The origins of musicality in the motion of primates.

Animals communicate acoustically to report location, identity, and emotive state to conspecifics. Acoustic signals can also function as displays to potential mates and as territorial advertisement. Music and song are terms often reserved only for humans and birds, but elements of both forms of acoustic display are also found in non-human primates. While culture, bonding, and side-effects all factor into the emergence of musicality, biophysical insights into what might be signaled by specific acoustic features are less well understood. OBJECTIVES: Here we probe the origins of musicality by evaluating the links between musical features (structural complexity, rhythm, interval, and tone) and a variety of potential ecological drivers of its evolution across primate species. Alongside other hypothesized causes (e.g. territoriality, sexual selection), we evaluated the hypothesis that perilous arboreal locomotion might favor musical calling in primates as a signal of capacities underlying spatio-temporal precision in motor tasks. MATERIALS AND METHODS: We used musical features found in spectrographs of vocalizations of 58 primate species and corresponding measures of locomotion, diet, ranging, and mating. Leveraging phylogenetic information helped us impute missing data and control for relatedness of species while selecting among candidate multivariate regression models. RESULTS: Results indicated that rapid inter-substrate arboreal locomotion is highly correlated with several metrics of music-like signaling. Diet, alongside mate-choice and range size, emerged as factors that also correlated with complex calling patterns. DISCUSSION: These results support the hypothesis that musical calling may function as a signal, to neighbors or potential mates, of accuracy in landing on relatively narrow targets.

On reappearance and complexity in musical calling.

Music is especially valued in human societies, but music-like behavior in the form of song also occurs in a variety of other animal groups including primates. The calling of our primate ancestors may well have evolved into the music of modern humans via multiple selective scenarios. But efforts to uncover these influences have been hindered by the challenge of precisely defining musical behavior in a way that could be more generally applied across species. We propose an acoustic focused reconsideration of "musicality" that could help enable independent inquiry into potential ecological pressures on the evolutionary emergence of such behavior. Using published spectrographic images (n = 832 vocalizations) from the primate vocalization literature, we developed a quantitative formulation that could be used to help recognize signatures of human-like musicality in the acoustic displays of other species. We visually scored each spectrogram along six structural features from human music-tone, interval, transposition, repetition, rhythm, and syllabic variation-and reduced this multivariate assessment into a concise measure of musical patterning, as informed by principal components analysis. The resulting acoustic reappearance diversity index (ARDI) estimates the number of different reappearing syllables within a call type. ARDI is in concordance with traditional measures of bird song complexity yet more readily identifies shorter, more subtly melodic primate vocalizations. We demonstrate the potential utility of this index by using it to corroborate several origins scenarios. When comparing ARDI scores with ecological features, our data suggest that vocalizations with diversely reappearing elements have a pronounced association with both social and environmental factors. Musical calls were moderately associated with wooded habitats and arboreal foraging, providing partial support for the acoustic adaptation hypothesis. But musical calling was most strongly associated with social monogamy, suggestive of selection for constituents of small family-sized groups by neighboring conspecifics. In sum, ARDI helps construe musical behavior along a continuum, accommodates non-human musicality, and enables gradualistic co-evolutionary paths between primate taxa-ranging from the more inhibited locational calls of archaic primates to the more exhibitional displays of modern apes.

Using Social Network Methods to Test for Assortment of Prosociality among Korean High School Students.

Assortative interaction among altruistic individuals is a necessary condition for the evolution of cooperation. The requirement for assortment holds regardless of whether a meta-population is subdivided into distinct and isolated subgroups or has ephemeral boundaries with a high migration rate. The assumption, however, is rarely tested directly. In this paper, we develop a method to test for assortment of prosociality in network-structured data. The method is applied to a friendship network collected from 238 Korean students attending the same high school. A mixing matrix was used to explore the presence of assortative friendship among more prosocial individuals. An exponential random graph model of network structure that accounts for additional observed relational propensities (higher-than-expected number of people nominating no friends) and sampling constraints (upper bound on friendship nominations) found that individual prosociality predicted friendship propensity, and that individuals with higher prosocial scores had a higher probability of befriending other more prosocial individuals. The results reveal that a considerable level of assortment of prosociality characterizes this population.

Palmar Creases: Classification, Reliability and Relationships to Fetal Alcohol Spectrum Disorders (FASD).

A normal human palm contains 3 major creases: the distal transverse crease; the proximal transverse crease; and the thenar crease. Because permanent crease patterns are thought to be laid down during the first trimester, researchers have speculated that deviations in crease patterns could be indicative of insults during fetal development. The purpose of this study was twofold: (1) to compare the efficacy and reliability of two coding methods, the first (M1) classifying both "simiana" and Sydney line variants and the second (M2) counting the total number of crease points of origin on the radial border of the hand; and (2) to ascertain the relationship between palmar crease patterns and fetal alcohol spectrum disorders (FASD). Bilateral palm prints were taken using the carbon paper and tape method from 237 individuals diagnosed with FASD and 190 unexposed controls. All prints were coded for crease variants under M1 and M2. Additionally, a random sample of 98 matched (right and left) prints was selected from the controls to determine the reliabilities of M1 and M2. For this analysis, each palm was read twice, at different times, by two readers. Intra-observer Kappa coefficients were similar under both methods, ranging from 0.804-0.910. Inter-observer Kappa coefficients ranged from 0.582-0.623 under M1 and from 0.647-0.757 under M2. Using data from the entire sample of 427 prints and controlling for sex and ethnicity (white v. non-white), no relationship was found between palmar crease variants and FASD. Our results suggest that palmar creases can be classified reliably, but palmar crease patterns may not be affected by fetal alcohol exposure.

Age-independent increases in male salivary testosterone during horticultural activity among Tsimane forager-farmers.

Testosterone plays an important role in mediating male reproductive trade-offs in many vertebrate species, augmenting muscle and influencing behavior necessary for male-male competition and mating-effort. Among humans, testosterone may also play a key role in facilitating male provisioning of offspring as muscular and neuromuscular performance are deeply influenced by acute changes in testosterone. This study examines acute changes in salivary testosterone among 63 Tsimane men ranging in age from 16-80 (mean 38.2) years during one-hour bouts of tree-chopping while clearing horticultural plots. The Tsimane forager-horticulturalists living in the Bolivian Amazon experience high energy expenditure associated with food production, have high levels of parasites and pathogens, and display significantly lower baseline salivary testosterone than age-matched US males. Mixed-effects models controlling for BMI and time of specimen collection reveal increased salivary testosterone (p<0.001) equivalent to a 48.6% rise, after one hour of tree chopping. Age had no effect on baseline (p=0.656) or change in testosterone (p=0.530); self-reported illness did not modify testosterone change (p=0.488). A comparison of these results to the relative change in testosterone during a competitive soccer tournament in the same population reveals larger relative changes in testosterone following resource production (tree chopping), compared to competition (soccer). These findings highlight the importance of moving beyond a unidimensional focus on changes in testosterone and male-male aggression to investigate the importance of testosterone-behavior interactions across additional male fitness-related activities. Acutely increased testosterone during muscularly intensive horticultural food production may facilitate male productivity and provisioning.

Progesterone and ovulation across stages of the transition to menopause.

OBJECTIVE: Detailed characterization of progesterone and ovulation across the menopausal transition provides insight into conception risk and mechanisms of reproductive aging. METHODS: Participants (n = 108, aged 25-58 y) collected daily urine specimens for 6-month intervals in each of 5 consecutive years. Specimens were assayed for pregnanediol glucuronide (PDG), luteinizing hormone (LH), follicle-stimulating hormone (FSH), and estrone glucuronide (E1G). Reproductive stage was determined using cycle length variance. A hierarchical algorithm was used to identify ovulation. Linear mixed-effects models estimated (1) the frequency and day of ovulation by age and stage; (2) differences in FSH, LH, and E1G levels between ovulatory (O) and anovulatory (AO) cycles; and (3) total PDG levels and PDG levels in O cycles by age and stage. RESULTS: The probability of AO cycles increased across the perimenopause (P < 0.0001); reproductive stage was a stronger predictor than age of the probability of anovulation. Most cycles in late perimenopause were AO (>60%), but one quarter of cycles longer than 60 days were O. Average day of ovulation was later in the late perimenopause (mean [SD] cycle day, 27 [25] d) compared with the premenopause. FSH and LH levels were higher and E1G levels were lower in AO than O cycles (P < 0.0001 for each). Total PDG decreased in the late perimenopause, but 95th percentile PDG in O cycles declined steadily across the transition. CONCLUSIONS: Exposure to the risk of conception in women experiencing cycles long enough to classify them as late perimenopausal is far from negligible. Reproductive stage is more informative than age about PDG levels and the likelihood of anovulation.

Total and unopposed estrogen exposure across stages of the transition to menopause.

Detailed characterization of estrogen dynamics during the transition to menopause is an important step toward understanding its potential implications for reproductive cancers developing in the transition years. We conducted a 5-year prospective study of endogenous levels of total and unopposed estrogen. Participants (n=108; ages 25-58 years) collected daily urine specimens for 6 months in each of 5 consecutive years. Specimens were assayed for estrone-3-glucuronide (E1G) and pregnanediol-3-glucuronide. Linear mixed-effects models were used to estimate exposure to total and unopposed estrogen by age and reproductive stage. Reproductive stage was estimated using menstrual cycle length variance. E1G mean area under the curve and mean E1G 5th and 95th percentiles represented total estrogen exposure. An algorithm identifying days of above-baseline E1G that coincided with the days of baseline pregnanediol-3-glucuronide was used to identify days of unopposed estrogen. Mean E1G area under the curve increased with age in the pretransition and early transition and decreased in the late transition. Ninety-fifth percentile E1G levels did not decline until after menopause, whereas 5th percentile levels declined from the early transition to the postmenopause. The number of days of unopposed estrogen was significantly higher during the transition compared with the pretransition. Given the length of time women spend in the transition, they are exposed to more total and unopposed estrogen than has been previously appreciated. Coupled with epidemiologic evidence on lifetime exposure to estrogen, these results suggest that variation in the amount of time spent in the transition may be an important risk factor for reproductive cancers.

Monitoring reproductive aging in a 5-year prospective study: aggregate and individual changes in luteinizing hormone and follicle-stimulating hormone with age.

OBJECTIVE: This study describes age-related changes in luteinizing hormone (LH) and follicle-stimulating hormone (FSH) in a 5-year prospective study of reproductive aging. DESIGN: Participants (n = 156 college-educated, white, US women; 25 to 58 y) were recruited from the TREMIN Research Program on Women's Health. They collected daily urine specimens for 6 months in each of 5 consecutive years. Specimens were assayed for LH and FSH. Aggregate changes were calculated in LH and FSH with age, and multilevel models were used to estimate individual hormone trajectories and within-woman and between-woman variances by age. RESULTS: Aggregate LH levels increased beginning after age 45; FSH increased at all ages, accelerating after age 45. Individual-level patterns with age included the following: reproductive-age LH and FSH levels, with increasing FSH and increasing or decreasing LH (ages 20 to 49); rapidly increasing LH and FSH (ages 40 to 59); and increasing or steady postmenopausal LH and FSH (ages 46 to 62). FSH levels were consistently high in the latter category, but LH levels overlapped with levels found in younger women (<45 y). Individual LH patterns showed more variability (5% to 35% of total variance) than FSH (3% to 22% of total variance). Both hormones had relatively low variation within individuals compared with between-woman differences (65% to 97% of total variance). CONCLUSIONS: Aggregate-level data do not reflect differences across women and oversimplify the age-related increases and variability in LH and FSH. Individual FSH levels are not distinguishable from reproductive-age levels until after rapid perimenopausal increases in FSH occur; individuals vary in whether their postmenopausal LH levels are distinguishable from reproductive-age levels.

The length of perimenopausal menstrual cycles increases later and to a greater degree than previously reported.

OBJECTIVE: To demonstrate that the perimenopausal increase in menstrual cycle length presented by Treloar et al. was biased by misidentified menopause dates, mean values classified by calendar year, and exclusion of menstrual cycles straddling two calendar years; and to use the revised data to investigate women's experiences of longer perimenopausal cycles. DESIGN: Secondary analysis of prospectively collected menstrual cycle data. SETTING: Center for Population and Health, Georgetown University. PATIENT(S): One hundred twenty white, college-educated, US women in the Tremin Research Program on Women's Health. INTERVENTION(S): None. MAIN OUTCOME MEASURE(S): Mean cycle length and time spent in >40-day cycles, by year before menopause. RESULT(S): Mean estimates for each of the 4 years before menopause were 30.48, 35.02, 45.15, and 80.22 days, respectively, compared with the original analysis: 33.60, 43.91, 55.87, and 54.58 days. In the year before menopause, the majority of women spent >or=75% of their time in cycles >40 days long. CONCLUSION(S): Treloar's estimates of mean cycle length were biased. Long cycles occurred throughout perimenopause, but the largest increase in mean cycle length did not occur until the final year before menopause. New estimates of the time spent in cycles >40 days may be useful clinically as well as epidemiologically for assessing menopausal onset and symptomatology.

Distribution of postpartum amenorrhea in rural Bangladeshi women.

Previous studies of postpartum amenorrhea (PPA) demonstrated distinct subgroups of women with short and long durations of amenorrhea. This phenomenon was attributed to cases where breastfeeding is absent because of pregnancy loss or infant death, or confusion of postpartum bleeding with resumption of menses. We explored these ideas using data from an 11-month prospective study in Bangladesh in which 858 women provided twice-weekly interviews and urine specimens for up to 9 months; 300 women were observed while experiencing PPA. The resulting exact, interval-censored, or right-censored durations were used to estimate parameters of two-component mixture models. A mixture of two Weibull distributions provided the best fit to the observations. The long-duration subgroup made up 84% (+/- 4% SE) of the population, with a mean duration of 457 (+/- 31) days. The short-duration subgroup had a mean duration of 94 (+/- 17) days. Three covariates were associated with the duration of PPA: women whose husbands had high-wage employment had a greater probability of falling in the short-duration subgroup; women in the long-duration subgroup whose husbands seasonally migrated had shorter periods of PPA within the subgroup; and mothers in the short-duration subgroup who gave birth during the monsoon season experienced a shortened duration of PPA within the subgroup. We conclude that the bimodal distribution of PPA reflects biological or behavioral heterogeneity rather than shortcomings of data collection.

Monitoring reproductive aging in a 5-year prospective study: aggregate and individual changes in steroid hormones and menstrual cycle lengths with age.

OBJECTIVE: We describe a 5-year prospective study of reproductive aging, and present analyses of steroid hormone and menstrual cycle changes with age. DESIGN: Participants were college-educated white women, primarily of northern European ancestry, recruited from the Tremin Research Program on Women's Health (n = 156, 25-58 years). In each of 5 consecutive years, they collected daily urine specimens for 6 months and recorded menstrual bleeds for all months. Urine specimens were assayed for estrone-3-glucuronide (E1G) and pregnanediol-3-glucuronide (PDG), urinary metabolites of estradiol and progesterone. Using multilevel models, we estimated hormone and cycle-length trajectories for individual women and within- and between-woman variance by age. RESULTS: At the aggregate level, PDG declined beginning in the 30s, E1G increased into the 40s before declining, and cycle length became more variable with age. Individual-level models revealed substantial hormonal variation across women, in both absolute levels and rates of change. Most women showed declining E1G by the late 40s, declining PDG in the 30s, and increasing mean cycle length in the 40s. Hormonal variation decreased with age; cycle length variation decreased and then increased. Within individual women, cycle lengths were highly variable while hormone levels were more stable. Women differed more from each other in hormone levels than for cycle lengths. CONCLUSIONS: Aggregate-level analyses show general changes in steroid hormones and cycle length but cannot show variation within and across women. Individuals' cycle lengths were too variable to predict hormone levels. Clinicians should obtain more data on individual women's hormonal patterns when determining fertility or menopause treatments.

Longitudinal analysis of deciduous tooth emergence: IV. Covariate effects in Japanese children.

Emergence of the deciduous teeth is generally considered to be robust to moderate environmental insults, malnutrition, and disease. Consequently, deciduous tooth emergence has been used to assess growth and development and for age estimation in children. In this paper, we examine the way in which nutritional status and other covariates affect deciduous tooth emergence in a sample of 114 Japanese children born in Tokyo in 1914 and 1924. Parametric survival analysis was used to quantify the effects of nutritional status, breastfeeding behavior, and sex on the hazard of deciduous tooth emergence. Children of poor nutritional status exhibited significantly delayed emergence of all deciduous teeth, with effects that ranged from 14-29% increases in mean emergence times. Children of medium nutritional status exhibited increases in mean emergence times of 5-9% for the canines and lower molars, and 13-17% for the incisors. Partial breastfeeding had no effect on tooth emergence, but children who were not breastfed at all showed delayed emergence of the upper incisors. No significant sex differences in emergence were found. The findings contradict the idea that moderate malnutrition has little effect on deciduous tooth emergence. Furthermore, nutritional differences may account for some of the observed differences among populations in the timing of tooth emergence.

Hormonal correlates for the initiation of breast-feeding in Bangladeshi women.

Hormonal changes that occur before or during parturition are known to trigger early postpartum maternal behaviors in many mammals. In humans, little evidence has been found for hormonal mediation of early postpartum maternal behavior. In this paper, we investigate associations between fetoplacental hormone concentrations in late pregnancy on the time from parturition to initiation of breast-feeding. A sample of 91 pregnant rural Bangladeshi women, enrolled in a 9-month prospective study, provided twice-weekly urine specimens and structured interviews. The subjects provided self-reports of time from parturition to initiation of breast-feeding. Specimens were assayed for urinary concentrations of human chorionic gonadotropin (hCG), pregnanediol-3alpha-glucuronide (PdG, a metabolite of progesterone), and urinary estrone conjugates (E1C). Parametric hazards analysis was used to investigate the effects of hCG, PdG, and E1C concentrations and other covariates (mother's age, parity, and child's sex) on the duration from parturition to breast-feeding. Mother's age, parity, the child's sex, hCG, and PdG showed no association with the onset of breast-feeding. Urinary E1C was significantly associated with time to initiation of breast-feeding, explaining about 4% of the variation in the behavior. The relationship was positive so that higher prepartum concentrations of EIC were associated with later times to initiation of breast-feeding. The direction of this relationship is opposite that found for many other species of mammals but is consistent with some recent findings in primates.

Comparison of specific gravity and creatinine for normalizing urinary reproductive hormone concentrations.

BACKGROUND: Specific gravity (SG) may perform as well as creatinine (CR) correction for adjusting urinary hormone concentrations, as well as offer some advantages. We compared the two methods and applied them to US and Bangladeshi specimens to evaluate their use in different populations. METHODS: Pearson correlations between serum concentrations and SG, CR, and uncorrected urinary concentrations were compared using paired daily urine and serum specimens from one menstrual cycle from 30 US women. Corrected urinary estrone conjugate and pregnanediol glucuronide concentrations were compared with serum estradiol and progesterone. Urine specimens across one menstrual cycle from 13 Bangladeshi women were used to evaluate the applicability of both methods to a nonindustrialized population. Linear mixed-effects models were used to compare CR and SG values in the Bangladeshi vs US specimens. RESULTS: There was no significant difference between SG-corrected vs serum and CR-corrected vs serum correlations for either assay. Usable CR results were obtained for all US specimens, but 37% of the Bangladeshi specimens were below the CR assay limit of detection. The Bangladeshi sample had significantly lower CR and higher inter- and intrasubject CR variability than the US sample. CONCLUSIONS: SG is a potentially useful alternative to CR correction for normalizing urinary steroid hormone concentrations, particularly in settings where CR values are highly variable or unusually low.

Patterns for the initiation of breastfeeding in humans.

Establishment of lactation has important biological and emotional health consequences for the newborn. Even so, substantial variation within a culture and among different cultures is seen in the onset of breastfeeding. Parametric mixture models are used to explore this variation and to uncover general human patterns for the initiation of breastfeeding. The model components reflect two hypothesized patterns of behavior. The first component is a "natural" pattern of breastfeeding that reflects, to some extent, a general mammalian behavior. The second component arises through culturally mediated behaviors that affect the initiation of breastfeeding. The model was fit by maximum likelihood to interval- and right-censored observations on 26220 mother-infant pairs collected from 25 previously published studies of breastfeeding behavior. Both model components were clearly statistically identified. Effects of cultural and geographic covariates were found to have significant effects on all components of the model. Although there is clear evidence for two distinct patterns of behavior in the initiation of breastfeeding, the results suggest that learned behaviors play an important role in mediating even "natural" breastfeeding behavior.

Longitudinal analysis of deciduous tooth emergence: III. Sexual dimorphism in Bangladeshi, Guatemalan, Japanese, and Javanese children.

Previous studies, mostly in European populations, found sex differences in the pattern of deciduous tooth emergence. Most studies find that the anterior dentition in males is precocial relative to the female dentition, and the pattern reverses so that females lead males in the emergence of the posterior deciduous dentition. Less is known about sex differences in the dental development and emergence of non-European populations. Here we examine the pattern of sex differences in deciduous tooth emergence in Japanese, Javanese, Guatemalan, and Bangladeshi children. The data come from four longitudinal or mixed longitudinal studies using similar study protocols. Survival analysis was used to estimate parameters of a log-normal distribution of emergence for each of the 10 teeth of the left dentition, and sexual dimorphism was assessed by sex-specific differences in mean emergence times and by Bennett's index. The results support the pattern of developmental cross-over observed in other populations. We conclude that there is little evidence to support the hypothesis of Tanguay et al. ([1984] J. Dent. Res. 63:65-68) that ethnic factors mediate sex differences in the emergence of deciduous teeth.

Regression-based prediction of net energy expenditure in children performing activities at high altitude.

We developed a simple, non-invasive, and affordable method for estimating net energy expenditure (EE) in children performing activities at high altitude. A regression-based method predicts net oxygen consumption (VO(2)) from net heart rate (HR) along with several covariates. The method is atypical in that, the "net" measures are taken as the difference between exercise and resting VO(2) (DeltaVO(2)) and the difference between exercise and resting HR (DeltaHR); DeltaVO(2) partially corrects for resting metabolic rate and for posture, and DeltaHR controls for inter-individual variation in physiology and for posture. Twenty children between 8 and 13 years of age, born and raised in La Paz, Bolivia (altitude 3,600m), made up the reference sample. Anthropometric measures were taken, and VO(2) was assessed while the children performed graded exercise tests on a cycle ergometer. A repeated-measures prediction equation was developed, and maximum likelihood estimates of parameters were found from 75 observations on 20 children. The final model included the variables DeltaHR, DeltaHR(2), weight, and sex. The effectiveness of the method was established using leave-one-out cross-validation, yielding a prediction error rate of 0.126 for a mean DeltaVO(2) of 0.693 (SD 0.315). The correlation between the predicted and measured DeltaVO(2) was r = 0.917, suggesting that a useful prediction equation can be produced using paired VO(2) and HR measurements on a relatively small reference sample. The resulting prediction equation can be used for estimating EE from HR in free-living children performing habitual activities in the Bolivian Andes.

Urinary estrone conjugate and pregnanediol 3-glucuronide enzyme immunoassays for population research.

BACKGROUND: Monitoring of reproductive steroid hormones at the population level requires frequent measurements, hormones or metabolites that remain stable under less than ideal collection and storage conditions, a long-term supply of antibodies, and assays useful for a range of populations. We developed enzyme immunoassays for urinary pregnanediol 3-glucuronide (PDG) and estrone conjugates (E1Cs) that meet these criteria. METHODS: Enzyme immunoassays based on monoclonal antibodies were evaluated for specificity, detection limit, parallelism, recovery, and imprecision. Paired urine and serum specimens were analyzed throughout menstrual cycles of 30 US women. Assay application in different populations was examined with 23 US and 42 Bangladeshi specimens. Metabolite stability in urine was evaluated for 0-8 days at room temperature and for 0-10 freeze-thaw cycles. RESULTS: Recoveries were 108% for the PDG assay and 105% for the E1C assay. Serially diluted specimens exhibited parallelism with calibration curves in both assays. Inter- and intraassay CVs were <11%. Urinary and serum concentrations were highly correlated: r = 0.93 for E1C-estradiol; r = 0.98 for PDG-progesterone. All Bangladeshi and US specimens were above detection limits (PDG, 21 nmol/L; E1C, 0.27 nmol/L). Bangladeshi women had lower follicular phase PDG and lower luteal phase PDG and E1Cs than US women. Stability experiments showed a maximum decrease in concentration for each metabolite of <4% per day at room temperature and no significant decrease associated with number of freeze-thaw cycles. CONCLUSIONS: These enzyme immunoassays can be used for the field conditions and population variation in hormone metabolite concentrations encountered in cross-cultural research.

Longitudinal analysis of deciduous tooth emergence in Indonesian children. I. Life table methodology.

Deciduous tooth emergence was investigated in a sample of 468 children from central Java. The subjects were examined every 35 days for up to 2 years. Statistics on the timing of emergence for each tooth were derived from actuarial life table estimates, and male and female survivorship curves of tooth emergence were compared for statistical sex differences. Javanese children exhibited delayed emergence compared to other populations. No overall pattern of sexual dimorphism was detected, although upper first molars emerged significantly earlier in females. A comparison of emergence sequence polymorphisms between Javanese, Finnish, and Japanese children revealed that the i1-i2-m1-c-m2 sequence is most common in all three populations, but each shows a uniquely higher proportion of one or more less common polymorphisms.