New insights from bone microanatomy of the Late Triassic Hyperodapedon (Archosauromorpha,Rhynchosauria): implications for archosauromorph growth strategy

Bone microanatomy of multiple postcranial skeletal elements of several individuals of Hyperodapedon collected from India is reported. This reveals that fibrolamellar bone tissue is predominant in the mid‐ and inner cortices, whereas the peripheral region of the cortex is composed of either parallel‐fibred and/or lamellar bone. The pattern of primary osteons mostly ranges between laminar and subplexiform. Such predominance of fibrolamellar bone tissue in the cortex suggests an overall fast growth, which slowed down considerably later in ontogeny. Four distinct ontogenetic stages are identified based on the bone microstructure. During the juvenile stage, growth was fast and continuous, but it became punctuated during the early and late sub‐adult stages. In adult individuals, growth was slow and showed periodic interruption but did not stop completely, suggesting that Hyperodapedon had an indeterminate growth strategy. Interelemental histovariations affecting cortical thickness, organization of the vascular network, incidence of growth rings and extent of secondary reconstruction are noted. Throughout ontogeny, the femora show higher cortical thickness than humeri and tibiae, suggesting differential appositional growth rate between the skeletal elements. Differences in cortical thickness are noted in the ribs, which suggest differential functional constraints based on anatomical site‐specific occurrences. Although fibrolamellar bone tissue became progressively more dominant towards the archosaurs, there are considerable variations in the growth patterns of the archosauromorphs. This is exemplified by the bone microstructure of Hyperodapedon, which deviates from the generalized slow‐growth pattern proposed for all basal archosauromorphs, suggesting that rapid growth was already present in the archosauromorphs. The cortical thickness of various long bones of Hyperodapedon bears similarity with that of several extant terrestrial quadrupeds, suggesting that Hyperodapedon was essentially a terrestrial quadruped.     

Skeletal Growth Marks and Testis Lobulation as Criteria for Age in Triturus spp. (Amphibia) in Central Norway

Two methods of age determination in newts. Triturus vulgaris and T. cristatus, have been examined and compared: (1) Counting growth rings in the bone tissue of humerus or femur is an accurate method, but possible supplementary rings may make the reading difficult. The innermost periosteal zone corresponds to the first year of life. Later, new zones are added each year. (2) Every second year after maturing a new lobe is developed on the testes. Knowing the juvenile life span in a population, it is possible to determine the age of male newts with a possible error of one year. The correlation between growth marks and testis lobes in T. vulgaris and T. cristatus is highly significant. The testis lobing method, however, is less applicable in older newts, since no more than three or four lobes can be developed on a testis.     

Aspects of reproductive biology of Labeo horie Heckel (Pisces: Cyprinidae) in Lake Chamo, Ethiopia

Labeo horie Heckel is increasingly becoming commercially important in Lake Chamo but its reproductive biology in the lake is not well enough to guide its management. Sex ratio, breeding season, length at first maturity and fecundity of L. horie were studied from 1197 fish samples collected between August 1998 and October 1999 in Lake Chamo, Ethiopia. The sex ratio was significantly different (χ ² = 12.12, P < 0.001). The peak‐breeding period was during the rainy months of March to May, during which time more than 90% of both sexes had ripe gonads. The size at maturity ( L _m50 ) of males was 52 cm while the L _m50 of females was 62 cm. The smallest ripe male in the catch was 46.7 cm and weighed 890 g while the smallest ripe female caught was 49.5 cm and weighed 1145 g. The weight of ripe ovaries ranged from 54.3 g to 991.8 g and contained between 68,400 and 1,320,400 eggs. Relative fecundity ranged between 60 and 290 eggs per g of body weight. The relationships between fecundity and total length and between fecundity and total weight were curvilinear, while the relationship between fecundity and ovary weight was linear. L. horie conformed to the general pattern of reproduction in a tropical environment where peak‐breeding activity occurred during the rainy season. In order to protect the spawning population, fishing pressure should be minimized during breeding time at the shallower littoral regions.     

Skeletal development in Pan paniscus with comparisons to Pan troglodytes

Fusion of skeletal elements provides markers for timing of growth and is one component of a chimpanzee's physical development. Epiphyseal closure defines bone growth and signals a mature skeleton. Most of what we know about timing of development in chimpanzees derives from dental studies on Pan troglodytes. Much less is known about the sister species, Pan paniscus, with few in captivity and a wild range restricted to central Africa. Here, we report on the timing of skeletal fusion for female captive P. paniscus (n = 5) whose known ages range from 0.83 to age 11.68 years. Observations on the skeletons were made after the individuals were dissected and bones cleaned. Comparisons with 10 female captive P. troglodytes confirm a generally uniform pattern in the sequence of skeletal fusion in the two captive species. We also compared the P. paniscus to a sample of three unknown‐aged female wild P. paniscus, and 10 female wild P. troglodytes of known age from the Taï National Park, Côte d'Ivoire. The sequence of teeth emergence to bone fusion is generally consistent between the two species, with slight variations in late juvenile and subadult stages. The direct‐age comparisons show that skeletal growth in captive P. paniscus is accelerated compared with both captive and wild P. troglodytes populations. The skeletal data combined with dental stages have implications for estimating the life stage of immature skeletal materials of wild P. paniscus and for more broadly comparing the skeletal growth rates among captive and wild chimpanzees (Pan), Homo sapiens, and fossil hominins. Am J Phys Anthropol 2012. © 2012 Wiley Periodicals, Inc.     

Assessing the utility of direct and indirect methods for estimating tropical tree age in the Western Ghats, India

The knowledge of tree age is important for understanding tree growth and forest dynamics. It may be estimated by ‘direct’ methods involving growth ring counts, or by ‘indirect’ methods involving field measurements of growth rates. Direct methods are considered more accurate, but it is not clear if they are appropriate for all species, notably from the humid tropics. In this paper we assess the occurrence of annual growth rings and their utility for age estimation in three tropical tree species, Acrocarpus fraxinifolius, Dalbergia latifolia (Fabaceae) and Syzygium cumini (Myrtaceae), growing in traditional shade coffee plantations of the southern Western Ghats, India. These species previously were described as having “indistinct or absent” growth rings. We used anatomical studies, field measurements and computational methods to characterise growth rings and assess similarities between directly and indirectly estimated tree ages. Our study revealed that annual growth rings were characterised by different sets of anatomical features per species and were most distinct in the fast-growing deciduous A. fraxinifolius. Growth rates measured in the field showed annual periodicity in all three species, and reflected annual rainfall-drought cycles in D. latifolia and S. cumini. Direct age estimates were most similar to indirect estimates in D. latifolia, and least so in S. cumini. The results of direct age estimation by counting rings are consistent with them being annual in nature in tropical species with distinct and reliable annual growth ring formation. However, for species with poorly defined growth rings, indirect age estimation methods might be more useful.     

The effects of socioeconomic status on endochondral and appositional bone growth,and acquisition of cortical bone in children from 19th century Birmingham,England

Endochondral growth, appositional growth, and acquisition of cortical bone thickness in the femur are investigated in subadult skeletons (N = 43, dental age range birth to 12 years) from the 19^th‐century AD burial site of St. Martin's churchyard, Birmingham, England. Endochondral growth is monitored using diaphyseal femoral length. Appositional growth is monitored using radiographic midshaft mediolateral width and acquisition of cortical bone using combined mediolateral cortical thickness measured at the midshaft from radiographs. The methodology involves plotting these variables against dental age. Growth is compared in children of differing socioeconomic status. Higher and lower status individuals are identified in the assemblage by their burial in brick vaults in the case of the former and in earth‐cut graves in the case of the latter. The relationships between bone dimensions and dental age are described using a polynomial regression procedure, and analysis of regression residuals is used to evaluate differences in bone dimension‐for‐dental age between the two status groups. Results show that lower socioeconomic status individuals had lower cortical thickness‐for‐dental age than those of higher status. This was interpreted as likely reflecting poorer nutrition in the children of lower socioeconomic backgrounds. There was no patterning with respect to socioeconomic status in femur diaphyseal length or midshaft width. The results support the idea that, for skeletal populations, growth in cortical thickness may be a more sensitive indicator of adverse conditions in childhood than growth in bone length or width. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Short-term progressive early diagenesis in density bands of recent corals:Porites Colonies,Mauritius Island,Indian Ocean

Summary The growth history of some recentPorites colonies of Mauritius Island (Indian Ocean) was dated by sclerochronological methods. Couples of high-density and low-density bands represent the annual growth rate of the corals and allow the growth pattern of every year in the corallum to be counted. The growth and structure of the skeletons ofPorites solida andPorites lutea were investigated. Older parts of the aragonitic skeleton in these 10 to 20 year old corals show various secondary microstructures resulting from alterations and thickenings of the elements of the skeleton. The primary needle-like aragonite crystals are absent in older parts of the corallum and blocky aragonitic cements can occur. Pores and primary skeletal elements are overgrown by new microstructures. These microstructures are caused by secondary cementation and exhibit frontal zones (Stirnzonen), zigzag-like and pseudolamellar-structures. The lamellar structures can be compared with similar structures in the exoskeleton of some Rugosa. A very short early diagenesis within the recent corals is responsible for the thickening and alteration of skeletal elements. It occurs only 4 to 5 years after the formation of the skeleton and tends to increase in importance in older parts of the corallum. Nevertheless, there is no proof for any alteration of aragonite to calcite.     

GROWTH PATTERNS OF THRINAXODON LIORHINUS, A NON-MAMMALIAN CYNODONT FROM THE LOWER TRIASSIC OF SOUTH AFRICA

Abstract:  The growth dynamics of the Early Triassic non-mammalian cynodont Thrinaxodon liorhinus were assessed through bone histology. Several limb bones of various sizes were examined, revealing a rapidly deposited, uninterrupted, fibro-lamellar bone tissue. A region of slowly deposited parallel-fibred bone occurs peripherally in most skeletal elements studied, becoming more extensively developed in the larger limb bones. On the basis of the bone histology, it is proposed that Thrinaxodon liorhinus grew rapidly during early ontogeny, and at a slower rate with increasing age, possibly once sexual maturity was reached. Variation in bone tissue patterns at different stages of ontogeny is noted and discussed. Given that growth rings are generally absent from the skeletal elements studied, and that the environment was seasonal, it appears that Thrinaxodon liorhinus growth was unaffected by environmental fluctuations.     

A comparative view on mechanisms and functions of skeletal remodelling in teleost fish, with special emphasis on osteoclasts and their function

Resorption and remodelling of skeletal tissues is required for development and growth, mechanical adaptation, repair, and mineral homeostasis of the vertebrate skeleton. Here we review for the first time the current knowledge about resorption and remodelling of the skeleton in teleost fish, the largest and most diverse group of extant vertebrates. Teleost species are increasingly used in aquaculture and as models in biomedical skeletal research. Thus, detailed knowledge is required to establish the differences and similarities between mammalian and teleost skeletal remodelling, and between distantly related species such as zebrafish (Danio rerio) and medaka (Oryzias latipes). The cellular mechanisms of differentiation and activation of osteoclasts and the functions of teleost skeletal remodelling are described. Several characteristics, related to skeletal remodelling, distinguish teleosts from mammals. These characteristics include (a) the absence of osteocytes in most species; (b) the absence of haematopoietic bone marrow tissue; (c) the abundance of small mononucleated osteoclasts performing non‐lacunar (smooth) bone resorption, in addition to or instead of multinucleated osteoclasts; and (d) a phosphorus‐ rather than calcium‐driven mineral homeostasis (mainly affecting the postcranial dermal skeleton). Furthermore, (e) skeletal resorption is often absent from particular sites, due to sparse or lacking endochondral ossification. Based on the mode of skeletal remodelling in early ontogeny of all teleosts and in later stages of development of teleosts with acellular bone we suggest a link between acellular bone and the predominance of mononucleated osteoclasts, on the one hand, and cellular bone and multinucleated osteoclasts on the other. The evolutionary origin of skeletal remodelling is discussed and whether mononucleated osteoclasts represent an ancestral type of resorbing cells. Revealing the differentiation and activation of teleost skeletal resorbing cells, in the absence of several factors that trigger mammalian osteoclast differentiation, is a current challenge. Understanding which characters of teleost bone remodelling are derived and which characters are conserved should enhance our understanding of the process in fish and may provide insights into alternative pathways of bone remodelling in mammals.     

The potential use of scales for estimating age and growth of Mediterranean albacore (Thunnus alalunga)

The ability to accurately estimate the age of fishes is critical for conducting stock assessments and developing fishery management policies. Scales were collected from albacore, Thunnus alalunga, caught in the Mediterranean Sea during the years 1989–1995 to estimate their age and growth. Ages, which ranged from 1+ to 6+ years, were estimated from the interpretation of the concentric rings on the scales of 473 individuals that ranged in fork length from 55.5 to 89 cm. Males reached a greater size and age than females. The relatively close agreement in the mean lengths at ages estimated by scales and other techniques constituted a preliminary verification of the method. The von Bertalanffy growth model was fitted to mean lengths at estimated ages, resulting in the following growth parameters for the combined sexes: L_∞ = 86 cm, K = 0.4, t_o = ?0.8 years. Parameter estimates were in agreement with what is known about life history of the species in the Mediterranean. Moreover, the growth rates were consistent with length increment observations from five tag returns, which lend support to our working hypothesis that the scale‐rings are annual structures. When the Mediterranean albacore growth parameters were compared with those of Atlantic Ocean albacore using scale age estimates, there were significant differences between the two populations, and Mediterranean albacore remain significantly smaller than Atlantic Ocean albacore.     

The first record of anguine lizards (Anguimorpha,Anguidae) from the early Miocene locality Ulm – Westtangente in Germany

Abstract

The first fossil anguine material from the lower Miocene (MN 2) locality Ulm – Westtangente in Germany is described. The parietal and compound bone of the lower jaw can be attributed to Ophisaurus holeci, previously known only from younger age (MN 3–MN 7). Moreover, the parietal represents the largest parietal of this species. In other disarticulated material, such as frontal, maxilla, dentary and osteoderms, alpha taxonomy is not possible and these elements cannot be allocated at the species level. Despite a limited data source, a phylogenetic analysis was done (16 taxa, 36 characters) producing four equally parsimonious trees. The analysis shows a close relationship of O. holeci and the Eocene Ophisauriscus quadrupes. These two taxa form a monophyletic clade, a sister-clade to Ophisaurus + Anguis. However, more complete skeletal material of O. holeci is needed to support such a statement. We used our phylogenetic analysis to analyze trace character history for one frontal and three parietal characters. The palaeoenvironmental conditions of the locality Ulm – Westtangente bring further support of the previous hypothesis that O. holeci was adapted to environments with high ground water levels – environments around lakes or rivers.     

Chemistry and structure of skeletal growth rings in the black coral Antipathes fiordensis (Cnidaria,Antipatharia)

The skeleton of the black coral Antipathes fiordensis is a laminar structure displaying major and minor growth rings. The skeleton is composed primarily of protein and chitin organized into successive microlayers. Each microlayer is tightly bound to the next by an organic cement that is periodically opaque in section, and occurs as clusters throughout the skeleton. In contrast, the skeletal microlayers themselves are never opaque. The apposition of multiple, opaque cement lines between skeletal layers is primarily responsible for the growth ring pattern.     

The growth of the common two-banded seabream, Diplodus vulgaris (Teleostei, Sparidae), in Canarian waters, estimated by reading otoliths and by back-calculation

The yearly nature of increment formation in the otoliths of 1–9‐year‐old seabream, Diplodus vulgaris (E. Geoffrey Saint‐Hilaire 1817), from the Canary Islands was validated. The marginal increment method showed that the opaque rings were formed in summer, and the translucent rings in winter. The Brody Proportional Hypothesis and the power length–radius relationship used to back‐calculate the growth trajectories of D. vulgaris showed that this growth model could provide reasonable growth estimates in this species. Growth back‐calculation and growth estimates obtained by direct otolith readings were similar. Data on age and size used to estimate the parameters of the von Bertalanffy growth model for D. vulagris from the Canary Islands showed that males and females had similar growth rates.     

Appendicular skeletal morphology in minute salamanders,genus Thorius (amphibia: Plethodontidae): Growth regulation,adult size determination,and natural variation

Patterns of growth and variation of the appendicular skeleton were examined in Thorius, a speciose genus of minute terrestrial plethodontid salamanders from southern Mexico. Observations were based primarily on ontogenetic series of each of five species that collectively span the range of adult body size in the genus; samples of adults of each of seven additional species provided supplemental estimates of the full range of variation of limb skeletal morphology. Limbs are generally reduced, i.e., pedomorphic, in both overall size and development, and they are characterized by a pattern of extreme variation in the composition of the limb skeleton, especially mesopodial elements, both within and between species. Fifteen different combinations of fused carpal or tarsal elements are variably present in the genus, producing at least 18 different overall carpal or tarsal arrangements, many of which occur in no other plethodontid genus. As many as four carpal or tarsal arrangements were observed in single population samples of each of several; five tarsal arrangements were observed in one population of T. minutissimus. Left-right asymmetry of mesopodial arrangement in a given specimen is also common. In contrast, several unique, nonpedomorphic features of the limb skeleton, including ossification of the typically cartilaginous adult mesopodial elements and ontogenetic increase in the degree of ossification of long bones, are characteristic of all species and distinguish Thorius from most related genera. They form part of a mechanism of determinate skeletal growth that restricts skeletal growth after sexual maturity. Interspecific differences in the timing of the processes of appendicular skeletal maturation relative to body size are well correlated with interspecific differences in mean adult size and size at sexual maturity, suggesting that shifts in the timing of skeletal maturation provide a mechanism of achieving adult size differentiation among species. Processes of skeletal maturation that confer determinate skeletal growth in Thorius are analogous to those typical of most amniotes – both groups exhibit ontogenetic reduction and eventual disappearance of the complex of stratified layers of proliferating and maturing cartilage in long bone epiphyses – but, unlike most amniotes, Thorius lacks secondary ossification centers. Thus, the presence of secondary ossification centers cannot be used as a criterion for establishing determinate skeletal growth in all vertebrates.     

Ontogenetic Patterning of Skeletal Fluctuating Asymmetry in Rhesus Macaques and Humans: Evolutionary and Developmental Implications

I address the question of how fluctuating asymmetry (FA)—the distribution of random deviations from bilateral symmetry—varies ontogenetically in the mammalian skeleton. This question is significant because of the light that such patterns can shed on the causes of variation in developmental stability in bone as well as other structures. Based on large ontogenetic skeletal series of Macaca mulatta and Homo sapiens, I report that the FA variances of skeletal metric traits increase ontogenetically. Coupled with the finding that FA variances also accumulate to greater magnitudes in slower growing mammals, this result is consistent with the hypotheses that FA in bone is primarily caused by (a) cumulative effects of asymmetrical mechanical factors, (b) accumulation of variation in the (local) regulation of growth, or (c) a tendency for bone morphology to drift through undirected remodeling. I discuss the implications of these optional explanations for primate evolution and bone development.     

Shaping Skeletal Growth by Modular Regulatory Elements in the Bmp5 Gene

Cartilage and bone are formed into a remarkable range of shapes and sizes that underlie many anatomical adaptations to different lifestyles in vertebrates. Although the morphological blueprints for individual cartilage and bony structures must somehow be encoded in the genome, we currently know little about the detailed genomic mechanisms that direct precise growth patterns for particular bones. We have carried out large-scale enhancer surveys to identify the regulatory architecture controlling developmental expression of the mouse Bmp5 gene, which encodes a secreted signaling molecule required for normal morphology of specific skeletal features. Although Bmp5 is expressed in many skeletal precursors, different enhancers control expression in individual bones. Remarkably, we show here that different enhancers also exist for highly restricted spatial subdomains along the surface of individual skeletal structures, including ribs and nasal cartilages. Transgenic, null, and regulatory mutations confirm that these anatomy-specific sequences are sufficient to trigger local changes in skeletal morphology and are required for establishing normal growth rates on separate bone surfaces. Our findings suggest that individual bones are composite structures whose detailed growth patterns are built from many smaller lineage and gene expression domains. Individual enhancers in BMP genes provide a genomic mechanism for controlling precise growth domains in particular cartilages and bones, making it possible to separately regulate skeletal anatomy at highly specific locations in the body.     

New information on age and growth in length of Micromesistius australis,Norman 1937 (Pisces,Gadidae), in the South-West Atlantic

Summary Sagittal otoliths were used for age determination in 1121 specimens of Micromesistius australis, caught in waters of the South-West Atlantic during the cruises made by the ships Walther Herwig and Shinkai Maru in agreement with the Argentine Republic, during the years 1978 and 1978/1979, respectively. The interpretation of growth rings required the otoliths to undergo a paraffin wax embedding and cutting technique. The relations otolith length-fish length and fish length-half otolith width were linear. Thus, back calculations, when necessary, were made by a simple extrapolation. Numbers of males and females were nearly equal, resulting in a sex ratio of 1:1.09. Length frequencies and age distributions were calculated for each sex separately. A significant difference in growth between sexes was found, curves for each were derived. A linear, regression was made and the Von Bertalanffy equation estimated as: Lt=56.90 [1-e^{-0.1983 (t+1.84)}], in males, and Lt=60.08 [1-e^{-0.1769 (t+2.07)}] in females, with Lt in cm and t in years. For estimation of these equation parameters age groups 2–16 years in males and 2–22 years in females were taken into account. All theoretical Lt values and almost all observed lengths for females were greater than those of males in corresponding age groups. Previously published observations about M. australis on this subject, are not in agreement or only in partial accord with present results. These are discussed. Original photographs of otolith cross sections illustrating age determination methods are presented for the first time in age and growth studies of this species.     

A requirement for Fgfr2 in middle ear development

The skeletal structure of the mammalian middle ear, which is composed of three endochondral ossicles suspended within a membranous air‐filled capsule, plays a critical role in conducting sound. Gene mutations that alter skeletal development in the middle ear result in auditory impairment. Mutations in fibroblast growth factor receptor 2 (FGFR2), an important regulator of endochondral and intramembranous bone formation, cause a spectrum of congenital skeletal disorders featuring conductive hearing loss. Although the middle ear malformations in multiple FGFR2 gain‐of‐function disorders are clinically characterized, those in the FGFR2 loss‐of‐function disorder lacrimo‐auriculo‐dento‐digital (LADD) syndrome are relatively undescribed. To better understand conductive hearing loss in LADD, we examined the middle ear skeleton of mice with conditional loss of Fgfr2. We find that decreased auditory function in Fgfr2 mutant mice correlates with hypoplasia of the auditory bulla and ectopic bone growth at sites of tendon/ligament attachment. We show that ectopic bone associated with the intra‐articular ligaments of the incudomalleal joint is derived from Scx‐expressing cells and preceded by decreased expression of the joint progenitor marker Gdf5. Together, these results identify a role for Fgfr2 in development of the middle ear skeletal tissues and suggest potential causes for conductive hearing loss in LADD syndrome.     

Age and growth of damselfish Chromis notata (Temminck & Schlegel, 1843), Jeju Island,Korea

This study investigated the age and growth of damselfish, Chromis notate, from Jeju Island in Korea. Samples were collected monthly by lift net from September 2013 to August 2014. Total lengths of the damselfish ranged from 6.4 to 15.3 cm. The relationship between total length and wet weight was WW = 0.0125TL^3.1631 for females, and WW = 0.0091TL^3.2769 for males. The slopes in the relationship between length and weight were not significantly different between sexes, but were significantly different in the intercepts. There were more female than male specimens (1.3:1). Age determination was conducted using the otoliths. Marginal increment (MI) declined in summer and winter, which suggests that two rings are formed each year. Ages of sampled individuals ranged from 1 to 5 years. Length‐at‐age data were fitted using the von Bertalanffy growth model. The estimated growth functions were L_t = 19.93 [1 ? exp^{?0.21 (t + 0.811)}] total length and wet weight was females, and L_t = 16.47 [1 ? exp^{?0.32 (t + 0.499)}] for males.     

Skeletal growth in school children: maturation and bone mass

Skeletal growth and development was evaluated in 322 white children (age 6 to 14) using three different methods: (1) ¹²⁵I photon absorptiometry, (2) compact bone measures on radiographs, and (3) Greulich-Pyle skeletal age from hand-wrist radiographs. Bone mineral content, measured by photon absorptiometry, increased at an incremental rate of about 8.5% each year. Skeletal age was a poor predictor of skeletal status, i.e., bone mineral content (14% error), and did not decrease the predictive error substantially more than did chronological age. Gross morphology (height and weight) was in fact a better predictor of bone mineral content than were skeletal age, chronological age, and radiographic morphometry. Skeletal age deviations were correlated with deviations in body size. A bone mineral index was devised which was independent of body size and this index was also independent of skeletal age. Skeletal age is imprecise (3 to 6 months error) and the range of variation in normal children (13 months) overlaps the maturational delay of the malnourished and diseased. The difficulties in using skeletal maturation are discussed and it is suggested that particular maturational indices be used which better indicate skeletal growth than does a composite skeletal age.