Malawi Rift (Chiwondo Beds, Karonga Basin) fill a major spatial gap in our knowledge of hominin adaptations on a continental scale. Oxygen (δ18O), carbon (δ13C), and clumped (Δ47) isotope data on paleosols, hominins, and selected fauna elucidate an unexpected diversity in the Pleistocene hominin diet in the various habitats of the East African Rift System (EARS). Food sources of early Homo and Paranthropus thriving in relatively cool and wet wooded savanna ecosystems along the western shore of paleolake Malawi contained a large fraction of C3 plant material. Complementary water consumption reconstructions suggest that ca. 2.4 Ma, early Homo (Homo rudolfensis) and Paranthropus (Paranthropus boisei) remained rather stationary near freshwater sources along the lake margins. Time-equivalent Paranthropus aethiopicus from the Eastern Rift further north in the EARS consumed a higher fraction of C4 resources, an adaptation that grew more pronounced with increasing openness of the savanna setting after 2 Ma, while Homo maintained a high versatility. However, southern African Paranthropus robustus had, similar to the Malawi Rift individuals, C3-dominated feeding strategies throughout the Early Pleistocene. Collectively, the stable isotope and faunal data presented here document that early Homo and Paranthropus were dietary opportunists and able to cope with a wide range of paleohabitats, which clearly demonstrates their high behavioral flexibility in the African Early Pleistocene.

Inner tooth morphology of Homo erectus from Zhoukoudian. New evidence from an old collection housed at Uppsala University, Sweden.

Locality 1, in the Lower Cave of the Zhoukoudian cave complex, China, is one of the most important Middle Pleistocene paleoanthropological and archaeological sites worldwide, with the remains of c. 45 Homo erectus individuals, 98 mammalian taxa, and thousands of lithic tools recovered. Most of the material collected before World War II was lost. However, besides two postcranial elements rediscovered in China in 1951, four human permanent teeth from the 'Dragon Bone Hill,' collected by O. Zdansky between 1921 and 1923, were at the time brought to the Paleontological Institute of Uppsala University, Sweden, where they are still stored. This small sample consists of an upper canine (PMU 25719), an upper third molar (PMU M3550), a lower third premolar crown (PMU M3549), and a lower fourth premolar (PMU M3887). Some researchers have noted the existence of morpho-dimensional differences between the Zhoukoudian and the H. erectus dental assemblage from Sangiran, Java. However, compared to its chrono-geographical distribution, the Early to Middle Pleistocene dental material currently forming the Chinese-Indonesian H. erectus hypodigm is quantitatively meager and still poorly characterized for the extent of its endostructural variation. We used micro-focus X-ray tomography techniques of virtual imaging coupled with geometric morphometrics for comparatively investigating the endostructural conformation (tissue proportions, enamel thickness distribution, enamel-dentine junction morphology, pulp cavity shape) of the four specimens stored in Uppsala, all previously reported for their outer features. The results suggest the existence of time-related differences between continental and insular Southeast Asian dental assemblages, the Middle Pleistocene Chinese teeth apparently retaining an inner signature closer to the likely primitive condition represented by the Early Pleistocene remains from Java, while the Indonesian stock evolved toward tooth structural simplification.

New cranium of the large cercopithecid primate Theropithecus oswaldi leakeyi (Hopwood, 1934) from the paleoanthropological site of Makuyuni, Tanzania.

The Pleistocene hominin site of Makuyuni, near Lake Manyara, Tanzania, is known for fossils attributable to Homo and Acheulean artifacts (Ring et al., 2005; Kaiser et al., 2010; Frost et al., 2012). Here we describe the fossil primate material from the Manyara Beds, which includes the first nearly complete female cranium of Theropithecus oswaldi leakeyi and a proximal tibia from the same taxon. The cranium is dated to between 633 and 780 Ka and the tibia to the Pleistocene. The T. oswaldi lineage is one of the most important among Neogene mammals of Africa: it is both widespread and abundant. The size of the dentition, cranium, and tibia all confirm the previously recognized trend of increasing body size in this lineage and make their taxonomic assignments secure. The morphology of this specimen provides new insights into the evolution of this lineage through time, as well as its geographic variation and sexual dimorphism. The cranium also shows damage consistent with a mammalian carnivore, most likely a felid. The identification of this material as representing T. o. leakeyi agrees with the Middle Pleistocene age estimates for the MK4 locality in particular and the Manyara Beds in general.

Persistent C3 vegetation accompanied Plio-Pleistocene hominin evolution in the Malawi Rift (Chiwondo Beds, Malawi).

The development of East African savannas is crucial for the origin and evolution of early hominins. These ecosystems, however, vary widely in their fraction of woody cover and today range from closed woodland to open grassland savanna. Here, we present the first Plio-Pleistocene long-term carbon isotope (δ(13)C) record from pedogenic carbonate and Suidae teeth in the southern East African Rift (EAR). These δ(13)C data from the Chiwondo and Chitimwe Beds (Karonga Basin, Northern Malawi) represent a southern hemisphere record in the EAR, a key region for reconstructing vegetation patterns in today's Zambezian Savanna, and permit correlation with data on the evolution and migration of early hominins in today's Somali-Masai Endemic Zone. The sediments along the northwestern shore of Lake Malawi contain fossils attributed to Homo rudolfensis and Paranthropus boisei. The associated hominin localities (Uraha, Malema) are situated between the well-known hominin bearing sites of the Somali-Masai Endemic Zone in the Eastern Rift and the Highveld Grassland in southern Africa, and fill an important geographical gap for hominin research. Persistent δ(13)C values around -9‰ from pedogenic carbonate and suid enamel covering the last ∼4.3 Ma indicate a C3-dominated closed environment with regional patches of C4-grasslands in the Karonga Basin. The overall fraction of woody cover of 60-70% reflects significantly higher canopy density in the Malawi Rift than the Eastern Rift through time. The discrepancy between the two savanna types originated in the Late Pliocene, when the Somali-Masai ecosystem started to show increasing evidence for open, C4-dominated landscapes. Based on the Malawi δ(13)C data, the evolution of savanna ecosystems in Eastern Africa followed different patterns along the north-south extent of the EAR. The appearance of C4-grasses is considered a driver of evolutionary faunal shifts, but despite the difference of ecosystem evolution in the north, similar hominins and suids occurred in both landscapes, pointing to distinct habitat flexibility and also nutritional versatility.

The early Pleistocene deciduous hominid molar FS-72 from the Sangiran Dome of Java, Indonesia: A taxonomic reappraisal based on its comparative endostructural characterization.

OBJECTIVES: Among the ten fossil hominid deciduous teeth reported so far from the Pleistocene sediments of the Sangiran Dome of Java are two isolated lower second molars: specimens PCG.2 from the Kabuh Formation and FS-72 from the Pucangan Formation. While PCG.2 appears to be certainly attributable to Homo erectus, FS-72 is somewhat more problematic, even though it is commonly listed within the Indonesian H. erectus hypodigm. Largely because of its large size, it was originally attributed to Meganthropus paleojavanicus. Subsequent study highlighted a set of metric and nonmetric crown features also found in Australopith and African early Homo (notably H. habilis) homologues. An additional problem with the taxonomic assignment of isolated teeth from the Pleistocene of Java is the presence of Pongo in these same deposits. METHODS: To assess the taxonomic affinity of FS-72, we investigated its inner structure (tissue proportions and enamel-dentine junction morphology) by using techniques of 2-3D virtual imaging coupled with geometric morphometric analyses. RESULTS: The results show that FS-72 has thinner enamel compared to fossil and recent humans and that its topographic repartition more closely follows the pongine pattern. It also exhibits a Pongo-like elongated morphology of the enamel-dentine junction, with proportionally lower and mesiodistally spaced dentine horns. CONCLUSIONS: Given the morphological and metric similarities between fossil orangutan and H. erectus molars, we tested the hypothesis that its internal morphology more closely resembles the patterns evinced by PCG.2 and modern humans than Pongo. Accordingly, we consider that FS-72 more likely represents a dm2 of Pongo rather than Homo.

Correction: The Swine Plasma Metabolome Chronicles "Many Days" Biological Timing and Functions Linked to Growth.

[This corrects the article DOI: 10.1371/journal.pone.0145919.].

The ROCEEH Out of Africa Database (ROAD): A large-scale research database serves as an indispensable tool for human evolutionary studies.

Large scale databases are critical for helping scientists decipher long-term patterns in human evolution. This paper describes the conception and development of such a research database and illustrates how big data can be harnessed to formulate new ideas about the past. The Role of Culture in Early Expansions of Humans (ROCEEH) is a transdisciplinary research center whose aim is to study the origins of culture and the multifaceted aspects of human expansions across Africa and Eurasia over the last three million years. To support its research, the ROCEEH team developed an online tool named the ROCEEH Out of Africa Database (ROAD) and implemented its web-based applications. ROAD integrates geographical data as well as archaeological, paleoanthropological, paleontological and paleobotanical content within a robust chronological framework. In fact, a unique feature of ROAD is its ability to dynamically link scientific data both spatially and temporally, thereby allowing its reuse in ways that were not originally conceived. The data stem from published sources spanning the last 150 years, including those generated by the research team. Descriptions of these data rely on the development of a standardized vocabulary and profit from online explanations of each table and attribute. By synthesizing legacy data, ROAD facilitates the reuse of heritage data in novel ways. Database queries yield structured information in a variety of interoperable formats. By visualizing data on maps, users can explore this vast dataset and develop their own theories. By downloading data, users can conduct further quantitative analyses, for example with Geographic Information Systems, modeling programs and artificial intelligence. In this paper, we demonstrate the innovative nature of ROAD and show how it helps scientists studying human evolution to access datasets from different fields, thereby connecting the social and natural sciences. Because it permits the reuse of "old" data in new ways, ROAD is now an indispensable tool for researchers of human evolution and paleogeography.

Dietary strategies of Pleistocene Pongo sp. and Homo erectus on Java (Indonesia).

During the Early to Middle Pleistocene, Java was inhabited by hominid taxa of great diversity. However, their seasonal dietary strategies have never been explored. We undertook geochemical analyses of orangutan (Pongo sp.), Homo erectus and other mammalian Pleistocene teeth from Sangiran. We reconstructed past dietary strategies at subweekly resolution and inferred seasonal ecological patterns. Histologically controlled spatially resolved elemental analyses by laser-based plasma mass spectrometry confirmed the preservation of authentic biogenic signals despite the effect of spatially restricted diagenetic overprint. The Sr/Ca record of faunal remains is in line with expected trophic positions, contextualizing fossil hominid diet. Pongo sp. displays marked seasonal cycles with ~3 month-long strongly elevated Sr/Ca peaks, reflecting contrasting plant food consumption presumably during the monsoon season, while lower Sr/Ca ratios suggest different food availability during the dry season. In contrast, omnivorous H. erectus shows low and less accentuated intra-annual Sr/Ca variability compared to Pongo sp., with δ13C data of one individual indicating a dietary shift from C4 to a mix of C3 and C4 plants. Our data suggest that H. erectus on Java was maximizing the resources available in more open mosaic habitats and was less dependent on variations in seasonal resource availability. While still influenced by seasonal food availability, we infer that H. erectus was affected to a lesser degree than Pongo sp., which inhabited monsoonal rain forests on Java. We suggest that H. erectus maintained a greater degree of nutritional independence by exploiting the regional diversity of food resources across the seasons.

Geometric morphometrics and paleoproteomics enlighten the paleodiversity of Pongo.

Pleistocene Pongo teeth show substantial variation in size and morphology, fueling taxonomic debates about the paleodiversity of the genus. We investigated prominent features of the enamel-dentine-junction junction (EDJ)-phylogenetically informative internal structures-of 71 fossil Pongo lower molars from various sites by applying geometric morphometrics and conducted paleoproteomic analyses from enamel proteins to attempt to identify extinct orangutan species. Forty-three orangutan lower molars representing Pongo pygmaeus and Pongo abelii were included for comparison. The shape of the EDJ was analyzed by placing five landmarks on the tip of the main dentine horns, and 142 semilandmarks along the marginal ridges connecting the dentine horns. Paleoproteomic analyses were conducted on 15 teeth of Late Pleistocene Pongo using high-resolution tandem mass spectrometry. The geometric morphometric results show variations in EDJ shape regarding aspects of the height and position of the dentine horns and connecting ridges. Despite the issue of molar position and sample size, modern molars are distinguished from fossil counterparts by their elongated tooth outline and narrowly positioned dentine horns. Proteomic results show that neither a distinction of P. pygmaeus and P. abelii, nor a consistent allocation of fossil specimens to extant species is feasible. Based on the EDJ shape, the (late) Middle to Late Pleistocene Pongo samples from Vietnam share the same morphospace, supporting the previous allocation to P. devosi, although substantial overlap with Chinese fossils could also indicate close affinities with P. weidenreichi. The hypothesis that both species represent one chronospecies cannot be ruled out. Two fossil specimens, one from Tam Hay Marklot (Laos, Late Pleistocene), and another from Sangiran (Java, Early to Middle Pleistocene), along with some specimens within the Punung sample (Java), exhibit affinities with Pongo abelii. The Punung fossils might represent a mix of early Late Pleistocene and later specimens (terminal Pleistocene to Holocene) related to modern Pongo. The taxonomy and phylogeny of the complete Punung sample needs to be further investigated.

Variation in enamel thickness within the genus Homo.

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having 'thick enamel' thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.

Brief communication: Identification reassessment of the isolated tooth Krapina D58 through occlusal fingerprint analysis.

High variability in the dentition of Homo can create uncertainties in the correct identification of isolated teeth. For instance, standard tooth identification criteria cannot determine with absolute certainty if an isolated tooth is a second or third maxillary molar. In this contribution, using occlusal fingerprint analysis, we reassess the identification of Krapina D58 (Homo neanderthalensis), which is catalogued as a third maxillary molar. We have hypothesized that the presence/absence of the distal occlusal wear facets can be used to differentiate second from third maxillary molars. The results obtained confirm our hypothesis, showing a significant difference between second and third maxillary molars. In particular we note the complete absence of Facets 7 and 10 in all third molars included in this analysis. The presence of these facets in Krapina D58 eliminates the possibility that it is a third maxillary molar. Consequently it should be reclassified as a second molar. Although this method is limited by the degree of dental wear (i.e., unworn teeth cannot be analyzed) and to individual molars in full occlusion, it can be used for tooth identification when other common criteria are not sufficient to discriminate between second and third maxillary molars.

Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia.

Since the first discovery of Pithecanthropus (Homo) erectus by E. Dubois at Trinil in 1891, over 200 hominid dentognathic remains have been collected from the Early to Middle Pleistocene deposits of Java, Indonesia, forming the largest palaeoanthropological collection in South East Asia. Most of these fossils are currently attributed to H. erectus. However, because of the substantial morphological and metric variation in the Indonesian assemblage, some robust specimens, such as the partial mandibles Sangiran 5 and Sangiran 6a, were formerly variably allocated to other taxa (Meganthropus palaeojavanicus, Pithecanthropus dubius, Pongo sp.). To resolve the taxonomic uncertainty surrounding these and other contentious Indonesian hominid specimens, we used occlusal fingerprint analysis (OFA) to reconstruct their chewing kinematics; we also used various morphometric approaches based on microtomography to examine the internal dental structures. Our results confirm the presence of Meganthropus as a Pleistocene Indonesian hominid distinct from Pongo, Gigantopithecus and Homo, and further reveal that Dubois's H. erectus paratype molars from 1891 are not hominin (human lineage), but instead are more likely to belong to Meganthropus.

Craniofacial architectural constraints and their importance for reconstructing the early Homo skull KNM-ER 1470.

OBJECTIVE: Our objective is to exploit architectural constraint for the analysis and interpretation of craniofacial form, which we apply here to the reconstruction of the early Homo cranium KNM-ER 1470. We are motivated to perform this study because in the absence of biological criteria our preconceptions are likely to govern our concept of craniofacial form. STUDY DESIGN: We reassembled the fragmented parts--left and right halves of the calvaria and the face--according to mammalian craniofacial architectural constraints described by Donald H. Enlow and colleagues. RESULTS: When evaluated on a biological premise, KNM-ER 1470 is found to have a more prognathic midface than commonly appreciated. The relationship between facial prognathism and cranial capacity also provides an estimate downward for this specimen, from 752cc to ca. 700cc. CONCLUSION: Awareness of our preconceptions is critical to the performance of relatively unbiased research in fields characterized by interpretations of morphology. When perceptual bias is relatively minimized, applied here as an architecturally constrained of KNM-ER 1470 craniofacial skeleton, we are able to provide the scientific community with a more tractable Gestalt perspective of form.

Disentangling isolated dental remains of Asian Pleistocene hominins and pongines.

Scholars have debated the taxonomic identity of isolated primate teeth from the Asian Pleistocene for over a century, which is complicated by morphological and metric convergence between orangutan (Pongo) and hominin (Homo) molariform teeth. Like Homo erectus, Pongo once showed considerable dental variation and a wide distribution throughout mainland and insular Asia. In order to clarify the utility of isolated dental remains to document the presence of hominins during Asian prehistory, we examined enamel thickness, enamel-dentine junction shape, and crown development in 33 molars from G. H. R. von Koenigswald's Chinese Apothecary collection (11 Sinanthropus officinalis [= Homo erectus], 21 "Hemanthropus peii," and 1 "Hemanthropus peii" or Pongo) and 7 molars from Sangiran dome (either Homo erectus or Pongo). All fossil teeth were imaged with non-destructive conventional and/or synchrotron micro-computed tomography. These were compared to H. erectus teeth from Zhoukoudian, Sangiran and Trinil, and a large comparative sample of fossil Pongo, recent Pongo, and recent human teeth. We find that Homo and Pongo molars overlap substantially in relative enamel thickness; molar enamel-dentine junction shape is more distinctive, with Pongo showing relatively shorter dentine horns and wider crowns than Homo. Long-period line periodicity values are significantly greater in Pongo than in H. erectus, leading to longer crown formation times in the former. Most of the sample originally assigned to S. officinalis and H. erectus shows greater affinity to Pongo than to the hominin comparative sample. Moreover, enamel thickness, enamel-dentine junction shape, and a long-period line periodicity value in the "Hemanthropus peii" sample are indistinguishable from fossil Pongo. These results underscore the need for additional recovery and study of associated dentitions prior to erecting new taxa from isolated teeth.

The Swine Plasma Metabolome Chronicles "Many Days" Biological Timing and Functions Linked to Growth.

The paradigm of chronobiology is based almost wholly upon the daily biological clock, or circadian rhythm, which has been the focus of intense molecular, cellular, pharmacological, and behavioral, research. However, the circadian rhythm does not explain biological timings related to fundamental aspects of life history such as rates of tissue/organ/body size development and control of the timing of life stages such as gestation length, age at maturity, and lifespan. This suggests that another biological timing mechanism is at work. Here we focus on a "many days" (multidien) chronobiological period first observed as enigmatic recurring growth lines in developing mammalian tooth enamel that is strongly associate with all adult tissue, organ, and body masses as well as life history attributes such as gestation length, age at maturity, weaning, and lifespan, particularly among the well studied primates. Yet, knowledge of the biological factors regulating the patterning of mammalian life, such as the development of body size and life history structure, does not exist. To identify underlying molecular mechanisms we performed metabolome and genome analyses from blood plasma in domestic pigs. We show that blood plasma metabolites and small non-coding RNA (sncRNA) drawn from 33 domestic pigs over a two-week period strongly oscillate on a 5-day multidien rhythm, as does the pig enamel rhythm. Metabolomics and genomics pathway analyses actually reveal two 5-day rhythms, one related to growth in which biological functions include cell proliferation, apoptosis, and transcription regulation/protein synthesis, and another 5-day rhythm related to degradative pathways that follows three days later. Our results provide experimental confirmation of a 5-day multidien rhythm in the domestic pig linking the periodic growth of enamel with oscillations of the metabolome and genome. This association reveals a new class of chronobiological rhythm and a snapshot of the biological bases that regulate mammalian growth, body size, and life history.

Wide faces or large canines? The attractive versus the aggressive primate.

Hominids display marked body size dimorphism, suggestive of strong sexual selection, yet they lack significant sex differences in canine size that are commonly associated with intrasexual competition in primates. We resolve this paradox by examining sex differences in hominoid facial morphology. We show that chimpanzees, but not gorillas, exhibit clear sexual dimorphism in face width, over and above that expected based on sex differences in body size. We show that this facial dimorphism, expressed as an index, is negatively correlated with canine dimorphism among anthropoid primates. Our findings suggest that a lack of canine dimorphism in anthropoids is not owing to weak sexual selection, but rather is associated with strong sexual selection for broader face width. Enlarged cheek-bones are linked with attractiveness in humans, and we propose that the evolution of a broad face and loss of large canines in hominid males results from mate choice.