PALEOANTHROPOLOGY The Growth Pattern Of Neandertals .
R ES E A RC HPALEOANTHROPOLOGYThe growth pattern of Neandertals,reconstructed from a juvenileskeleton from El Sidrón (Spain)Antonio Rosas,1*† Luis Ríos,1,2† Almudena Estalrrich,1,3 Helen Liversidge,4Antonio García-Tabernero,1 Rosa Huguet,5 Hugo Cardoso,6 Markus Bastir,1Carles Lalueza-Fox,7 Marco de la Rasilla,8 Christopher Dean9Neandertals provide us with an importantperspective on our own biology (1). Bothmodern humans and Neandertals arosefrom a recent common ancestor along independent evolutionary lines, becominglarge-brained hominins but with contrastingbody forms. Developing a large brain is energetically expensive and places a constraint onsomatic growth (2). The unusually high cost ofmodern human brain development is greatestduring the infant and childhood periods andseems to require a compensatory slowing ofchildhood body growth (2, 3). Neandertals hadlarger average cranial capacity than modernhumans, but little is known about the ontogenetic trajectories of brain and body underlyingthis difference.Some studies have proposed that a larger brainin Neandertals can be explained by a faster rateof early postnatal growth (4), yet others have1Paleoanthropology Group, Department of Paleobiology,Museo Nacional de Ciencias Naturales (MNCN)–ConsejoSuperior de Investigaciones Científicas (CSIC), Calle JoséGutiérrez Abascal 2, 28006 Madrid, Spain. 2Department ofPhysical Anthropology, Aranzadi Society of Sciences,Zorroagagaina 11, 20014 Donostia–San Sebastián, Gipuzkoa,Spain. 3Department of Paleoanthropology, SenckenbergResearch Institute and Natural History Museum Frankfurt,Senckenberganlage 25, 60325 Franckfurta, Germany. 4QueenMary University of London, Institute of Dentistry, TurnerStreet, London E1 2AD, UK. 5Institut Català de PaleoecologiaHumana i Evolució Social–Unidad Asociada al CSIC, CampusSescelades (Edifici W3), Universitat Rovira i Virgili, CarrerMarcel.lí Domingo s/n, 43007 Tarragona, Spain.6Department of Archaeology, Simon Fraser University,Burnaby, British Columbia V5A1S6, Canada. 7Institute ofEvolutionary Biology (CSIC–Universitat Pompeu Fabra),Carrer Dr. Aiguader 88, 08003 Barcelona, Spain. 8Área dePrehistoria Departamento de Historia, Universidad de Oviedo,Calle Teniente Alfonso Martínez s/n, 33011 Oviedo, Spain.9Department of Cell and Developmental Biology, UniversityCollege London, Gower Street, London WC1E 6BT, UK.*Corresponding author. Email: arosas@mncn.csic.es †Theseauthors contributed equally to this work.Rosas et al., Science 357, 1282–1287 (2017)proposed a longer period of growth as an explanation (5, 6). However, in large-brained hominins like modern humans and Neandertals, anaccelerated pace of brain growth, coincident withaccelerated somatic growth, would impose a highenergetic cost (2). Yet the trade-off between thedifferent aspects of somatic and neural growthin Neandertals, particularly during the juvenileperiod, remains unclear.Here we describe a partial juvenile Neandertalskeleton from the 49-thousand-year-old site ofEl Sidrón (Asturias, Spain). The specimen has amixed dentition of deciduous and permanentteeth and preserves cranial, dental, and postcranial remains (Figs. 1 and 2A and supplementary text 1 and 2), providing a rare opportunityto estimate an age at death from daily dentalincremental markings preserved in teeth, againstwhich to compare many aspects of its dentoskeletal maturation. Chronological age is fundamental for assessing patterns of maturation indifferent dento-skeletal systems, both within individuals and between species. This approachallowed us to ask what the probability is thatthis specimen would fit within or lie beyondthe ranges of modern human variation and represent its own pattern of dental and skeletalmaturation.The El Sidrón cave system (Asturias, Spain)(Fig. 1C and supplementary text 1) has providedmore than 2500 remains of seven adults andsix immature individuals belonging to a singleNeandertal group (7) with close kinship relations (8). Among them, a partial immature skeleton was recovered with up to 36% (left side)preserved. Virtually all of the remains associated with this individual come from the 1-m2G-6 square grid of the archaeological site (supplementary text 2), and importantly, several werefound in anatomical association. From the three22 September 20171 of 6Downloaded from http://science.sciencemag.org/ on December 4, 2017Ontogenetic studies help us understand the processes of evolutionary change. Previousstudies on Neandertals have focused mainly on dental development and inferred anaccelerated pace of general growth. We report on a juvenile partial skeleton (El Sidrón J1)preserving cranio-dental and postcranial remains. We used dental histology to estimatethe age at death to be 7.7 years. Maturation of most elements fell within the expected range ofmodern humans at this age. The exceptions were the atlas and mid-thoracic vertebrae,which remained at the 5- to 6-year stage of development. Furthermore, endocranial featuressuggest that brain growth was not yet completed. The vertebral maturation pattern andextended brain growth most likely reflect Neandertal physiology and ontogenetic energyconstraints rather than any fundamental difference in the overall pace of growth in thisextinct human.mitochondrial DNA lineages detected withinthis Neandertal group, this individual belongs toline C of the group and was tentatively identifiedas the child of adult female 4 and the oldersibling of infant 1 (8).A number of diagnostic Neandertal featuresare present throughout the skeleton (supplementary text 2). Although ancient DNA failed to confirm the sex, group-specific evaluation of caninesize and bone robusticity strongly suggests that itwas male (supplementary text 2). Dental development, with a near-complete first molar (M1)root, would place him in the juvenile stage ofhominin life history (3). Height and weightestimates indicate that he was a sturdy individual, weighing 26 kg and standing 111 cm tall atthe time of death (supplementary text 2). Biosocialmarkers indicate that El Sidrón juvenile 1 (J1) wasright-handed, with evidence that he was involvedwith, or learning, adult behaviors and economicactivities (9). Apart from mild linear dental enamelhypoplasia around the age of 2 to 3 years, there isno other evidence of pathology. Several postmortem cut marks appear on some of the bones.Age at death was first established by dentalhistology. Daily incremental markings in twosections of El Sidrón J1 first left upper molar(see materials and methods, figs. S1 and S2, andsupplementary text 3) were used to estimate anaverage age at death of 7.69 years (range: 7.61to 7.78 years). Biological maturity was then assessed using modern human references for dental,skeletal, and somatic maturation (supplementarytext 4 to 7).Dental maturity was assessed in two ways.Individuals from two reference samples of modern children of known age (n 4072 and 6829)were assigned a radiographic stage of development for each tooth (supplementary text 4).Compared with the first sample, dental maturityof El Sidrón J1 was judged to be 71.7 to 72.1%complete. Probability density plots for mean ageof transition entering each tooth stage werecomputed from the second sample, and El SidrónJ1 fell well within the modern human range forall tooth types represented (Fig. 2B). Skeletalmaturity (SM) and skeletal age (SA) were assessedfrom six secondary ossification centers from theelbow, hand, wrist, and knee, by applying established pediatric methods (figs. S3 to S5 andsupplementary text 5). The SA interval rangedfrom 6 to 10 years, with an average of 7.62 2 years(table S7). Maturity of each individual vertebra wasassessed in two ways. Individuals from a sampleof 106 immature modern human skeletons (ofwhich 70 were of known age and sex) were assigneda stage of fusion of the neurocentral synchondrosis(NS) and a radiographic stage of development forthe lower M1 (materials and methods). Probabilitydensity functions for the mean age of transitionentering fusion of the NS of the first cervicalvertebra (C1) and the 3rd to 11th thoracic vertebrae (T3 to T11) were computed from the knownage sample (Fig. 2C). The same procedure wasapplied to the total sample using the mean ageentering the respective M1 stage scored (Fig. 2D).Compared with chronological and dental age,
R ES E A RC H R E PO R TRosas et al., Science 357, 1282–1287 (2017)Downloaded from http://science.sciencemag.org/ on December 4, 2017maturation of each available vertebra of El SidrónJ1 fell at the extreme end of the modern humanrange (fig. S6 and supplementary text 6). SM of ElSidrón J1 vertebral column [fused C3-C5-C6, T1-T2,and L2-L3 (second and third lumbar vertebrae);unfused C1 and four middle thoracic vertebrae]fits the modern human observed sequence of NSvertebral fusion but corresponds chronologically to younger individuals between 4 to 6 yearsof age (Fig. 2C). Percentage of adult size (PAS)attained (10) was calculated as a measure ofsomatic maturation for 53 measurements throughthe cranial, axial, and appendicular skeleton (supplementary text 7). In comparison with a sampleof 11 modern human skeletons with chronological age (CA) between 6.5 and 8.5 years, valuesof El Sidrón J1 fell within (49 variables) or veryclose to (4 variables) the modern human range(Fig. 3A). The height-for-age of El Sidrón J1 alsofell within the range of modern humans (11)(Fig. 3B), with Neandertal body shape featuresalready observable at 7.7 years (12, 13) (Fig. 3C).Clearly visible bone resorption areas on theinner aspect of the occipital poles provide someevidence that brain expansion was still ongoing(Fig. 4 and supplementary text 8). Resorptionactivity is a characteristic of the period of braingrowth in modern humans (14). These observations suggest that specific locations on the occipital lobe and cerebellum of El Sidrón J1 werestill increasing in size. The extremely well-definedimprints of the gyri and sinus impressions on theinternal aspect of SD-2300, as well as the narrowdural sinus grooves (supplementary text 8), furthersuggest that the encephalon was still exertinggrowing pressure on the neurocranium.A consensus value for endocranial volume of 1330 cm3 (supplementary text 8) was computed,which represents 87.5% of mean Neandertal adultendocranial volume (1520 cm3). On average, modern humans achieve 90% of adult brain weightby 5 years old (15) and 95% by 7 years (16). Thissuggests that further brain growth in El SidrónJ1 would likely have continued beyond the timeexpected in modern humans at 7.7 years.The dental and skeletal maturity of El SidrónJ1 were compared with modern humans. Dentaldevelopment is what one would expect for achild of his age. This contrasts with previousfindings from isolated cranio-dental materialthat have reported a faster pace of dental development (17, 18). Compared with early Homospecimens at a comparable stage of dental development, El Sidrón J1 is at least 2.7 yearsolder than a 2-million-year-old Homo specimen, StW 151 (19–22), but almost identical inage (7.78 years) to a 315-thousand-year-old Homosapiens specimen from Jebel Irhoud, Morocco(23), that shows a prolonged modern human–like period of dental developmental (24). At7.7 years of age, El Sidrón J1 shows a secondincisor (I2) at the stage of alveolar eruption, anM2 at the stage of crown completion, and anM3 crypt present in the mandible. It is, therefore,no longer possible to assume that these eventsoccurred earlier at 6 years of age, or that M2erupted at 8 years of age, in all Neandertals (18).Fig. 1. El Sidrón J1 Neandertal skeleton. (A) The138 specimens, 30 of which are deciduous andpermanent teeth, that compose the El Sidrón J1skeleton. Cranial, axial, and appendicular elements arewell represented, but the legs (especially the right one) are less well represented. (B) Virtualreconstruction of the El Sidrón J1 skull and endocast based on the same cranial bonespecimens shown in (A). (C) Localization of the El Sidrón site in the Iberian Peninsula(Asturias, Spain). A map of the karst system depicts the 25-m-long Osario gallery(shadowed), as well as the excavated area of this gallery. Most of the specimens of El SidrónJ1 were recovered from the G-6 1-m2grid of the excavation.22 September 20172 of 6
R ES E A RC H R E PO R T0.00.30.6M3 CryptM2 RiM1 RcP4 RiP3 R1/4C R1/4I2 R3/4I1 RcI2 AE444.8355.6677.76Cr 3/4 Cr Ri Rcl8C1C2C3C4C5C6C7T1T28R1/410R1/2R3/4 Rc12A 1/2AcC1Downloaded from http://science.sciencemag.org/ on December 4, 4L5T9Stage 1T10Stage 2Stage 3T11T12Fig. 2. Dental and vertebral maturation of El Sidrón J1. (A) Computedtomography (CT) scan image of the mandible of El Sidrón J1, with the enamelshown in green. (B) Probability density plots (PDPs) for mean age oftransition entering each mandibular tooth stage scored for El Sidrón J1 in aradiographic sample (n 6829 individuals) of modern children of knownchronological age (CA). Red vertical lines represent the CA of El SidrónJ1 from dental histology (7.69 years; range: 7.61 to 7.78 years). (C) Maturationof the spine relative to CA in El Sidrón J1 and modern humans. Thevertical axis represents the presacral vertebral column; the horizontal axisrepresents age in years. For each vertebra, the three successive maturationstages are represented (see vertebral diagrams in the figure): stage one,unfused posterior synchondrosis (PS) and neurocentral synchondrosis (AS);stage two, fused PS and unfused neurocentral synchondrosis (NS); stagethree, fused PS and NS. A sample of 70 known CA skeletons was used todevelop PDPs for mean age of transition entering fusion of the NS for eachRosas et al., Science 357, 1282–1287 (2017)22 September 2017vertebra (from stage two to three). El Sidrón J1 is displayed in red, andthe two oldest modern human cases (4.83 and 5.6 years) with a spinematuration similar to that of El Sidrón J1 (unfused C1 and middle thoracicvertebrae) are represented in black. The C1 falls within the P 0.01shaded area of the PDP, whereas the thoracic vertebrae would fall outside(T3 and T4), in the P 0.05 shaded area (T5, T6, T7, and T9), or underthe PDP (T8). (D) Maturation of the spine relative to dental maturation in ElSidrón J1 and in modern humans. The vertical axis represents C1 and thethoracic vertebrae, whereas stages of formation of the first permanentmandibular molar are represented on the horizontal axis. A sample of106 modern human skeletons of diverse origins was used to develop PDPsfor mean first molar formation stage entering fusion of the NS for eachvertebra (from stage two to three). The vertical red line, representingcomplete root formation of the first permanent molar of El Sidrón J1, fallsin the P 0.05 area in all PDPs.3 of 6
R ES E A RC H R E PO R TPercentage of adult 105250Femur diaphysis length (mm)El Sidrón J110200798150651004321NeandertalsModern humans5012345678910El Sidrón J1100880Clavicle length (mm)7601140NeandertalsModern humansEl Sidrón J1Modern humansx 2 SD012345678910Age (years)As with El Sidrón J1, new ages at death (18) foryounger Neandertal specimens (Engis 2, Gibraltar2, Krapina B, and Obi-Rakhmat 1) fall withinmodern ranges, but two older specimens, Scladinaand Le Moustier 1 (18, 25), now seem unexpectedly young. An assumed early age, 2155 days (18),of initial M3 mineralization (18, 25) or foreshortened estimates of root formation times (26)might explain this. Clearly, variation in Neandertal dental development would have been asgreat as today but may generally have tendedlengths of El Sidrón J1 and a Neandertal ontogenetic series (17), with 80modern human skeletons with CA of 0 to 9 years, with fitted quadraticmodels (Neandertals, R2 0.968; modern humans, R2 0.952). (C) Claviclelength of El Sidrón J1 and a Neandertal ontogenetic series (18), with 51modern human skeletons with CA of 0 to 9 years, with fitted quadraticmodels (Neandertals, R2 1; modern humans, R2 0.889). 1, La Ferrassie4/Le Moustier 2; 2, La Ferrassie 4b/La Ferrassie 4; 3, Mezmaiskaya;4, Kiik-Koba 2; 5, Shanidar 10; 6, Dederiyeh 2; 7, Dederiyeh 1; 8, Roc de Marsal1; 9, La Ferrassie 6; 10, Cova Negra; 11, Amud 7.toward the more advanced end of the modernhuman spectrum.A Homo erectus juvenile aged between 7.6 to8.8 years (KNM WT 15000) shows evidence ofboth advanced dental development and earlierattainment of body mass and stature than is typical of modern humans of a similar age (22, 27 ).However, SA and PAS are also within the modern range, given the limited level of biologicalresolution of SA and PAS estimation. Growthand development in this juvenile Neandertal fit22 September 2017the typical features of human ontogeny, wherethere is slow somatic growth between weaningand puberty (3, 28) that may offset the cost ofgrowing a large brain. Moreover, a slower pace ofgrowth provides an opportunity for shifts in boththe rate and timing of brain growth (4–6, 15).Even so, divergent morphogenetic trajectoriesunderlying shape differences, such as brain development (29–31) and cranio-facial morphology(32, 33), can exist within this broadly human growthpattern.4 of 6Downloaded from http://science.sciencemag.org/ on December 4, 20170Fig. 3. Somatic maturation and size-by-age of El Sidrón J1. (A) Percentageof adult size (PAS) of El Sidrón J1 in comparison with 11 modern humanskeletons with CA between 6.5 and 8.5 years (supplementary text 7). L,length variables, including bones from the appendicular and axial skeletoncontributing to stature (i.e., vertebral body height); W, width variables,including diaphysis and epiphysis from the appendicular and axial (articularwidths, diaphyseal circumferences, vertebral body widths); C, craniofacial andcentral nervous system–associated variables from cranial bones, mandible,and vertebrae. Variables are listed in supplementary text 7. (B) FemoralRosas et al., Science 357, 1282–1287 (2017)300115
R ES E A RC H R E PO R TThe one divergent aspect of ontogeny is thetiming of maturation within the vertebral column. In all hominoids, the NS of the middlethoracic vertebrae and the atlas are the last tofuse, but in this Neandertal, it appears that fusionoccurs 2 years later than in modern humans (orcloser to M1 root closure than to the M1 rootbeing a quarter to half formed).At 1.5 to 2 years old, the state of maturation ofthe complete spine of the Dederiyeh 1 child(34, 35) suggests that, earlier in ontogeny, whenthe posterior synchondroses fuse, Neandertalsfollowed a vertebral maturation schedule similarto that of modern humans. The later fusion ofthe NS could reflect a decoupling of certainRosas et al., Science 357, 1282–1287 (2017)fiber bundles and insertion of Sharpey’s fibers, whereas resorption areas arerecognized as anisotropic resorption bays (Howship’s lacunae). Bonedeposition (blue) is easily illustrated by collagen fiber bundles, at timeschanging direction to form a wavy impression (1B and 4A). Resting depositioncan be identified by a dense and uniform bright surface in which thedeposition of bone matrix masks other histological features, includingcollagen fiber and Sharpey’s fibers (4D and 5D). Insertions of Sharpey’s fibersare spread out over the deposition surfaces (2B and 5D). A well-definedreversal line can be seen in 3C, and an example of taphonomic alteration (e.g.,scratches) is depicted in 3B. Scale bar (bottom of central panel), 3 cm.smaller-scale aspects of growth and maturationin these extinct humans in the transition fromthe childhood to the juvenile stage. Although theimplications of this are unknown, they may berelated to the characteristically expanded Neandertal torso (36, 37) or to ongoing growth of theneuraxis. Together, these findings suggest thatlate Neandertal neural growth pattern exhibits adegree of modularity relative to dent
PALEOANTHROPOLOGY The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain) Antonio Rosas,1*† Luis Ríos,1,2† Almudena Estalrrich,1,3 Helen Liversidge,4 Antonio García-Tabernero, 1Rosa Huguet,5 Hugo Cardoso,6 Markus Bastir, Carles Lalueza-Fox,7 Marco de la Rasilla,8 Christopher Dean9 Ontogenetic studies help us understand the processes of .
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