Diversity among African pygmies

Although dissimilarities in cranial and post-cranial morphology among African pygmies groups have been recognized, comparative studies on skull morphology usually pull all pygmies together assuming that morphological characters are similar among them and different with respect to other populations. The main aim of this study is to compare cranial morphology between African pygmies and non-pygmies populations from Equatorial Africa derived from both the Eastern and the Western regions in order to test if the greatest morphological difference is obtained in the comparison between pygmies and non-pygmies. Thirty three-dimensional (3D) landmarks registered with Microscribe in four cranial samples (Western and Eastern pygmies and non-pygmies) were obtained. Multivariate analysis (generalized Procrustes analysis, Mahalanobis distances, multivariate regression) and complementary dimensions of size were evaluated with ANOVA and post hoc LSD. Results suggest that important cranial shape differentiation does occur between pygmies and non-pygmies but also between Eastern and Western populations and that size changes and allometries do not affect similarly Eastern and Western pygmies. Therefore, our findings raise serious doubt about the fact to consider African pygmies as a homogenous group in studies on skull morphology. Differences in cranial morphology among pygmies would suggest differentiation after divergence. Although not directly related to skull differentiation, the diversity among pygmies would probably suggest that the process responsible for reduced stature occurred after the split of the ancestors of modern Eastern and Western pygmies.     

Allometry and adaptation of body proportions and stature in African pygmies

We have analyzed the growth allometry of external body proportions in Efe pygmies from Zaire and combined these data with values from the literature for comparable dimensions in adult pygmies and nonpygmies. We sequentially tested the hypotheses that adult proportion differences between 1) male vs. female Efe, and 2) pygmies vs. nonpygmies result from ontogenetic scaling, or the differential extension of common patterns of growth allometry. Results indicate an almost complete concordance of allometric trajectories for male and female Efe. These preliminary analyses also strongly suggest that adult nonpygmy Africans generally differ from pygmies in their terminal size and correlated allometric consequences, rather than in more fundamental alterations of underlying patterns of growth. Biacromial diameter emerges as the measurement most likely to depart from this general pattern. These results provide further evidence that shifts in systemic growth hormones yielding differences in terminal overall body size may be accompanied by global and coordinated allometric transformations. Certain proportion differences previously interpreted by some as specific evidence of primitive retention in pygmies in fact reflect simple growth allometric correlates of the derived rapid size decrease in these groups. Selected divergent body proportions characterizing adult pygmies, previously interpreted by some as independent evidence of climatic adaptation, also reflect such allometric correlates of ontogenetic scaling. We critically assess arguments that the small overall body size of pygmies was specifically selected for reasons of thermoregulatory efficiency, and consider an alternative or complementary scenario, based on selection for small size in order to reduce caloric requirements. © 1996 Wiley-Liss, Inc.     

Energetics of foraging and locomotion in the platypus Ornithorhynchus anatinus

We measured the energy requirements of platypuses foraging, diving and resting in a swim tank using flow-through respirometry. Also, walking metabolic rates were obtained from platypuses walking on a conventional treadmill. Energy requirements while foraging were found to depend on water temperature, body weight and dive duration and averaged 8.48 W kg(-1). Rates for subsurface swimming averaged 6.71 W kg(-1). Minimal cost of transport for subsurface swimming platypuses was 1.85 J N(-1)m(-1) at a speed of 0.4 m s(-1). Aerobic dive limit of the platypus amounted to 59 s. Metabolic rate of platypuses resting on the water surface was minimal with 3.91 W kg(-1) while minimal RMR on land was 2.08 W kg(-1). The metabolic rate for walking was 8.80 W kg(-1) and 10.56 W kg(-1) at speeds of 0.2 m s(-1) and 0.3 m s(-1), respectively. A formula was derived, which allows prediction of power requirements of platypuses in the wild from measurements of body weight, dive duration and water temperature. Platypuses were found to expend energy at only half the rate of semiaquatic eutherians of comparable body sizes during both walking and diving. However, costs of transport at optimal speed were in line with findings for eutherians. These patterns suggest that underwater locomotion of semiaquatic mammals have converged on very similar efficiencies despite differences in phylogeny and locomotor mode.     

Short stature in African pygmies is not explained by sexual selection

African pygmies' short stature has been studied for more than a century, but the evolution of this extreme phenotype remains unknown. The present study tests the hypothesis that sexual selection, through preference for short partners, may have contributed to the evolution of pygmies' stature. We gathered anthropometric and familial data from 72 Baka pygmy couples and 27 neighboring Nzimé nonpygmy couples from Cameroon. We found evidence for positive assortative mating and partial evidence for the male-taller norm in both groups. This is surprisingly close to results reported for many modern occidental populations, in which sexual selection is thought to exert a positive selective pressure on men height. Semistructured interviews of Baka pygmies concerning height and mate choice suggested that the male-taller norm matches mating preferences. Stature was also positively correlated with the number of serial marriages contracted by men of both populations, while the stature of women was not related to their mating success. Finally, we did not detect any linear or quadratic effect of height on reproductive success for either men or women. Altogether, our results demonstrate that stature influences mate choice in pygmies, and we argue that, if of any influence for sexual selection, mate choice should have favored tallness rather than shortness in our pygmy population. Consequently, this study establishes that sexual selection is a very unlikely candidate to account for the evolution of pygmies' short stature.     

Energetics of East African Pycnonotids1

Rate of oxygen consumption was measured in five bulbuls (Family Pycnonotidae) from western Uganda to evaluate whether this group is indeed characterized by the very low basal rates of metabolism previously reported. For three of these species, body temperature and rate of metabolism were measured as a function of ambient temperature from 10°C to 35°C. In these species body temperature was highly variable, and declined with ambient temperature in Andropadus virens. Such variation, in conjunction with behavioral adjustments, may reduce heat loss at low ambient temperatures. Body mass accounted for 98 percent of the variation in the basal rates of metabolism presented here. Basal rates in these species ranged from 81 to 90 percent of values predicted by the Aschoff–Pohl relationship for passerines, whereas previous measurements ranged from 56 to 72 percent of predicted values. This difference may reflect differences in species or measurement techniques, which, if the latter, suggests that the reduction in metabolic rate in this family may be less than originally thought. These data underline the importance of continued data collection on the metabolism of tropical birds, few of which have been measured to date.     

Studies on African pygmies. V. Red cell acid phosphatase polymorphism in Babinga pygmies: high frequency of ACPR allele.

A group of Babinga Pygmies from the Central African Republic have been analyzed for the acid phosphatase polymorphism with special reference to the ACPR allele. The frequency of this allele (17%) is one of the highest observed in Africa and is comparable only with those reported for the Khoikhoi and the San.     

Energetics of actively powered locomotion using the simplest walking model

We modified an irreducibly simple model of passive dynamic walking to walk on level ground, and used it to study the energetics of walking and the preferred relationship between speed and step length in humans. Powered walking was explored using an impulse applied at toe-off immediately before heel strike, and a torque applied on the stance leg. Although both methods can supply energy through mechanical work on the center of mass, the toe-off impulse is four times less costly because it decreases the collision loss at heel strike. We also studied the use of a hip torque on the swing leg that tunes its frequency but adds no propulsive energy to gait. This spring-like actuation can further reduce the collision loss at heel strike, improving walking energetics. An idealized model yields a set of simple power laws relating the toe-off impulses and effective spring constant to the speed and step length of the corresponding gait. Simulations incorporating nonlinear equations of motion and more realistic inertial parameters show that these power laws apply to more complex models as well.     

Fast locomotion of some African ungulates

Ten species of ungulate were filmed, galloping in their natural habitat. They ranged in size from Thomson's gazelle (about 20 kg) to giraffe (about 1000 kg). They were pursued to make them run as fast as possible. The films have been analysed to determine speed, stride frequency, stride and step lengths, and duty factors. The dependence of these quantities on body size is discussed.  

Summary:

Fast locomotion of zebra, giraffe, warthog and seven species of Bovidae has been studied. The animals were filmed from a pursuing vehicle while galloping in their natural habitat.
Stride frequency was more closely correlated with limb length (represented by hip height) than with body mass. Mean stride frequency was proportional to (hip height)^-0·51 and maximum stride frequency to (hip height) ^-0·63.
Maximum speed was between 10 and 14 m s ^-1 for all species except buffalo (7 m s ^-1). It was not significantly correlated with body mass.
Since the small species ran at least as fast as the large ones they attained higher Froude numbers. Relative stride length was approximately 1·8 (Froude number)^0·39 for all species, irrespective of size. Relative step length was approximately 0·65 (Froude number)^0·2, both for the fore feet and for the hind ones. The vertical forces exerted by the feet are proportional to (body weight)×(Froude number)^0·2 so the forces at maximum speed are larger multiples of body weight for small species than for large ones.     

Energetics of locomotion and load carriage in the nectar feeding ant, Camponotus rufipes

Abstract.  To investigate if there is an energetic constraint influencing a nectar feeding ant's decision to come back to the nest with partial loads, the energetic costs of running and carrying a load in the ant Camponotus rufipes are measured. Metabolic rates of individuals are measured in a running tube respirometer while they are unladen and laden at 25 °C. Workers voluntarily collect a load of 6 µL of a 30% sucrose solution (mass = 6.8 mg), which results in an internal load of about 50% of the ant mass and is close to a full load for ants within this size range. The gross cost of unladen running is 264 J kg⁻¹ m⁻¹, while that of laden running is 225 J kg⁻¹ m⁻¹. The mass used to calculate the cost of laden running includes body mass of ant and load carried. Load carriage cost in C. rufipes foragers is calculated to be about 60% as much as body carriage per unit mass. Internal load carriage in C. rufipes is energetically cheaper compared with external carriage in other ant species. Such low carriage costs make it unlikely that the collection of partial crop loads in C. rufipes foragers is based on a minimization of foraging costs, as suggested for honeybees.     

Patterns of ancestry, signatures of natural selection, and genetic association with stature in Western African pygmies

African Pygmy groups show a distinctive pattern of phenotypic variation, including short stature, which is thought to reflect past adaptation to a tropical environment. Here, we analyze Illumina 1M SNP array data in three Western Pygmy populations from Cameroon and three neighboring Bantu-speaking agricultural populations with whom they have admixed. We infer genome-wide ancestry, scan for signals of positive selection, and perform targeted genetic association with measured height variation. We identify multiple regions throughout the genome that may have played a role in adaptive evolution, many of which contain loci with roles in growth hormone, insulin, and insulin-like growth factor signaling pathways, as well as immunity and neuroendocrine signaling involved in reproduction and metabolism. The most striking results are found on chromosome 3, which harbors a cluster of selection and association signals between approximately 45 and 60 Mb. This region also includes the positional candidate genes DOCK3, which is known to be associated with height variation in Europeans, and CISH, a negative regulator of cytokine signaling known to inhibit growth hormone-stimulated STAT5 signaling. Finally, pathway analysis for genes near the strongest signals of association with height indicates enrichment for loci involved in insulin and insulin-like growth factor signaling.     

Digital ridge-counts of Efe pygmies

The mean total digital ridge-counts among a sample of Efe of the Ituri forest is 96.4 in males (n = 153) and 96.6 in females (n = 54). These are the lowest values yet recorded and probably represent the lower extreme in the spectrum of ridge-counts of non-pathological populations.     

Capacity for sustained terrestrial locomotion in an insect: Energetics,thermal dependence,and kinematics

Summary The capacity for sustained, terrestrial locomotion in the cockroach. Blaberus discoidalis, was determined in relation to running speed, metabolic cost, aerobic capacity, and ambient temperature (T _a=15, 23, and 34°C; acclimation temperature=24°C). Steady-state thoracic temperature (T _tss) increased linearly with speed at each T _a.The difference between T _tss and T _awas similar at each experimental temperature with a maximum increase of 7°C. Steady-state oxygen consumption (VO_2ss) increased linearly with speed at each T _aand had a low thermal dependence (Q₁₀=1.0-1.4). The minimum cost of locomotion (the slope of the VO_2ss versus speed function) was independent of T _a.Cockroaches attained a maximal oxygen consumption (VO_2max). increased with T _afrom 2.1 ml O₂·g^-1·h^-1 at 15°C to 4.9 ml O₂·g^-1·h^-1 at 23°C, but showed no further increase at 34°C, VO_2max increased 23-fold over resting VO₂ at 23°C, 10-fold at 34°C, and 15-fold at 15°C. Endurance correlated with the speed at which VO_2max was attained (MAS, maximal aerobic speed). Temperature affected the kinematics of locomotion. compared to cockroaches running at the same speed, but higher temperatures (23–34°C), low temperature (15°C) increased protraction time, reduced stride frequency, and reduced stability by increasing body pitching. The thermal independence of the minimum cost of locomotion (C_min), the low thermal dependence of VO_2ss (i.e., y-intercept of the VO_2ss versus speed function), and a typical Q₁₀ of 2.0 for VO_2max combined to increase MAS and endurance in B. discoidalis when T _awas increased from 15 to 23°C. Exerciserelated endothermy enabled running cockroaches to attain a greater VO_2max, metabolic scope, and endurance capacity at 23°C than would be possible if T _tss remained equal to T _a. The MAS of B. discoidalis was similar to that of other arthropods that use trachea, but was 2-fold greater than ectotherms, such as salamanders, frogs, and crabs of a comparable body mass.Abbreviations T _a ambient temperature - T _t thoracic temperature - T _tss steady state thoracic temperature during exercise - T _trest thoracic temperature during rest - VO₂ oxygen consumption - VO_2rest oxygen consumption during rest - VO_2ss steady-state oxygen consumption during exercise - VO_2max maximal oxygen consumption; MAS maximum aerobic speed - C _min minimum cost of locomotion - t _end endurance time     

Dermatoglyphics of the Efe pygmies.

This paper deals with the dermatoglyphics in a group of Efe Pygmies, especially with the quantitative traits of fingers, as well as with palmar dermatoglyphics. This material was collected by P. Schebesta in Zaire and was partially published already by J.A. Valsík in 1938.     

Nutritional status of Efe pygmies and Lese horticulturists

We assessed the nutritional status of 90 pygmies (Efe) and 211 members of a neighboring horticultural Lese in the Ituri forest in northeast Zaire. Clinical signs of iron and iodine deficiency were highly prevalent, but the frequency of these and other clinical signs of nutritional deficiency did not differ between the two groups. Infectious diseases such as yaws, cellulitis, and filiariasis were common. Absolute heights and weights of both Efe males and females aged 2–18 years were less than the 5th percentile of the National Center for Health Statistics standards. Weights for height of both groups were within the normal range. Weights and heights, expressed as percentages of the 50th percentile for age and sex, were significantly lower (P < .001) in Efe males and females than among Lese males and females, but weights for height did not differ significantly. Z-Scores of height and weight were also significantly lower in Efe males and females than among their Lese counterparts. The differences in weight and height, expressed either as a percentage of the 50th percentile or as Z-scores, were most apparent in older adolescents and persisted among adults. The normal weight for height among both populations indicated that acute undernutrition did not exist at the time of the survey. Despite comparable levels of adult fatness, measured by triceps skinfold thickness, heights of Efe males and females were lower than those of the Lese. These observations support the hypothesis that genetic differences account for the short stature of the Ituri pygmy. However, because the environments shared by the two groups are not identical and because other practices differ, nutritional effects cannot be excluded.     

Beta-globin gene haplotypes and alpha-thalassemia analysis in Babinga pygmies from Congo-Brazzaville

We analyzed beta-globin gene cluster haplotypes and deletional alpha+-thalassemia (-alpha3.7kb) in 54 Babinga pygmy subjects from Congo-Brazzaville. The beta(S)-globin gene frequency was 0.065 and that of the deletional alpha-globin gene (-alpha3.7kb) was 0.29. Eighty-five percent of the beta(S) chromosomes and 13% of the beta(A) chromosomes were associated with the Bantu haplotype, 10% of beta(A) chromosomes with the Senegal haplotype, and the remaining beta chromosomes with atypical haplotypes. None of the chromosomes were of the Benin haplotype. These results are clearly of anthropological and evolutionary interest. They also support earlier observations that alpha+-thalassemia is prevalent at a high frequency in African populations.     

Anomalously low endemic goiter prevalence among Efe pygmies

The existence of endemic goiter, caused by iodine deficiency and the presence of a dietary goitrogen, has been noted in eastern Zaire by a number of authors (De Visscher et al.: Journal of Clinical Endocrinology and Metabolism 21:175-188, 1961; Delange et al.: Journal of Clinical Endocrinology and Metabolism 34:1059, 1972). In the Ituri Forest of Huate-Zaire, two distinct populations, the Efe (pygmy) and Lese (Bantu), live in association with each other and have similar diets. The goiter survey reported here documents differences in goiter prevalence and severity between the nomadic pygmy and village-living Bantu populations. While the Efe have an overall goiter prevalence of 9.4%, the Lese have a goiter prevalence of 42.9%. Furthermore, Efe women living in Lese villages and subsisting on a Lese diet have a prevalence of goiter similar to that of forest-living Efe women. Village-living individuals born of Efe mothers and Lese fathers have a prevalence of goiter greater than that of pure Efe but less than that of Lese. While our data cannot exclude dietary explanations for the difference in goiter prevalence between the Efe and Lese, they do support the hypothesis that the Efe possess an adaptation to an iodine-deficient environment that does not result in the development of goiters.