Modes of ontogenetic allometric shifts in crocodylian vertebrae

During postnatal ontogeny of vertebrates, allometric trends in certain morphological units or dimensions can shift drastically among isometry, positive allometry, and negative allometry. However, detailed patterns of allometric transitions in certain timings have not been explored well. Identifying the presence and nature of allometric shifts is essential for understanding the patterns of changes in relative size and shape and the proximal factors that are controlling these changes mechanistically. Allometric trends in 10 selected vertebrae (cervical 2–caudal 2) from hatchlings to very mature individuals of Alligator mississippiensis (Archosauria, Crocodylia) are reported in the present study. Allometric coefficients in 12 vertebral dimensions are calculated and compared relative to total body length, including centrum, neural spine, transverse process, zygapophysis, and neural pedicle. During the postnatal growth, positive allometry is the most common type of relative change (10 of the 12 dimensions), although the diameter of the neural canal shows a negative allometric trend. However, when using spurious breaks (i.e. allometric trends subdivided into growth stages using certain growth events, and key body sizes and/or ages), vertebral parts exhibit various pathways of allometric shifts. Based on allometric trends in three spurious breaks, separated by the end of endochondral ossification (body length: approximnately 0.9 m), sexual maturity (1.8 m), and the stoppage of body size increase (2.8 m), six types of ontogenetic allometric shifts are established. Allometric shifts exhibit a wide range from positive allometry restricted only in the early postnatal stage (Type I) to life‐long positive allometry (Type VI). This model of ontogenetic allometric shifts is then applied to interpret potential mechanisms (causes) of allometric changes, such as (1) growth itself (when allometric trend gradually decreases to isometric or negative allometric change: Type II–IV allometric shift); (2) developmental constraint (when positive allometry is limited only in the early growth stage: Type I allometric shift); and (3) functional or biomechanical drive (when positive allometry continues throughout ontogeny: Type VI allometric shift).     

Allometric Growth in Four Species of Digenetic Trematodes of Marine Fishes from Belize

Four species of digenetic trematodes of marine fishes from Belize are studied for allometric growth: Multitestis rotundus Sparks, 1954 (Lepocreadiidae), Stenopera equilata Manter, 1933 (Opecoelidae), Leurodera decora Linton, 1910 (Hemiuridae), Prosorhynchus pacificus Manter, 1940 (Bucephalidae). These are compared with six other species for which allometric growth is known. Positive growth is exhibited in all species only by the hindbody, whereas the forebody, suckers, and pharynx are negative in all. The body width, post-testicular body, and gonads are positive in some species and negative in others. At times the body proportion or organ is growing at about the same rate as the body length.     

眼斑双锯鱼仔稚鱼发育异速生长

运用生态学和传统理论生物学的研究方法, 对孵化后眼斑双锯(Amphiprion ocellaris)仔、稚鱼在早期生存和环境适应上的异速生长及器官优先发育生态学意义进行了研究, 以期为眼斑双锯鱼人工繁殖和育苗提供参考资料。以11日龄为眼斑双锯鱼仔、稚鱼的区分时期, 结果表明, 眼斑双锯鱼仔、稚鱼的感觉、摄食和游泳等器官快速分化, 均存在异速生长现象。在头部器官中, 吻长、眼间距、口宽和头高在仔鱼期均为正异速生长, 吻至鳃裂前缘长和眼径为负异速生长。在身体各部位中, 仔鱼期体高、躯干长、尾长、尾柄长、尾柄高和体厚均为正异速生长, 仅头长为负异速生长; 在游泳器官中, 仔鱼期眼斑双锯鱼尾鳍、背鳍、胸鳍、腹鳍和臀鳍均为正异速生长。稚鱼期眼斑双锯鱼头部、躯干及游泳等各器官均为负异速生长。眼斑双锯鱼这些关键器官的异速发育, 对适应环境因子变化具有重要的生态学意义。     

红鳍笛鲷仔、稚鱼异速生长

运用生态学和传统理论生物学的研究方法,对孵化后红鳍笛鲷（Lutjanus erythropterus）仔、稚鱼在早期生存和环境适应上的异速生长及器官优先发育生态学意义进行了研究,以期为红鳍笛鲷人工繁殖、育苗提供参考资料。以17日龄为红鳍笛鲷仔、稚鱼的区分时期,结果表明,红鳍笛鲷仔、稚鱼的感觉、呼吸摄食和游泳等器官快速分化,均存在异速生长现象。在头部器官中,吻长、口宽、眼径和头高在仔鱼期均为正异速生长,稚鱼期吻长为等速生长,口宽、眼径和头高为负异速生长。在身体各部位中,仔鱼期头长和体高为正异速生长,躯干部和尾长为负异速生长;稚鱼期体高和躯干长为正异速生长,头长和尾长为等速生长;在游泳器官中,仔鱼期红鳍笛鲷背鳍、腹鳍、尾鳍为正异速生长,胸鳍为等速生长,稚鱼期臀鳍为正异速生长,腹鳍、胸鳍和尾鳍为等速生长,背鳍为负异速生长。红鳍笛鲷这些关键器官的快速发育,使外源性营养开始后以最小的代谢损耗获得了生存能力的显著提升,对挑战和适应纷繁变换的外界压力具有重要的生态学意义。     

大麻哈鱼卵黄囊期仔鱼异速生长及其生态学意义

运用实验生态学的方法, 对大麻哈鱼(Oncorhynchus keta Walbaum)卵黄囊期仔鱼的异速生长及器官优先发育在早期生存和环境适应上的生态学意义进行了研究。结果表明, 大麻哈鱼卵黄囊期仔鱼的感觉、摄食, 呼吸和游泳等器官快速分化, 许多关键器官均存在异速生长现象。在身体各部分中, 头部和尾部为正异速生长, 躯干部为负异速生长, 体高有先增大后减小的趋势; 在头部器官中, 眼径、口宽、吻长和眼后头长均为正异速生长; 在游泳器官中, 胸鳍、腹鳍、背鳍、臀鳍、背鳍基、臀鳍基和尾鳍均为正异速生长, 脂鳍为负异速生长, 其中, 腹鳍在全长25.31 mm、12日龄出现生长拐点, 但拐点前后均为正异速生长。大麻哈鱼卵黄囊期仔鱼感觉、摄食, 呼吸和游泳等器官的快速发育, 使出膜后的仔鱼在最短的时间内获得了与早期生存密切相关的各种能力, 对适应复杂多变的外界环境具有重要的生态学意义。       

日本七鳃鳗胚胎发育及卵黄囊期仔鱼的异速生长

通过观察日本七鳃鳗Lampetra japonica (Martens, 1868)胚胎外部形态和内部组织结构变化, 描述受精卵从卵裂至器官形成以及仔鱼孵出的发育阶段, 采用实验生态学方法研究卵黄囊期仔鱼的异速生长模式。结果表明: 日本七鳃鳗卵子为乳白色, 呈卵圆形; 受精卵卵裂方式为全卵裂; 胚胎发育过程主要包括卵裂期、囊胚期、原肠胚期、神经胚期、头凸期、孵出前期和孵出期, 历时11—12d; 初孵仔鱼为乳白色, 全长约(3.41±0.24) mm, 体质量约为0.0006 g。日本七鳃鳗胚胎发育研究可为了解七鳃鳗胚胎发育过程, 早期脊椎动物的起源和发育进化研究提供参考。卵黄囊期内仔鱼身体各部分中, 头长和尾长均表现出快速生长, 同在7 日龄出现生长拐点, 且生长拐点后的生长速率都大于生长拐点前的生长速率; 而仔鱼体长在卵黄囊期内表现出慢速生长。在头部器官中, 吻长、鳃前长和鳃长均表现出快速生长现象, 吻长和鳃长分别在9日龄和8日龄出现生长拐点; 口笠长在3日龄时出现生长拐点, 在生长拐点前为等速生长, 而在生长拐点后表现出快速生长; 眼径和眼鳃间距则分别表现出等速生长和慢速生长。泄殖孔在卵黄囊期内未出现生长拐点, 生长速率相对于全长生长速率表现出快速生长现象。七鳃鳗卵黄囊期仔鱼的异速生长是在长期进化过程中, 适应早期生活环境和独特的生活方式而形成特有的发育机制。     

Relative growth and sexual dimorphism in the hermit crab Clibanarius signatus Heller, 1861 (Anomura,Diogenidae) from the northern coast of the Persian Gulf,Iran

The purpose of this study was to investigate the relative growth and sexual dimorphism in the hermit crab Clibanarius signatus. The evaluation was done with 955 specimens (494 males, 251 females, and 210 intersexes) captured in Persian Gulf (Iran) during January to December 2015. Animals were submitted to measurements related to weight (BW, total wet weight) and body size related to cephalic shield (SW, width; and SL, length) and propodus of both chelipeds (CPL, length; and CPW, width). Males were larger and heavier than females and intersexes. Both males and females showed a negative allometric growth for the SL–BW and SL–SW relationships, but a positive allometric growth to intersex specimens. To SL–CPL relationship, a negative allometric growth was confirmed in males and females independent of the laterality of the CPL, whereas a contrast was verified in intersexes, with a positive allometric growth occurred for both hands. To SL–CPW relationship, a negative allometric growth (b < 1) occurred in females, independent of the laterality of the CPW, while in males, a positive allometric pattern was confirmed. In intersexes, this relationship was positive except for the right CPW which was isometric. Sexual dimorphism was evident in Clibanarius signatus, with males being the largest and females the smallest specimens in the population.     

It's all relative: allometry and variation in the baculum (os penis) of the harp seal, Pagophilus groenlandicus (Carnivora: Phocidae)

We compared allometry and variation in the baculum (os penis), mandible, and humerus of the harp seal, Pagophilus groenlandicus. This species is presumed to have a promiscuous mating system in which choice of mate by females during intromission with different males is likely. The baculum is large and grows throughout life so may be an honest indicator of males' quality (size) or viability (age). We predicted that bacular size would exhibit stronger allometry relative to body size than mandibles or humeri. The baculum is less functionally (mechanically) constrained than mandibles or humeri so we also predicted it would be more variable, though less variable than sexually selected traits which do not function as honest indicators. Our sample (N=67 seals) represented broad ranges of size and age (0–35 yr) so we compared variation using residuals from allometric regressions of skeletal measurements on body length. Bacular size was isometric to body length until ∼ 137 cm (when some seals enter puberty) in body length then was highly positively allometric; mandibular and humeral size were negatively allometric to body length throughout growth. Bacula were more variable than mandibles or humeri. Bacular size in large specimens (>137 cm in body length) was related strongly to body length and weakly to age. We interpret bacular size to be an uncheatable honest indicator of male quality and viability. High bacular variation conforms with theoretical predictions of females' asymmetrical choice of mate and choice of extremes, and may reflect corresponding anatomical variation among females. Some bacular variation may also result incidentally from positive allometry coupled with lifelong bacular growth, which can amplify early differences between reproductive and somatic growth, enabled by weak selection on bacular form in relation to function.     

Allometric relations in three species of finches (Aves: Fringillidae)

Static nestling, adult and ontogenetic allometry were analysed in three species of finches. Static nestling allometry was very similar across age in early ontogeny and among species and could be approximated by a single matrix of phenotypic variances and covariances. The first eigenvector of this matrix showed negative allometry of bill and tarsus to mass, but positive for wing length to mass. Adult static allometry was also very similar among species, but differed from nestling pattern. In adults the bill had a positive allometry in relation to tarsus and wing, but negative to mass, while tarsus and wing were unrelated to mass. The ontogenetic allometry in each species was very similar to nestling static allometry. Viewed in relation to final size, bill characters grew more slowly than body characters, but for a longer time, which created the difference between adult and nestling allometric patterns. There were differences among species both with regard to elevation and slope of allometric coefficients, suggesting that the differences among species came about by changes in the three fundamental ontogenetic parameters namely growth rate, onset of growth and offset of growth.     

Allometric strategies for increasing respiratory surface area in the Mississippian blastoid Pentremites

The surface area of blastoid respiratory structures (hydrospires) shows positive allometry during ontogeny to offset the exponential increase in volume. Transverse cross-sections of thecae through an ontogenetic series in two blastoid species, Pentremites pyriformis and Pentremites godoni , were used to calculate surface area and volume within the hydrospires. These two congeneric species showed similar allometric change in hydrospire surface area, but this change was accomplished using different mechanisms. In P. godoni , increased hydrospire surface area was developed through an allometric increase in hydrospire length while keeping hydrospire fold count constant. In contrast, P. pyriformis , showed little allometric change in hydrospire length, but instead increased the number of hydrospire folds ontogenetically. This study shows that developmental patterns can be modified in different ways in order to solve the same functional problem.     

Ontogenetic allometry and body composition of the common bottlenose dolphin (Tursiops truncatus) from the U.S. mid‐Atlantic

Growth in common bottlenose dolphins (Tursiops truncatus) was investigated through examination of sex‐specific, ontogenetic changes in the mass of 38 discrete body compartments, utilizing stranded dolphins in good body condition (n = 145). Ontogenetic allometry and the body composition technique were used to quantitatively describe growth patterns. Although adult males were significantly larger than adult females in total body mass (TBM) and total length, overall patterns of growth were remarkably similar between sexes. The integument, locomotor muscle, and vertebral column together represented 50%–58% of TBM across all life history categories, although their relative contributions varied ontogenetically. Young dolphins invested the greatest percentage of TBM in integument, while locomotor muscle was the single largest body component in adults. In both sexes (1) most muscle groups displayed positive allometry, (2) most skeletal elements displayed negative allometric or isometric growth, (3) most abdominal viscera associated with digestion displayed positive allometry, and (4) the brain displayed negative allometric growth. Reproductive tissues exhibited the highest rates of growth in both sexes, and increased as a percentage of TBM with maturity. This study provides an integrated view of bottlenose dolphin growth and a quantitative baseline of body composition for future monitoring of this sentinel species of ecosystem health.     

A test of Allen's rule in ectotherms: the case of two south American Melanopline Grasshoppers (Orthoptera: Acrididae) with partially overlapping geographic ranges

We studied the geographic variation of three morphometric characters in relation to body size in two South American grasshoppers (Acrididae), Dichroplus vittatus Bruner and D. pratensis Bruner to test Allen's rule in these ectotherms. Since both species follow the converse to Bergmann's rule owing to latitudinal and/or altitudinal variation in time available for growth and reproduction, geographic variation in body size proportions of protruding parts may obey to differential allometric growth in different geographic areas. Alternatively, it could reflect true Allenian variation related to thermoregulation. Body proportions were studied by correlation/regression analyses with geographic and climatic variables. In D. pratensis, body proportions increased with latitude and decreased with altitude. These results probably obey to the effects of water balance and seasonality on final body size, and on the allometric growth of the three studied characters not being related to thermoregulation. In D. vittatus, a generally non-significant trend towards the decrease of the mean proportions of all three characters with increasing latitude was observed. Nevertheless, also in this species, it is probable that the environmental gradient responds to seasonality factors (although not to water balance) that affect the length of growing season and, in consequence, body size and its allometric relationships. We conclude that the regularities in the geographic distribution of body proportions of D. pratensis and D. vittatus do not follow Allen's rule in the sense of thermoregulation, and result from variables that determine growing season length and the allometric growth of different body parts.     

Condition factor,length–weight and length–length relationships of an endangered fish Ompok pabda (Hamilton 1822) (Siluriformes: Siluridae) from the River Gomti,a tributary of the River Ganga,India

The present study describes the condition factor, length‐weight and length‐length relationships for an endangered freshwater fish species Ompok pabda (Hamilton) of the family Siluridae from the River Gomti in Northern India. The values of regression parameter b ranged from 2.81 to 3.32 (r² > 0.90). During the pre‐monsoon, the allometric coefficient b of the LWR was close to isometric value (b = 3.08) allometric (b = 2.87), although it suggested negative allometric growth in monsoon periods while positive growth in post‐monsoon. The condition factor values ranged from 0.672 to 0.744. Results of the present study could be useful to help in conservation and sustainable fisheries management of this endangered species.     

Biology of the annular seabream, Diplodus annularis (Sparidae), in coastal waters of the Canary Islands

The biology of the Canary Islands annular seabream Diplodus annularis (Linnaeus 1758) was studied from samples collected between January and December 1998. Fish ranged from 82 to 209 mm total length in size and from 8.7 to 137.1 g in weight. The mean length showed an increase with increasing water depth. Males showed a negative allometric growth and females isometric growth. The species was characterized by protandric hermaphroditism. The overall sex ratio was unbalanced in favour of males (1 : 0.79). The reproductive season extended from January to May, with a peak in spawning activity in March–April. Males reached maturity at a smaller length (103 mm, 1-year-old) than females (128 mm, 2-year-old). Fish aged 0–6 years were found. The von Bertalanffy growth parameters for all individuals were: L_∞=247.9 mm, k=0.268 years^–1, and t₀=–0.879 years.     

Length–weight and length–length relationships of 10 small fish species from the Ganges, Bangladesh

This study describes the length–weight (LWR) and length–length (LLR) relationships for ten small indigenous fish species from the lower part of the Ganges, Bangladesh, namely Ailia coila , Amblypharyngodon mola , Aspidoparia morar , Clupisoma atherinoides , Eutropiichthyes vacha , Glossogobius giuris , Gudusia chapra , Lepidocephalus guntea , Mystus vittatus , and Puntius ticto . A total of 2142 specimens, representing 10 species of 5 families used for this study were caught by traditional fishing gear from March 2006 to February 2007. Standard length (SL), total length (TL) and fork length (FL) for each specimen were measured by digital slide calipers and each body weight (BW) was taken by a digital balance. The allometric coefficient b of the LWR was close to the isometric value ( b  =   3.001) in G. giuris , although it suggested negative allometric growth in A. coila, A. morar, C. atherinoides, E. vacha, and P. ticto, whilst positive allometric growth in rest of the species. The results further indicated that the LLRs were highly correlated ( r ²  >   0.890; P < 0.01). To the best our knowledge, this study presented the first reference on LWR and LLR for most of the species in Bangladesh.     

Growth, development and allometry of Philophthalmus nocturnus in the eyes of domestic chicks.

Fifty day-old chicks were each infected with 10 excysted metacercariae of Philophthalmus nocturnus Looss, 1907 around each orbit and growth, development and allometry were studied. The growth rate showed two phases over a period of 35 days, a limited lag phase lasting two days post-infection in which flukes did not exceed 440 microns in length, and a rapid phase during which growth was rapid and flukes reached a size of 3.008-3.504 mm on day 35. Five developmental stages were noticed during the course of development of the metacercaria to the egg-producing adult stage. Eggs appeared in the uterus on day 14 and oculate miracidia on day 25. The hindbody, testes and ovary showed positive allometric growth, the pharnyx less so, whereas negative allometric growth was shown by the forebody. Body width, oral sucker and ventral sucker were close to isometry, growing at the same rate as the body length.     

The relation between pituitary gland and thyroid growth during the lifespan of the annual fish Cynolebias whitei and Nothobranchius korthausae: gonadotropic and thyrotropic cells

Summary In the annual cyprinodont Cynolebias whitei the cell types responsible for the increase of pituitary growth at the onset of maturation and for pituitary hyperplasia in old specimens were identified as gonadotropic cells and thyrotropic cells, respectively. The gonadotropic cells showed a high affinity to anti-carp -GTH serum, both at light- and electron-microscopical levels. The allometric relation of total gonadotropic cell volume to body length, determined for fish from six weeks up to six months of age, showed no inflections. Therefore pituitary growth in maturing fish may be partly a result of proliferation of gonadotropes, although gonadotropic cells do not contribute to pituitary hyperplasia in old fish. Thyrotropic cells showed a weak affinity to anti-carp -GTH serum at light-microscopical level. Under the electron microscope thyrotropic cells displayed signs of activation in maturing fish and signs of proliferation in old fish. The allometric relation of thyroid gland volume to body length paralleled that of pituitary volume to body length. Histologically the thyroid gland showed signs of inactivity in adult fish and of hyperplasia in old fish. The possibility, that gonadal maturation, pituitary thyrotropic activity, and growth of the thyroid in maturing fish are related through the inhibitory action of gonadal steroids on thyroid hormone release, is discussed. Pituitary hyperplasia in old fish is the result of proliferation of thyrotropic cells. Similar hyperplasia of pituiary and thyroid glands was observed in old Nothobranchius korthausae.     

斑鰶仔、稚鱼形态发育及异速生长模式研究

运用光学显微镜技术和实验生态学方法, 对斑鰶(Konosirus punctatus)早期形态发育观察、异速生长模式及其生态学意义进行了研究。结果表明: 在水温(21.5±0.5)℃下, 初孵仔鱼全长(3.18±0.52) mm, 斑鰶仔鱼期从孵化出膜到43日龄棱鳞开始出现前, 稚鱼期从44日龄棱鳞出现到55日龄全身覆满鳞片。斑鰶的形态变化和器官分化主要发生在仔鱼期。斑鰶的吻长、躯干长、肠道长、胸鳍长、腹鳍长等重要形态学指标均存在异速生长现象, 其生长拐点依次为 42日龄(TL: 26.41 mm)、24日龄(TL: 15.57 mm)、31日龄(TL: 21.41 mm)、41日龄(TL: 25.47 mm)、42日龄(TL: 26.41 mm)。相对于全长、吻长和胸鳍长在拐点前后由正异速生长变为等速生长, 腹鳍长由正异速生长转为负异速生长, 这为呼吸、摄食和成功逃避捕食者提供有利条件; 而肠道长由负异速生长变为等速生长, 这可能与斑鰶的食性转化有关。综上所述, 为适应复杂多变的生存环境, 斑鰶在早期发育阶段优先发育对生长生存起关键作用的器官, 这对提高仔、稚鱼的存活率具有重要的生态学意义。研究将为今后进一步人工繁育和苗种培育提供理论依据。     

Negative allometry of docosahexaenoic acid in the fowl lung and pulmonary surfactant phospholipids

In a recent study (Comp. Biochem. Physiol. B. (2010)155: 301-308) we reported that the fatty acids (FA) of the avian (7 species) total lung phospholipids (PL) (i.e. lung parenchyma and surfactant together) provide allometric properties. To test whether this allometric scaling also occurs in either of the above components, in six gallinaceous species, in a body weight range from 150 g (Japanese quail, Coturnix coturnix japonica) to 19 kg (turkey, Meleagris gallopavo) the PL FA composition (mol%) was determined in the pulmonary surfactant, in native and in thoroughly lavaged lungs (referred to as lung parenchyma). In all three components docosahexaenoic acid (DHA) showed significant and negative allometric scaling (B = -0.056, -0.17 and -0.1, respectively). Surfactant PLs provided further negative allometry for palmitic acid and the opposite was found for palmitoleate and arachidonate. In the lung parenchymal PLs increasing body weight was matched with shorter chain FAs (average FA chain length) and competing n6 and n3 end-product fatty acids (positive allometry for arachidonic acid and negative for DHA). Negative allometric scaling was found for the tissue malondialdehyde concentration in the native and lavaged lungs (B = -0.1582 and -0.1594, respectively). In these tissues strong correlation was found between the MDA concentration and DHA proportion (r = 0.439 and 0.679, respectively), denoting the role of DHA in shaping the allometric properties and influencing the extent of in vivo lipid peroxidation of membrane lipids in fowl lungs.     

Scaling of the appendicular skeleton of the giraffe (Giraffa camelopardalis)

Giraffes have remarkably long and slender limb bones, but it is unknown how they grow with regard to body mass, sex, and neck length. In this study, we measured the length, mediolateral (ML) diameter, craniocaudal (CC) diameter and circumference of the humerus, radius, metacarpus, femur, tibia, and metatarsus in 10 fetuses, 21 females, and 23 males of known body masses. Allometric exponents were determined and compared. We found the average bone length increased from 340 ± 50 mm at birth to 700 ± 120 mm at maturity, while average diameters increased from 30 ± 3 to 70 ± 11 mm. Fetal bones increased with positive allometry in length (relative to body mass) and in diameter (relative to body mass and length). In postnatal giraffes bone lengths and diameters increased iso‐ or negatively allometric relative to increases in body mass, except for the humerus CC diameter which increased with positive allometry. Humerus circumference also increased with positive allometry, that of the radius and tibia isometrically and the femur and metapodials with negative allometry. Relative to increases in bone length, both the humerus and femur widened with positive allometry. In the distal limb bones, ML diameters increased isometrically (radius, metacarpus) or positively allometric (tibia, metatarsus) while the corresponding CC widths increased with negative allometry and isometrically, respectively. Except for the humerus and femur, exponents were not significantly different between corresponding front and hind limb segments. We concluded that the patterns of bone growth in males and females are identical. In fetuses, the growth of the appendicular skeleton is faster than it is after birth which is a pattern opposite to that reported for the neck. Allometric exponents seemed unremarkable compared to the few species described previously, and pointed to the importance of neck elongation rather than leg elongation during evolution. Nevertheless, the front limb bones and especially the humerus may show adaptation to behaviors such as drinking posture. J. Morphol. 276:503–516, 2015. © 2014 Wiley Periodicals, Inc.