Heterochrony and sexual dimorphism in the skull of the Liberian chimpanzee (Pan troglodytes verus)

Allometric methods and theory derived from principles of relative growth provide new and powerful approaches to an understanding of the nature and development of sexual dimorphism among living primates. The Frankfurt collection of Liberian chimpanzee skulls and mandibles provides a large skeletal sample from a single natural population of wild shot animals, including individuals of all ages and both sexes, and allows investigation of allometric and heterochronic patterns of sexual dimorphism. Univariate, bivariate, and multivariate analyses are utilized in this study in order to ascertain the ontogenetic nature of male-female differences in the skull of the Liberian chimpanzee. The results of univariate and multivariate analyses indicate that, while overall levels of sexual dimorphism in the chimpanzee skull are small, the greatest differences are in dimensions of the viscerocranium, while neurocranial dimensions and orbital size tend to be less dimorphic. Bivariate regressions of 21 cranial variables against basicranial length document positive allometry in many facial and mandibular dimensions, and isometry or negative allometry for most neurocranial dimensions. The data confirm previous work in chimpanzees and other anthropoid primates suggesting that males and females are “ontogenetically scaled” in most cranial traits. That is, males and females share the same cranial growth trajectories, although ending up at different points. Both rate and time hypermorphosis are suggested as underlying causes of ontogenetic scaling in the Liberian chimpanzee.     

Cranial ontogenetic variability,sex ratio and age structure of the Red fox

Describing the sex ratio, age structure of the population and ontogenetic variability of Red fox, Vulpes vulpes (Canidae, Carnivora) skull parameters, this study is based on 416 male and 289 female skulls collected in the Czech Republic. The skulls analysed came from feral individuals, that were shot by hunters. The male to female ratio was 1:0.69 regarding the whole population. Individuals younger than one year prevailed in the population (54% males, 48% females were in their first year of life). Four growth patterns of skull dimensions were described. The first group included mainly skull length dimensions (e.g. condylobasal length). They grew rapidly until the sixth month of life, becoming stabilised afterwards. The second group comprised parameters that were stable throughout the life (e.g. cheek tooth rows). Measurements representing the third growth pattern showed continual growth (mainly width dimensions, e.g. zygomatic breadth). Conversely, smaller dimensions of postorbital breadth were observed after the sixth month of life. Postorbital breadth represented the fourth growth pattern. It was concluded, that male and female Red foxes had similar ontogenetic skull development, even though there were some differences, e.g. in jugular breadth, which increased after the age of six months in males unlike in females.     

Growth of skeletal components in the young squirrel monkey (Saimiri sciureus boliviensis): a longitudinal experiment

A functional skeletal criterion, as an extension of the van der Klaauw's cranial theory, was adopted in the present study. The null hypothesis tested was: "The major skeletal components of the platyrrhine body grow linearly, regardless of their functional dependence to different demands." The acceptance of the hypothesis will imply that all Saimiri skeletal growth may be satisfactorily explained by independent variables in a single equation. The rejection will suggest that such skeletal growth patterns have to be explained by variables in several different equations, and perhaps these equations may vary with the effect of sex and undernutrition. Control and undernourished squirrel monkeys were radiographed monthly for 2 years; they were also measured; and their volumetric and morphometric neurocranial, facial, and pelvic indices were calculated. The curves that best described each of the 24-point sequences were obtained. Three main growth patterns were observed: 1) Simple linear (femur length for all groups, and pelvic index for control and undernourished females), for which the simple regression equation explained more than 95% of the variation; 2) Complex linear (pelvic index for control and undernourished males, and neurocranial and facial indices for all of the groups), for which more than 95% of the variation was explained by one of the four four-function type equations; and 3) Noncorrelated with age (neurofacial index for undernourished males, and pelviofemoral index for control females and undernourished males and females), which showed nonsignificant correlations with respect to age. The food intake and the oscillations of the environmental temperature might help to explain the undulating growth trajectory observed in the complex linear components.     

Morphometric variation in the Hooded seal (Cystophora cristata)

Intra- and intersexual variation in 16 skull dimensions and total body length of 233 aged Hooded seals caught in the north-west Atlantic were investigated. Absolute growth was described by asymptotic growth curves applied to single dimensions, as well as to scores on the first principal component of logarithmically transformed cranial data, which are believed to reflect the multivariate nature of growth. All dimensions were found to be fully developed later in males than in females. The growth of the male skulls was found to continue for more than 20 years, while the females approach final size about–7 years earlier than males, according to the scores on the first principal component. Female seals were found to reach 86 % of final total body length at the time of maturation, which is a generalized pinniped pattern. In both sexes, a yearling skull was characterized by a large brain-case, which decreased relatively with growth. The male skulls were further characterized by an increase in zygomatic width and orbital width in relation to basal length, a pattern which was not found in females.
All the asymptotic values were significantly larger in males than in females. The dimorphism develops mainly as a result of prolonged growth of males after the attainment of sexual maturity.     

Macroscopic and roentgenographic anatomy of the skull of the ferret (Mustela putorius furo)

Normal macroscopic and roentgenographic features of the skull of the ferret (Mustela putorius furo) were examined and described. Data were based on a sample of 100 (50 male and 50 female) adult ferrets of known body weight and age. The skull was described macroscopically according to six standard views, i.e. dorsal, lateral, ventral, caudal, cranial and midsagittal. The mandible was described separately. The roentgenographic characteristics of the ferret skull were demonstrated only in lateral and dorsoventral projections. Furthermore, the skull length and width as well as the minimum frontal width were measured, and skull indices were derived from relevant measurements. Sexual dimorphism was examined both morphologically and craniometrically. Besides the common features of a carnivore skull, the ferret skull is relatively elongated and flat with a short facial region. The skulls of adult male ferrets are about 17% longer and 22% wider than those of the females. Significant sexual dimorphism also exists regarding certain skull indices. The general features and some dimensional parameters of the adult ferret skull support the contention that the ferret would be an interesting and workable alternative animal model in craniofacial research.     

Craniofacial sequelae of lesions to facial and trigeminal motor nuclei in growing rats

Unilateral electrolytic lesions were made in the left-side facial motor nucleus (FMNu) of six Sprague-Dawley rats at 35 days of age in order to correlate craniofacial sequelae with changed motoneuron function. Experimental and control rats were killed at 22, 32, 42, and 52 days postoperatively to provide muscle weight, brain histology, and dry skull preparations for analyses. Dissection, muscle weight, motoneuron count, and osteometric data revealed that lesion-side facial and masticatory muscles were affected by the lesions. Paired t-tests indicated that significant differences existed between weights of experimental lesion- and nonlesion-side anterior digastric, temporalis, masseteric complex, and medial pterygoid muscles, numbers of facial and trigeminal motoneurons, and several skeletal dimensions of the skull. Basi-cranial dimensions of experimental animals were least affected by the lesion, whereas zygomatic arch, dorsal facial region, and mandibular condyle dimensions were most affected. Statistical analyses also detected significant differences between experimental and control groups for several skeletal dimensions of the skull. Data indicated that damage to the trigeminal motor nucleus (TMNu) was secondary to the primary lesion in the FMNu. Motoneurons within the facial and trigeminal neuromuscular complexes (FNC and TNC) play an important role in craniofacial growth and development.     

Biology of the rice rat (Oryzomys palustris natator) in a laborabory environment. X. Postweaning growth of the skull.

A survey of skull development of the rice rat (Oryzomys palustris natator) covering a period of 21 days to 16 months involving equal numbers of males and females (108) was undertaken using various skull parameters. Sexual dimorphism in relation to body weight and length emerged before that of the skull which was only clearly marked by 2 months. A number of the skull parameters by virtue of their individual growth potential did not show sexual differences in size for a long time, whereas others were clearly marked. Once the differences between the sexes had been established they were maintained. One parameter - lenght of cranium - represents those parameters which show changes and it was found that the male rat continued to show a steady linear increase in length up to 16 months, whereas the female cranial length shows a deceleration with termination around 12 months. The skull differences are mirrored in the weight and length findings and it suggested that the earlier sexual maturity of the females is compensated by an earlier cessation of growth compared to the males which mature later and have a correspondingly later cessation.     

Ontogeny of craniofacial sexual dimorphism in the orangutan (Pongo pygmaeus). I: face and palate

The orangutan is widely recognized as a highly dimorphic species. An ontogenetic approach to the study of sexual dimorphism can assist researchers in understanding both where and when these differences develop. In this study, 357 orangutans from Borneo were divided into five developmental stages representing infancy to mature adulthood. Three-dimensional (3D) coordinate data from 16 landmarks representing the face and palate were analyzed by means of a Euclidean distance matrix analysis (EDMA), a quantitative method for the comparison of forms. Three separate analyses (an age-specific static comparison of forms, a sex-specific analysis of growth trajectories, and an intersex comparison of patterns of relative growth) were carried out with the intent to describe the rate, timing, magnitude, and pattern of growth in the orangutan face and palate. The results indicate that generally males and females share a similar, but not identical, pattern of growth or local form change, but differ in growth rate, timing, and magnitude of difference. Dimorphism in the face and palate can be localized in infancy and traced throughout all age intervals. Orangutan females grow slightly faster than males from infancy to adolescence, at which time male growth exceeds female growth. Female growth ceases with the advent of adulthood, while male growth continues (i.e., both the number and magnitude of the dimorphic dimensions increase). Males and females are similar in facial dimensions and growth related to the orbits, upper face, and palate width. They maintain these similarities throughout development. However, they differ in facial and nasal height, palate length, snout projection, depth of the nasopharynx, and hafting of the face onto the skull. The face broadens and the zygomatic bone flares dramatically in adult males, corresponding to the development of cheek pads. While growth patterns are similar between the two sexes, they differ in the lateral orbit, snout projection, and hafting of the face onto the cranium. Adult dimorphism is the result of growth patterns experienced throughout life, and it is not equally expressed across the cranium. An understanding of patterns of dimorphism, along with the magnitude of difference, may be helpful for interpreting dimorphism in the fossil record.     

基于内壳的西非乌贼日龄与生长特性

乌贼主要分布在北大西洋东部及非洲沿岸海域,是拖网渔业的重要捕捞对象,也是大西洋乌贼科最重要的商业种。本研究根据2015年7—8月西非沿岸采集的乌贼样本,测量了283 尾乌贼内壳外部形态参数,结合内壳日龄信息,对不同性别乌贼生长特性进行了研究。结果表明: 不同性别乌贼胴长和体重存在显著差异。通过读取内壳日龄,发现雌雄性优势日龄组均为80～100 d。内壳外部形态与日龄的拟合函数中,Logistic函数的拟合效果最佳。雌性个体内壳外部形态参数的最大似然估计值均大于雄性。雄性和雌性乌贼内壳长和内壳宽的绝对和瞬时相对生长率分别在110～120 d 和100～110 d 达到峰值。雄性和雌性的初次性成熟日龄分别为111 d和104 d。不同性别乌贼的生长差异可能与栖息环境密切相关。     

Longitudinal roentgencephalometric study of the growth of the New Zealand white rabbit: cumulative and biweekly incremental growth rates for skull and mandible

A longitudinal roentgencephalometric study of the New Zealand white rabbit has documented postnatal skull and mandible growth from 2 to 34 weeks of age. Dorsoventral and lateral radiographs were performed using a specially constructed restrainer that allowed for standardized films and did not require the use of anesthesia. Assessments from 17 male and 12 female rabbits at biweekly intervals documented cumulative growth as well as biweekly incremental growth. Indices reported here include skull length, interzygomatic width, intercondylar width, and mandibular length. Mean skull length at 2 weeks was 54% of that at 34 weeks, and by 16 weeks 91% of adult skull length was achieved. Mean interzygomatic width at 2 weeks was 60% of that at 34 weeks, and by 16 weeks 91% of adult male and 94% of adult female widths were achieved. Mean mandibular length at 2 weeks was 47% of that at 34 weeks, and by age 16 weeks 90% of adult length was achieved. Mean intercondylar width at 2 weeks was 57% of that at 34 weeks, and by 16 weeks 89% of adult male and 93% of adult female widths were achieved. Biweekly increments decreased continually from 2 weeks of age on for all indices.     

Craniometric allometry in the thicktailed bushbabyOtolemur Crassicaudatus

Static adult craniometric allometry was evaluated in a sample of 66Otolemur crassicaudatus skulls (34 males, 32 females). Although cranial measures were equally well correlated to skull length in males and females, there were noteworthy differences in the exponential values between the sexes. These results underlined the need for caution when allometric analyses are based on pooled data. From the cranial allometric analyses it is concluded that the longer the skull, the shorter and the narrower the maxilla, and the broader the bizygomatic distance. Although cranial length increased proportionately to the increase in skull length, the cranial width in females was positively allometric whilst in males it was negatively allometric. Allometric analyses of mandibular dimensions suggest that larger animals will have proportionately longer mandibulae, which will, in turn, be relatively wider across the gonia, yet shallower behind the first molars. It is postulated that the disproportionate widening of the zygomata might be related to the widening across the gonia.     

Postnatal growth of skull linear measurements of Cape Hare Lepus capensis in northern China: an analysis in an adaptive context

304 skulls of Cape hare (Lepus capensis) were collected from two climatically distinct localities in northern China. With eye lens weight as a continuous age variable, postnatal growth patterns of 25 cranial linear measurements in relation to sex, growth season and region were analysed to understand the morphological basis of life history adaptation. In almost all the skull measurements, no significant differences were found between either sex or growth seasons. Principal component analysis revealed that facial elements accounted for the greatest proportion of skull morphological variation. Von Bertalanffy function was applied to describe growth trajectories of the skull elements. Based on this model, the growth rates of skull elements and the age at which they reached a certain proportion (95%) of asymptotes were compared. The results showed that skull growth exhibited an allometric pattern, with neural components attaining their final size more rapidly (at about 2–3 months old in tympanic bulla and 4–6 months old in others) than did the facial, which continued to grow well into postnatal life (at 6–10 months old). The earlier establishment of neurocranial morphology was associated with a fully developed central nervous system, which may play a key role in improving the survival of animals during the early phase of life. There was a regional difference in developmental rate of the hare skull. For all the skull parameters, northern hares had a more rapid rate of cranial growth compared to the southern, i.e. skull elements of juveniles from northern population were relatively larger at comparable ages and achieved adult size 0.5–4.0 months earlier than those from the south. In adult hares, however, no significant regional differences in any of the skull parameters were present. Adaptive explanations for the regional difference in ontogenetic pattern of skull morphology include age‐specific thermoregulation constraint, season‐related food availability and age‐dependent predation pressure. Based on the findings of this study, it is suggested that the postnatal growing period represents a crucial time of life, and that improvement of survivorship when young by growth adaptation forms an important aspect of the hare's life history strategies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 343–353.     

Changes in skull components of the squirrel monkey evoked by growth and nutrition: an experimental study

Twenty weanling 6-month-old male squirrel monkeys were allotted to the following treatments: 1) first control animals were killed at weaning; 2) second control animals were killed when 24 months old; and 3) malnourished animals were fed on a low-protein diet and killed at age 24 months. Lateral and vertical teleradiographies were taken. Growth of the neurocranial and splanchnocranial components were measured by volumetric (size estimators) and morphometric (shape estimators) indices. All facial components grew. The neurocranial components showed a heterogeneous behavior: The anteroneural component remained stable, and the increase of the midneural component was compensated by a decrease in the posteroneural component. Malnutrition affected the growths of 1) the craniofacial complex, 2) the splanchnocranium, and 3) the respiratory and midneural components. Growth influenced skull shape through 1) increases of the splanchnocranium and the midneural component relative to the neurocranium; 2) decreases of the masticatory and optic components relative to the splanchnocranium, and 3) decreases of the anteroneural and posteroneural components relative to the neurocranium. Malnutrition influenced skull shape through the relationship between the anteroneural component and the neurocranium. These results confirmed the existence of functional interrelationships among the cranial components. A new approach to craniological studies is suggested.     

A commentary on the growth of the human brain and skull

This paper tests the suggestion put forth by Tanner ('55) and Eichorn and Bayley ('62) to the effect that the brain participates in the parapubertal spurt of growth which characterizes many of the dimensions of the human body. To this end, longitudinal data consisting of oriented head roentgenograms of 11 boys were examined. Two measurements were taken directly from each lateral head film: (1) skull length, measured from glabella to opisthocranion, and (2) endocranial length, the maximum length of the endocranial contour in the mid-sagittal plane. While many of the individual cumulative curves depicting growth in skull length exhibit a parapubertal acceleration, all of the curves for endocranial length comprise segments of a parabolic arc representing a single decelerating phase of growth. Mean incremental curves, mathematically fitted, further emphasize the differences in velocity and pattern of size attainment for the two dimensions tested. The data here presented, then, fail to implicate the brain in the general spurt of growth evident for the external dimensions of the head at adolescence. It is suggested that two discrete systems are evident in the growth of the skull: a rapidly growing neural system essentially completed by adolescence, and a facial system of slower growth and longer duration. The conventional measurement of skull length cuts across both systems, appraising neural growth and the cerebral skeletal envelope prior to adolescence, and then superimposing the facial component, the forward projection of the frontal sinus, during adolescence and post-adolescence.     

The ontogeny of cranial sexual dimorphism in two old world monkeys:Macaca fascicularis (Cercopithecinae) andNasalis larvatus (Colobinae)

Allometric and heterochronic approaches to sexual dimorphism have contributed much to our understanding of the evolutionary morphology of the primate skull and dentition. To date, however, extensive studies of sexual dimorphism have been carried out only on the great apes and a few cercopithecine monkeys. To fill this gap, representative dimensions of the skull were collected among ontogenetic series of two dimorphic Old World monkeys:Macaca fascicularis (Cercopithecinae) andNasalis larvatus (Colobinae). The ontogeny of cranial sexual dimorphism was evaluated with least-squares bivariate regression, analysis of covariance (ANCOVA), and analysis of variance (ANOVA). Results indicate that within each species the sexes typically exhibit nonsignificant differences in ANCOVAs of ontogenetic trajectories, except for bivariate comparisons with bicanine breadth. AmongMacaca fascicularis, ANOVAs between males and females of common dental ages show that adult, and frequently subadult, males are significantly larger than females, i.e., sexual dimorphism develops via time and rate hypermorphosis (males primarily grow for a longer time period as well as faster). AmongNasalis larvatus, however, comparisons between males and females of common dental ages indicate that only adult males are significantly larger than females, i.e., sexual dimorphism develops primarily via time hypermorphosis (males grow for a longer time period). Within both species, females appear to exhibit an early growth spurt at dental age 2; that is, many cranial measures for females tend to be larger than those for males. Measures of the circumorbital region (e.g., browridge height), body weight, and bicanine breadth exhibit typically the highest sexual dimorphism ratios. The fact that postcanine toothrow length and neurocranial volume (less so inNasalis) demonstrate very low dimorphism ratios generally supports assertions that postnatal systemic growth (and associated selective pressures thereon) exerts a greater influence on facial, but not neural, dental, or orbital, development (Cochard, 1985, 1987; Shea, 1985a,b, 1986; Shea and Gomez, 1988; Sheaet al., 1990). Additional consideration of ontogenetic differences between species generally supports previous functional interpretations of subfamilial differences in cranial form related to agonistic displays in cercopithecine monkeys (Ravosa, 1990).     

Intraspecific variation in the external morphology of the sand skate

In male sand skate Psammobatis extenta , from the commercial fishery of Puerto Quequén, Argentina, six of 13 proportional dimensions analysed were constant (isometric) with total length, while seven proportions remained constant in females. The nature of the allometric and isometric relationships of disc length, disc width, interspiracular distance and interorbital distance did not vary between the sexes, and no sexual dimorphism occurred between these morphometric dimensions. A stepwise discriminant function analysis separated the sexes and was able to correctly classify 90·8% of the original grouped cases. The variables that best discriminated between sexes were prenasal length, distance from the cloaca to caudal fin tip, and distance from the snout to cloaca. The study also revealed that several of the meristic characters studied showed much intraspecific variation and, therefore, should be used with caution in taxonomic studies.     

高原鼢鼠(Myospalax baileyi) 不同地理种群的形态变异

本文采用主成分分析和聚类分析两种方法,分析了高原鼢鼠(Myospalax baileyi)12 个地理种群17 项形态特征的变异。主成分分析结果显示,分析构建的雄性前3 个主成分累积贡献率为78.483% ,而雌性前4 个主成分累积贡献率为79.587% 。对种群间形态差异分析贡献最大的指标为颅全长、基长、后头宽等反映头骨大小性状,以及顶嵴、额嵴最小间距体现的颧弓扩张程度指标。主成分分析结果与聚类分析结果一致,12 个地理种群分化构成两个大的地域性群体:一个由甘肃种群构成的甘肃群体;另一个由四川种群和青海种群构成的混合群体。两个地域性群体实际上反映出两个不同的分化方向:一个方向表现为头骨较小,顶嵴、额嵴相对分开即颧弓扩张程度小,如甘肃群体;另一个则表现为头骨较大,顶嵴、额嵴相对靠拢即颧弓扩张程度大,如混合群体。这种地域群体的形态变异可以解释为,地下洞道系统中环境因子选择作用对高原鼢鼠形态变化产生了重要影响。     

Craniofacial suture stenosis: morphologic effects

Craniofacial anomalies, such as Apert's and Crouzon's syndromes, are presumed to be related to premature growth arrest of cranial base growth sites. However, premature growth arrest at cranial vault sutures in animals appears to play a causative role in the development of cranial deformities characteristic of single-suture, or simple, craniosynostosis in humans. To study the possible causative role of cranial vault and other (interface) suture stenoses on the development of craniofacial deformity, a vault suture and an interface suture between the cranial vault and facial skeleton were simultaneously immobilized. Thirty-one New Zealand White rabbits at 9 days of age underwent implantation of dental amalgam growth markers adjacent to cranial vault and facial sutures. In the experimental group (n = 15), methylcyanoacrylate adhesive was applied over the coronal (vault) and frontonasal (interface suture between vault and facial skeleton) sutures to immobilize them. The remaining 16 animals served as sham-treated controls. All animals underwent serial radiographic cephalometry to document growth effects in the cranial vault, cranial base, and facial skeleton. Application of adhesive resulted in statistically significant (p less than 0.05) reduction in growth at the coronal and frontonasal sutures. This was accompanied by an overall significant reduction in neurocranial vault length during the first 30 days of development.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sex Ratio and Sexual Size Dimorphism in a Toad-headed Lizard,Phrynocephalus guinanensis

Phrynocephalus guinanensis has sexual dimorphism in abdominal coloration, but its ontogenetic development of sexual size dimorphism(SSD) is unknown. Using mark-recapture data during four days each year from August from 2014 to 2016, we investigated the development of sex ratios, SSD, sex-specific survivorship and growth rates in a population of P. guinanensis. Our results indicated that the sex ratio of males to females was 1:2.8. Males had a lower survival rate(6%) than females(14%) across the age range from hatchling to adult, which supported the discovered female-biased sex ratio potentially associated with the low survival rate of males between hatchlings and juveniles. Male-biased SSD in tail length and head width existed in adults rather than in hatchling or juvenile lizards. The growth rates in body dimensions were undistinguishable between the sexes during the age from hatchling to juvenile, but the growth rate in head length from juvenile to adult was significantly larger in males than females. Average growth rate of all morphological measurements from hatchling to juvenile were larger compared with corresponding measurements from juvenile to adult, but only being significant in tail length, head width, abdomen length in females and snout-vent length in males. We provided a case study to strengthen our understanding of the important life history traits on how a viviparous lizard population can survive and develop their morphology in cold climates.     

Observation of marking-like behavior, marking behavior, and growth of the scent gland in young Mongolian gerbils (Meriones unguiculatus) of an inbred strain.

A marking-like behavior (defined by authors), a marking behavior, and growth of the scent glands were observed in young Mongolian gerbils of an inbred strain. In males and females, a marking-like behavior, in which animals rub their abdominal scent glands on the floor, began to be seen at the age of 19 days and could be seen in almost all the gerbils at 22 days of age during the suckling period. The frequency of this behavior was highest at 60 days of age (males: 17.9/10 min, females: 15.4/10 min) and there was no sex difference. Marking behavior, in which animals rub their abdominal scent glands on small protruding objects, began to be seen at the age of 40 days in males and 50 days in females. The frequency of this behavior tended to increase until 90 days of age in males (13.7/10 min), but the levels were low (2.5-5.0/10 min) in females. The values in the male group therefore tended to be higher than that in the female group. Macroscopic scent gland pads were clearly observed at the age of 30 days in males, but not until 45 days of age in females. At the age of 45-90 days, the length of the scent gland pad in males and females was 2.1-2.8 and 1.6-1.7 cm, respectively and the width was 0.3-0.5 in males and 0.2-0.3 cm in females. During this period, the length and depth of the pads in males were significantly greater than those in females (p < 0.05). Histological examination of the structure of the scent glands after the age of 45 days showed that the development of clusters of acinar cells in females occurred much later than that in males, but the basic structure of these glands was similar in both sexes. These results suggest that the marking-like behavior was manifested although during the period when the scent glands had not yet developed, whereas true marking behavior first occurred when the glands were moderately well developed.