Allometric aspects of population dynamics: A symmetry approach

Summary The maximum intrinsic rate of increase (r _max ) shows a –0.25 scaling with adult body weight (W) in mammals (and others). Average adult life span (1/M) and age at maturity () show –0.25 scalings, independent of population size; these two lead to ther _max scaling, providedR _o is invariant with body size in rarified populations (=R _om ). Thus ther _max scaling follows from the existence of two population size symmetries (i.e. 1/M and ) and one body size symmetry (R _o ). The theory provides a formula to calculate the height (A ₃) of the scalingr _max =A ₃ ·W ^–0.25. The theory also helps to explain Fowler's Rules, which linkR _om to the relative position of the inflection point of the population growth curve.     

Life‐history constraints on maximum population growth for loggerhead turtles in the northwest Atlantic

Conservation planning for protected species often relies on estimates of life‐history parameters. A commonly used parameter is the instantaneous maximum population growth rate (r_max) that can be used to limit removals and design recovery targets. Estimation of r_max can be challenging because of limited availability of species‐ and population‐specific data and life‐history information. We applied a method proposed by Neil and Lebreton, originally developed for birds, to loggerhead turtles. The method uses age‐at‐first‐reproduction and adult survival to estimate r_max. We used a variety of datasets and matrix population models to confirm an allometric assumption required by the method, and to generate estimates of age‐at‐first‐reproduction and adult survival. A meta‐analysis was applied to parameters from reported growth curves, which were then combined with the size distribution of neophyte nesters to derive estimates of age‐at‐first‐reproduction. Adult survival rates were obtained from an existing matrix population model. Monte Carlo simulation was then used to combine the estimates of the allometric coefficients, age‐at‐first‐reproduction, and adult survival to obtain a probability distribution of approximate r_max values. Estimated annual maximum population growth rates averaged 0.024, with a mode of 0.017 and a 95% highest density interval of 0.006–0.047. These estimates were similar to values reported by others using different methods and captured the variability in positive, annual change estimates across nesting beach sites for the northwest Atlantic loggerhead population. The use of life‐history parameters has a long history in wildlife and fisheries management and conservation planning. Our estimates of r_max, while having some biases and uncertainty, encompassed values presently used in recovery planning for loggerhead turtles and offer additional information for the management of endangered and threatened species.     

Optimum body size: effects of food quality and temperature,when reproductive growth rate is restricted,with examples from aphids

Most models of optimum energy partitioning predict variability in adult size, although not always explicitly. Increase in size is usually attributed to an increase in the growth rate or decline in mortality. The model presented shows that this may not always be the case. Even when mortality is kept constant in organisms with overlapping generations, a constraint on the maximum reproductive growth rate may lead, when the rate of overall growth increases, to either an increase or a decline in the optimum adult body size. It is shown that adult size could be a consequence of the differential responses of life history traits to changes in temperature and food quality. This is clearly advantageous for short lived organisms, like aphids, each generation of which only experience a very small part of the great seasonal range in conditions. This hypothesis complements Iwasa's (1991) explanation of the phenotypic plasticity observed in long lived organisms. The predictions are illustrated with empirical data from aphids. The model presented, which has been verified against a very large data set, indicates that for aphids the adult weight observed at a particular combination of temperature and food quality is that at which the population growth rate, r_m, is maximized. We conclude that predictions about adult size from models based on the partitioning of energy are more likely to apply to organisms that scramble for resources, i.e., “r” selected species. The size of organisms that contest for resources is more likely to be determined by competitive status and avoidance of natural enemies.     

DENSITY-DEPENDENT EVOLUTION OF LIFE-HISTORY TRAITS IN DROSOPHILA MELANOGASTER

Populations of Drosophila melanogaster were maintained for 36 generations in r- and K-selected environments in order to test the life-history predictions of theories on density-dependent selection. In the r-selection environment, populations were reduced to low densities by density-independent adult mortality, whereas populations in the K-selection environment were maintained at their carrying capacity. Some of the experimental results support the predictions or r- and K-selection theory; relative to the r-selected populations, the K-selected populations evolved an increased larval-to-adult viability, larger body size, and longer development time at high larval densities. Mueller and Ayala (1981) found that K-selected populations also have a higher rate of population growth at high densities. Other predictions of the thoery are contradicted by the lack of differences between the r and K populations in adult longevity and fecundity and a slower rate of development for r-selected individuals at low densities. The differences between selected populations in larval survivorship, larval-to-adult development time, and adult body size are strongly dependent on larval density, and there is a significant interaction between populations and larval density for each trait. This manifests an inadequacy of the theory on r- and K-selection, which does not take into account such interactions between genotypes and environments. We describe mechanisms that may explain the evolution of preadult life-history traits in our experiment and discuss the need for changes in theories of density-dependent selection.     

Perinatal life history traits in New World monkeys

Gestation length, neonatal and maternal body weight, and neonatal and adult brain weight data were collected for New World monkeys in an attempt to establish typical patterns of perinatal life history. This study attempts to illuminate the most accurate values from the available data, which suggest that gestation length and prenatal growth rate are broadly conserved in relation to maternal size in New World monkeys. Exceptions to the patterns evident in the data point to derivations in life history strategies. In particular, this study suggests that the extended gestation length of callitrichines is a function of minimum viable neonate size and not exclusively energy minimization associated with simultaneous lactation. Cebus is shown to undergo more postnatal brain growth relative to other New World monkeys, but not as much as previously believed. Alouatta is shown to be relatively small brained at birth as well as in adulthood. Saimiri is shown to present the most unusual package of perinatal life history traits, in which precocial neonates are gestated for a relatively long time and at a slightly faster growth rate than is typical for New World monkeys. © 1996 Wiley-Liss, Inc.     

Genetic variance and covariance patterns of larval development in the small white butterflyPieris rapae crucivora Boisduval

Quantitative genetic theory indicates that genetic covariance patterns among life history characters should have played an important role as genetic constraint in life history evolution. Highly positve (and negative) genetic correlations between larval development time (or larval growth rate) and adult size characters were detected by means of sib analysis for the small white butterfly Pieris rapae crucivora. The genetic associations suggested that evolution of developmental characteristics and adult phenotypic traits were constrained by pleiotropy. The positive genetic correlations between development time and adult body size may be compatible with the trade-off between them, but the negative genetic correlations between larval growth rate and adult body size are not predicted from theories of optimal energy allocation. That phenotypic correlations drastically differed from the genetic correlations indicates limitations of evolutionary inferences based only on phenotypic variation.     

Optimum body size in aphids

Abstract.

• 1 Smallness in British plant-sucking bugs is associated with feeding on the contents of individual plant cells, especially phloem.
• 2 The sizes of species of aphids living in similar habitats and the rates of water loss from those that have and lack a filter chamber do not support the contention that small size in aphids is a means of maximizing the loss of water by cuticular transpiration.
• 3 An optimum energy partitioning model, previously developed for aphids, was used to predict the interspecifie relation between adult weight and birth weight that would maximize the population growth rate, r_m. Using the observed r_m/IRGR ratio (0.8–0.9) and that the gonads are smaller than the soma at birth the predicted relation between the logarithm of the adult weight and the logarithm of the birth weight is linear and has a slope of 1.
• 4 The relation between the logarithm of the adult weight and the logarithm of the birth weight for eighteen species of aphids is linear and has a slope of 1.
• 5 Birth size differs between species of aphids and in each species is assumed to be the minimum size necessary for feeding. Accepting this constraint then the optimum adult size for maximizing r_m is approximately 15 times the birth size.

     

Intrinsic rate of natural increase in Neotropical forest mammals: relationship to phylogeny and diet

Summary The relationship of diet and phylogeny to the intrinsic rate of population increase (r _max) was examined in a sample of 39 mammalian species that live in Neotropical forests. Diets of species did not predict their r _max, contrary to published predictions based on associations between basal metabolic rate and diet and between basal metabolic rate and r _max. Phylogeny did however, apparently because life history characteristics and susceptibility to predation vary predictably with phylogeny and with one another.     

Ontogeny and limb bone scaling in two new world marsupials,Monodelphis domestica and Didelphis virginiana

This study examines the growth of two species of marsupials who share common ancestry and are born at the same neonatal size of a little less than 1 g. Despite this similarity at birth, adult size of these two species differs by about 50 times, with the smaller species believed to be the more ancestral. We quantified the growth in the limb bones (humerus, femur, ulna, tibia, metacarpal, and metatarsal) beginning around 40 days of age until adult size was reached. Results indicate that the larger species grows at a higher rate of growth as well as for a longer period of time to reach its larger adult size. Despite these differences in growth, there were few differences observed in the scaling over time of length to width in the various limb bones that were measured. The two species, although different in their adult size and the patterns of growth, maintain the same length to width proportions in each limb bone. The biggest difference between species in scaling was observed in the bones of the hands and feet, which may suggest adaptation to size and/or locomotor performance as body size increases. Despite variation in size, these heterochronic patterns do not affect the shape among adults or over evolutionary time. J Morphol 231:117–130, 1997. © 1997 Wiley-Liss, Inc.     

Ability of Young Women to Recall Past Body Size and Age at Menarche

Objective: The ability of young women (n = 132, average age 17 years) to recall body size and age at menarche was examined. The use of body silhouettes to assist women in recalling their body size at menarche and to represent their current body size was also evaluated. Research Methods and Procedures: Subjects, who previously participated in a cohort study, were asked to recall height and weight at the time of menarche, to select body silhouettes that best reflected their body shape at the time of menarche and their current body shape, and to recall age at menarche. Two sets of body silhouettes were developed, one representative of an adult body shape and another representative of an adolescent body shape. Results: Pearson correlation coefficients between the adult and adolescent body figures and actual body mass index (BMI; kg/m²) at the time of menarche were not significantly different (r = 0.82 for adult figures vs. r = 0.72 for adolescent figures, p > 0.05). The correlation between actual BMI at the time of menarche and body silhouette (r = 0.77, all subjects) was similar to the correlation between actual and recalled BMI at the time of menarche (r = 0.83) as well as the correlation between current BMI and current body silhouette (r = 0.75). Recalled and actual ages at menarche were highly correlated (r = 0.83). Discussion: The recall of body shape was considered to be a less precise measure of body size than asking about height and weight, but use of body silhouettes may offer advantages in certain situations.     

外寄生性花绒寄甲的寄生选择及其发育表现

"选择-表现"假说认为,成虫应该选择有利于子代发育的高品质寄主,但在寄主选择中,除了寄主品质外,其他因素也可能影响寄主选择决策。寄主选择研究通常以成虫为对象,而对那些初龄幼虫选择寄主的寄生性昆虫很少关注。以1龄幼虫积极搜寻寄主的寄生性花绒寄甲为模式生物,采用双选试验设计,观察了花绒寄甲初孵幼虫在不同体重青杨天牛幼虫之间、在已被寄生与健康的黄粉虫蛹之间的寄生选择性;然后采用回归设计,观察了花绒寄甲寄生若干不同体重的青杨天牛幼虫后的发育表现。研究结果表明,花绒寄甲1龄幼虫对体型较大的青杨天牛幼虫的选择偏好显著大于对体型较小的寄主幼虫的选择,选择大体型幼虫的比值比是选择小体型幼虫的4.55倍;对已被寄生的寄主黄粉虫蛹的选择偏好显著大于对健康寄主蛹的选择,选择已被寄生寄主的比值比是选择健康寄主的12.57倍。寄生青杨天牛幼虫的花绒寄甲幼虫发育历期平均为11.49 d、蛹历期为26.67 d、幼虫发育至成虫的羽化率50%,这些发育表现与寄生时青杨天牛幼虫的体重没有显著关系。但刚羽化寄甲成虫体重与寄生时寄主的体重存在显著的正直线关系:寄生时的寄主体重每增大0.01 g,羽化出的寄甲成虫体重增大近0.08%;方差分析寄甲成虫体重在不同寄主体重水平之间的差异表明,从体型较大寄主中羽化的寄甲成虫体重显著大于从体型较小寄主中羽化的成虫。研究结果说明,花绒寄甲初孵幼虫在寄主选择决策时,在寄主体型大小与被寄生状态之间可能采取折衷对策,而且对体型大小不同的寄主选择与子代发育适合度表现存在一致性,从而支持"选择-表现"假说。     

The potential for population growth of Ascomorpha ecaudis

The intrinsic rate of natural increase (r_m) of the rotifer Ascomorpha ecaudis was found to be a hyperbolic function of food concentration. The threshold food concentration and r_max/2-food concentration determined for this species were significantly lower than values predicted from allometric models. These growth characteristics may be related to the mucus house in which Ascomorpha lives and/or the symbiotic algae living in its body tissues. The maximum rate of population growth recorded (0.71 d^–1) was similar to that of other soft-bodied rotifers of similar body mass. These population growth characteristics and the resistance of this species to invertebrate predation should allow it to become a dominant member of freshwater zooplankton communities. However, field observations suggest that it is not. Reasons for this are suggested.     

City limits: Heat tolerance is influenced by body size and hydration state in an urban ant community

Cities are rapidly expanding, and global warming is intensified in urban environments due to the urban heat island effect. Therefore, urban animals may be particularly susceptible to warming associated with ongoing climate change. We used a comparative and manipulative approach to test three related hypotheses about the determinants of heat tolerance or critical thermal maximum (CT_max) in urban ants—specifically, that (a) body size, (b) hydration status, and (c) chosen microenvironments influence CT_max. We further tested a fourth hypothesis that native species are particularly physiologically vulnerable in urban environments. We manipulated water access and determined CT_max for 11 species common to cities in California's Central Valley that exhibit nearly 300‐fold variation in body size. There was a moderate phylogenetic signal influencing CT_max, and inter (but not intra) specific variation in body size influenced CT_max where larger species had higher CT_max. The sensitivity of ants’ CT_max to water availability exhibited species‐specific thresholds where short‐term water limitation (8 hr) reduced CT_max and body water content in some species while longer‐term water limitation (32 hr) was required to reduce these traits in other species. However, CT_max was not related to the temperatures chosen by ants during activity. Further, we found support for our fourth hypothesis because CT_max and estimates of thermal safety margin in native species were more sensitive to water availability relative to non‐native species. In sum, we provide evidence of links between heat tolerance and water availability, which will become critically important in an increasingly warm, dry, and urbanized world that others have shown may be selecting for smaller (not larger) body size.     

Intraspecific alkaloid variation in ladybird eggs and its effects on con- and hetero-specific intraguild predators

Sexual size dimorphism (SSD) is a common phenomenon in animals. In many species females are substantially larger than males. Because body size plays a central role in modulating the body temperature (T _b) of ectotherms, intersexual differences in body size may lead to important intersexual differences in thermoregulation. In addition, because SSD is realized by differences in growth rate and because growth rate is strongly temperature dependent in ectotherms, a conflict between male reproductive behaviour and thermoregulation may affect the expression of SSD. In this study, we investigated the thermal implications of SSD in a reptile exhibiting spectacular female-biased SSD: the northern map turtle (Graptemys geographica). Over three seasons, we collected >150,000 measurements of T _b in free-ranging adult and juvenile northern map turtles using surgically implanted miniature temperature loggers. Northern map turtles exhibited seasonal patterns of thermoregulation typical of reptiles in northern latitudes, but we found that large adult females experienced a lower daily maximum T _b and a narrower daily range of T _b than adult males and small juvenile females. In addition, despite more time spent basking, large adult females were not able to thermoregulate as accurately as small turtles. Our findings strongly suggest that body size limits the ability to thermoregulate accurately in large females. By comparing thermoregulatory patterns between adult males and juvenile females of similar body size, we found no evidence that male reproductive behaviours are an impediment to thermoregulation. We also quantified the thermal significance of basking behaviour. We found, contrary to previous findings, that aerial basking allows northern map turtles to raise their T _b substantially above water temperature, indicating that basking behaviour likely plays an important role in thermoregulation.     

Brain size at birth throughout human evolution: a new method for estimating neonatal brain size in hominins

An increase in brain size is a hallmark of human evolution. Questions regarding the evolution of brain development and obstetric constraints in the human lineage can be addressed with accurate estimates of the size of the brain at birth in hominins. Previous estimates of brain size at birth in fossil hominins have been calculated from regressions of neonatal body or brain mass to adult body mass, but this approach is problematic for two reasons: modern humans are outliers for these regressions, and hominin adult body masses are difficult to estimate. To accurately estimate the brain size at birth in extinct human ancestors, an equation is needed for which modern humans fit the anthropoid regression and one in which the hominin variable entered into the regression equation has limited error. Using phylogenetically sensitive statistics, a resampling approach, and brain-mass data from the literature and from National Primate Research Centers on 362 neonates and 2802 adults from eight different anthropoid species, we found that the size of the adult brain can strongly predict the size of the neonatal brain (r² = 0.97). This regression predicts human brain size, indicating that humans have precisely the brain size expected as an adult given the size of the brain at birth. We estimated the size of the neonatal brain in fossil hominins from a reduced major axis regression equation using published cranial capacities of 89 adult fossil crania. We suggest that australopiths gave birth to infants with cranial capacities that were on average 180 cc (95% CI: 158–205 cc), slightly larger than the average neonatal brain size of chimpanzees. Neonatal brain size increased in early Homo to 225 cc (95% CI: 198–257 cc) and in Homo erectus to approximately 270 cc (95% CI: 237–310 cc). These results have implications for interpreting the evolution of the birth process and brain development in all hominins from the australopiths and early Homo, through H. erectus, to Homo sapiens.     

Inter- and intraspecific diversity of ontogeny and fecundity patterns in relation to reproductive strategy choice in Myomorpha (Rodentia: Calomyscidae,Cricetidae, Muridae)

Studying the life history of various organisms is essential for ecological and evolutionary inferences. However, publications regarding models of evolutionary shifts to either r- or K-reproductive strategies are scarce. Here, the combined original and previously published data on reproduction and development of Goodwin’s brush-tailed mouse (Calomyscus elburzensis), golden hamster (Mesocricetus auratus), and house mouse (Mus musculus) were compared. An adult male and female in estrous state (from each species) were coupled with each other for a night and embryos were harvested at embryonic days (E) 10–17 from pregnant females. The mean gestation period in Goodwin’s brush-tailed mouse was 31.5?±?2.1 days; growth was rapid at the first half of this period and after that it was reduced by half. During the postnatal period, growth was rapid up to weaning, but after day 35, the rate of growth decreased abruptly. Growth rate of head and body length was very rapid prior to weaning (PN35) in comparison with weight but the rates of the two characters were inversed following weaning. Totally, weight gain was more dynamic in rate, as compared with head and body length, up to adultness. For golden hamster and house mouse, gestation period was 15.6?±?0.8 and 20.8?±?0.8 days, respectively. Growth was slow during approximately the first half of the gestation period in both species but increased during the second half. During postnatal period in golden hamster, growth rate of head and body length was rapid both before and after weaning as compared with that of weight. However, head and body length showed a stable rate of growth before and after weaning (around PN15), but for body weight, the growth rate was slightly faster after weaning. Similarly, changes in head and body length in house mouse were rapid both before and after weaning (PN16) as compared with that of weight. Since approximately two-fold increases were observed in the growth rates of weight and head and body length in the second stage of postnatal development (PN16-adult) compared with the rates during the first stage (PN0-PN16), both showed dynamic changes during the postnatal development in this species. Typically, r-strategic house mouse appeared to be more similar in early morphological development to comparatively K-strategic Goodwin’s brush-tailed mouse, than to comparatively r-strategic golden hamster. Nevertheless, growth rate and pattern were different in the three species after birth. Hence, results showed that peculiarities of association between an ontogenetic pattern and a tendency in reproductive strategy can differ in diverse groups of myomorph rodents.

     

Nestling growth in the African fish eagle in Uganda

Nestling African fish eagles Haliaeetus vocifer (Daudin, 1858) in Queen Elizabeth National Park (QEP), Uganda, attained and exceeded the adult body weights of their respective sexes by fledging time. The growth curves conformed with the sigmoid form of most animal species. The overall growth rate, K (g/day), was low, averaging 0.077 ± 0.017 g/day and the time t_10–90 required to complete growth from 10 to 90% of the asymptotic weight was c. 42 days. Brood size and hatching order did not significantly influence growth rates but the sex of the young did. The tarsus, femur and humerus reached adult sizes by fledging time but the standard wing did not.     

Relationship Between Birth Weight and Body Composition,Energy Metabolism,and Sympathetic Nervous System Activity Later in Life

Objective: Epidemiological studies suggest that high birth weight might be associated with an increased risk of obesity later in life. Programming of metabolic, endocrine, and/or autonomic pathways during intrauterine development has been proposed to explain this association. Research Methods and Procedures: To determine the relationship between birth weight and body composition and energy metabolism later in life, we measured fat mass and fat‐free mass (hydrodensitometry or double‐energy X‐ray absorptiometry), 24‐hour energy expenditure, sleeping metabolic rate, and 24‐hour respiratory quotient (respiratory chamber) in 272 adult nondiabetic Pima Indians (161 males/111 females, age 25 ± 5 years, mean ± SD). In these subjects, birth weight varied over a wide range (2000 to 5000 g). Individuals known to be offspring of diabetic pregnancies were excluded. In 44 of the 272 subjects, muscle sympathetic nerve activity was assessed by microneurography. Results: Birth weight was positively correlated with adult height (r = 0.20, p < 0.001) and fat‐free mass (r = 0.21, p < 0.001), but not with fat mass (r = 0.01, not significant). Sleeping metabolic rate, adjusted for age, sex, fat‐free mass, and fat mass, was negatively related to birth weight (r = ?0.13, p < 0.05), whereas adjusted 24‐hour energy expenditure (r = 0.07, not significant) and 24‐hour respiratory quotient (r = ?0.09, not significant) were not. There was no relationship between birth weight and muscle sympathetic nerve activity (r = 0.12, not significant, n = 44). Discussion: In Pima Indians who are not offspring of diabetic pregnancies, high birth weight is associated with increased height and lean body mass, but not with increased adiposity later in life. Although high birth weight may be associated with relatively low resting energy expenditure, it is not associated with major abnormalities in 24‐hour energy metabolism or with low muscle sympathetic nerve activity later in life.     

Intraspecific competition of Glyptotendipes paripes (Diptera: Chironomidae) larvae under laboratory conditions

Glyptotendipes paripes larvae were reared in wells of tissue culture plates, in groups of 2, 4, 8, 16, and 32 (representing densities of about 1,300, 2,600, 5,200, 10,400, and 20,800 larvae per m², respectively). Larval groups were supplied with one of two concentrations (low or high) of food and larvae were individually observed to evaluate the effects of density on mortality, growth, development, behavior, and adult body size. Increased larval densities resulted in higher mortality, as well as slower larval growth and development. The distribution of developmental time became flatter at higher density, with a wider range of values, or even became bimodal. This was a consequence of the most rapidly developing individuals at higher densities emerging as adults sooner than the fastest developing individuals at lower densities, although overall mean developmental time was longer at higher densities. At higher densities, growth and development of smaller larvae were slowed, based on the relative difference in body length between competitors. When larger competitors emerged as adults or died, the growth of smaller larvae may have accelerated, resulting in increased variability of developmental times. The effect of larval density on adult body size was complex, with the largest body size found at the lowest density and a second peak of adult size at high-middle densities, with smaller adult body sizes found at low-middle, and high densities. Similarly, as with developmental time, the range of body size increased with increasing density. Examined food concentrations had no effect on larval mortality, but significantly affected developmental time, growth rate, and adult body size. At higher densities, larvae spent more time gathering food and were engaged in aggressive or antagonistic behaviors.     

Diet and a developmental time constraint alter life‐history trade‐offs in a caddis fly (Trichoptera: Limnephilidae)

Environmental factors influence variation in life histories by affecting growth, development, and reproduction. We conducted an experiment in outdoor mesocosms to examine how diet and a time constraint on juvenile development (pond‐drying) influence life‐history trade‐offs (growth, development, adult body mass) in the caddis fly Limnephilus externus (Trichoptera: Limnephilidae). We predicted that: (1) diet supplementation would accelerate larval growth and development, and enhance survival to adulthood; (2) pond‐drying would accelerate development and increase larval mortality; and (3) the relationship between adult mass and age at maturity would be negative. Diet supplementation did lead to larger adult mass under nondrying conditions, but did not significantly alter growth or development rates. Contrary to predictions, pond‐drying reduced growth rates and delayed development. The slope (positive or negative) of the female mass–age at maturity relationship depended on interactions with diet or pond‐drying, but the male mass–age relationship was negative and independent of treatment. Our results suggest that pond‐drying can have negative effects on the future fitness of individuals by increasing the risk of desiccation‐induced, pre‐reproductive mortality and decreasing adult body size at maturity. These negative effects on life history cannot be overcome with additional nutritional resources in this species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 495–504.