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Age at First Molar Emergence in Lufengpithecus Lufengensis and Its Available online at www.sciencedirect.com Journal of Human Evolution 54 (2008) 251e257 Age at first molar emergence in Lufengpithecus lufengensis and its implications for life-history evolution Lingxia Zhao*, Qingwu Lu, Wending Zhang Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China Received 9 November 2006; accepted 30 September 2007 Abstract The late Miocene hominoid Lufengpithecus from Yunnan Province, China, is crucial for understanding hominoid evolution in Asia. Given that age at first permanent molar emergence is a key life-history trait in primates, the present study determined the age at death of the Lufeng- pithecus lufengensis juvenile PA868, which was in the process of erupting its first molar. Using a perikymata periodicity of 7e11 days, along with estimation of cusp formation time and the postnatal delay of crown mineralization, perikymata counts obtained from the permanent central incisor and canine germs indicate that the age at death of PA868 was 2.4e4.5 years based on the central incisor germ, and 2.5e4.7 years based on the canine germ. The age at the first molar emergence was actually slightly younger (by about 0.3 years), as demonstrated by tiny wear facets on this tooth, which indicate that gingival emergence had occurred sometime before death. The average age at first molar emergence of Lufeng- pithecus lufengensis PA868 is estimated to be 3.2e3.3 years, with a range of 2.1e4.4 years. In comparison to extant primates and other fossil hominoids, the life history of Lufengpithecus lufengensis is similar to that of extant great apes and the Miocene hominoids Afropithecus turka- nensis and Sivapithecus parvada, as well as Plio-Pleistocene Australopithecus, and different from monkeys, gibbons, and modern humans. Ó 2007 Elsevier Ltd. All rights reserved. Keywords: Age at first molar emergence; Dental development; Life history; Lufengpithecus lufengensis Introduction likely belongs to an ape, and its closest affinities are to Lufeng- pithecus (Wu, 2000). However, archaeological evidence Large-bodied hominoid fossils (Woo, 1957, 1958; Xu and (Huang et al., 1995; Hou et al., 1999, 2002, 2006) from Long- Lu, 1979; Wu et al., 1986; Zheng and Zhang, 1997; Qi and gupo makes this issue complex. More comprehensive research Dong, 2006), all attributable to the genus Lufengpithecus on Lufengpithecus is necessary to discern its appropriate phy- (Wu, 1987; Gao, 1998; Qi et al., 2006) from the late Miocene logenetic placement. The present study aims to assess in of Yunnan Province, China, are critical for understanding greater detail the dental development and life history of Lu- hominoid evolution in Asia. Lufengpithecus has been generally fengpithecus lufengensis. grouped with the Sivapithecus-Pongo lineage, but further One of the profound changes in the evolution of the human study and new discoveries of Lufengpithecus suggest that lineage has been prolongation of the maturational, or life- this genus may not a member of the Ponginae, but may in history, profile. Extensive debate has centered on the questions fact be a member of the Homininae (Gao, 1998; Kelley, of when in the course of human evolution and in which species 2003; Chaimanee et al., 2003, 2004; Gao et al., 2004, 2006). the major part of this change took place (Kelley, 2002; Anem- The early Pleistocene ‘‘Homo’’ mandible found at the Long- one, 2002). Therefore, more data on life history of fossil hom- gupo site of Wushan County, China (Huang et al., 1995), inoids are essential for resolving this issue. ‘‘Life history’’ refers to the timing or scheduling of events from conception to death. An animal’s life history can be sum- * Corresponding author. marized by key variables such as gestation length, neonatal E-mail address: [email protected] (L. Zhao). weight, prenatal and postnatal growth rates, weaning age, 0047-2484/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2007.09.019 252 L. Zhao et al. / Journal of Human Evolution 54 (2008) 251e257 the age at first reproduction, and life span. Clearly, it is diffi- based on the fauna (Flynn and Qi, 1982; Qi, 1985, 1986). Re- cult to obtain such direct data from the fossil record. Dental cent faunal comparisons and paleomagnetic stratigraphic work development can serve as a reliable predictor of a series of indicate that L. lufengensis from Lufeng is somewhat younger critical life-history parameters. Among primates, age at first than L. hudiensis from Yuanmou (Qi et al., 2006; Yue and molar emergence is highly correlated with a variety of life- Zhang, 2006). history traits (Smith, 1991, 1992), and it can be used to Specimen PA868 preserves the right deciduous fourth pre- broadly infer the life histories of fossil primate species. molar and permanent first molar, as well as five right perma- Based on the dental development of the central incisor and nent tooth crown germs within the crypts of the mandible: canine germs, the present study estimates the age at death and I1,I2,C,P3, and P4 (Figs. 1 and 2). All five crown germs the age at first molar emergence of the Lufengpithecus lufengen- are incomplete and roots are not visible radiographically sis individual represented by the juvenile mandible PA868, and (Fig. 2). The M1 was in the process of erupting when the in- explores the life-history profile of this species. Because L. lufen- dividual died, with the crown already beyond the mandibular gensis is the latest Miocene large-bodied hominoid, this analy- alveolar margin and nearly up to the level of the fully func- sis provides important data for understanding the evolution of tional occlusal plane (Figs. 1 and 2). life history in Miocene and Plio-Pleistocene hominoids, as Two size clusters are evident in the L. lufengensis sample, well as in extant great apes and humans. and the size of the PA868 M1 crown (9.1 mm in length and 8.3 mm in breath) places it among the smaller specimens. Kel- Materials and methods ley and Xu (1991) suggested that the two clusters represent males and females, indicating that L. lufengensis was more Zhao and He (2005) previously gave a preliminary report sexually dimorphic in its postcanine dentition than living hom- on the estimation of the age at first molar emergence in inoids, perhaps exceeding all extant anthropoids. Thus, if we PA868 based only on an X-ray photograph as part of a review consider the whole assemblage to represent a single highly di- paper on dental development in the large-bodied Miocene morphic species, then it is likely that PA868 belongs to a fe- hominoids from Yunnan. In the present study, we have done male individual. more direct work on the specimen to determine its age at death, particularly using scanning electron microscopy to ex- Estimation of the age at death of PA868 amine two developing tooth crown germs (the right central in- cisor and canine) for estimating crown formation time, which The method used here to determine age at death is similar to is the key process for determining the age at death. In the fol- the one employed by Kelley (1997, 2002) and Kelley and Smith lowing analysis, we revise the earlier estimate of age at first (2003) to reconstruct age at death for Sivapithecus parvada molar emergence for the PA868 individual with a full descrip- specimen GSP 11536 and Afropithecus turkanensis specimen tion of the methods and a more thorough analysis. KNM-MO 26. The age at death for individual PA868 could be determined using crown formation times of the tooth germs, Condition of the specimen PA868 which were still developing when the individual died. In lingual view, the three anterior teethdI1,I2, and the caninedare ex- Specimen PA868 is a juvenile right mandibular corpus posed due to damage to the outer bone (Fig. 1). The I1 and ca- (Fig. 1), discovered in 1980 in stratigraphic level D-3, H nine are more suitable for estimating the age of this individual. square, of the Shihuiba site in Lufeng County of Yunnan Prov- Age at death was calculated by adding the time between 0 0 ince, China. The Shihuiba site (E104 4 , N25 1 ) is located birth and the initiation of I1 or canine crown mineralization about 9 km north of Lufeng town, 60 km northwest of Kunm- to the time of I1 or canine crown formation until the individ- ing. The age of the L. lufengensis layers is approximately 8 Ma ual’s death. Crown formation time in teeth that have not Fig. 1. Juvenile mandible of L. lufengensis specimen PA868: (a) buccal view, (b) lingual view. L. Zhao et al. / Journal of Human Evolution 54 (2008) 251e257 253 with a dental pick and needle probe, exposing much of the sur- faces of the tooth crowns (Fig. 1). The perikymata on the ex- posed surfaces were clear under a microscope. The exposed I1 and canine were molded using silica, and a replica was made using epoxy resin, coated with a thin layer of gold-palladium, and examined with a JEOL1600 scanning electron micro- scope. Montage photos were taken of the tooth surfaces from the developing cervical region to cusp edge (Fig. 3). From the montage, we obtained the perikymata number on the surface from the cusp to the developing cervix. The peri- odicity of perikymata, or Retzius lines, was determined from sections made using other L. lufengensis teeth (Zhao et al., 2000; Schwartz et al., 2003). Ages at initiation of I1 and canine mineralization in PA868 were estimated according to compar- Fig. 2. Radiograph of specimen PA868. ative data from all extant apes and humans for which histolog- ical data are available (Beynon et al., 1998; Reid et al., completed their development was calculated as the sum of the 1998a,b; Reid and Dean, 2006).
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