Phylogenetic analyses of dimorphism in primates: evidence for stronger selection on canine size than on body size

Phylogenetic comparative methods were used to analyze the consequences of sexual selection on canine size and canine size dimorphism in primates. Our analyses of previously published body mass and canine size data revealed that the degree of sexual selection is correlated with canine size dimorphism, as well as with canine size in both sexes, in haplorhine but not in strepsirrhine primates. Consistent with these results, male and female canine size was found to be highly correlated in all primates. Since canine dimorphism and canine size in both sexes in haplorhines were found to be not only related to mating system but also to body size and body size dimorphism (characters which are also subject to or the result of sexual selection), it was not apparent whether the degree of canine dimorphism is the result of sexual selection on canine size itself, or whether canine dimorphism is instead a consequence of selection on body size, or vice versa. To distinguish among these possibilities, we conducted matched-pairs analyses on canine size after correcting for the effects of body size. These tests revealed significant effects of sexual selection on relative canine size, indicating that canine size is more important in haplorhine male-male competition than body size. Further analyses showed, however, that it was not possible to detect any evolutionary lag between canine size and body size, or between canine size dimorphism and body size dimorphism. Additional support for the notion of special selection on canine size consisted of allometric relationships in haplorhines between canine size and canine size dimorphism in males, as well as between canine size dimorphism and body size dimorphism. In conclusion, these analyses revealed that the effects of sexual selection on canine size are stronger than those on body size, perhaps indicating that canines are more important than body size in haplorhine male-male competition.     

The interspecific range size–body size relationship in Australian frogs

Aim There is substantial residual scatter about the positive range size–body size relationship in Australian frogs. We test whether species’ life history and abundance can account for this residual scatter. Location Australia. Methods Multiple regressions were performed using both cross‐species and independent contrasts analyses to determine whether clutch size, egg size and species abundance account for variation in range size over and above the effects of body size. Results In both cross‐species and independents contrasts models with body size, clutch size and egg size as predictors, partial r² values revealed that only egg size was significantly and uniquely related to range size. Contrary to expectation, neither body size nor clutch size could account for significant variation in range size. Incorporating species abundance as a predictor in further multiple regression analysis demonstrated that while abundance accounted for a significant proportion of range size variation, the contribution of egg size was reduced but still significant. Notably, non‐significant relationships persisted between range size and both body size and clutch size. Conclusions The weak positive correlation between body size and range size in Australian frogs disappears after accounting for species abundance and egg size. Our findings demonstrate that species with both high local abundance and small eggs occupy comparatively wider geographical ranges than species with low abundance and large eggs.     

The effective size of fluctuating populations

We consider a Wright-Fisher model whose population size is an autocorrelated stochastic process. Our interest is in the effects of autocorrelated fluctuations of the population size on the effective size. We define the inbreeding effective size and the variance effective size and show that these effective sizes are the same for this model. In the literature, it is said that the effective size is equal to the harmonic mean of population size when the size fluctuates. We will show, however, that the effective size is not the same as the harmonic mean of population size unless the fluctuations of population size are uncorrelated. The effective size is larger (resp. smaller) than the harmonic mean when the fluctuations of population size are positively (resp. negatively) autocorrelated. Further, we obtain some asymptotic expressions for effective size when the population size is very large and/or the autocorrelation of the fluctuation is very strong.     

Egg size variation in European Satyrine butterflies (Nymphalidae, Satyrinae)

The interspecific relationships between egg size and female size, sexual size dimorphism, latitude, preoviposition and egg-laying behaviour, was investigated in European Satyrine butterflies by controlling for taxonomic relatedness and by simple cross-species comparisons. Egg size scales positively to female size, although the relationship is not significant (but nearly so) when taxonomic effects are controlled for. Egg size is correlated with latitude. When the effects of latitude are removed, the relationship between egg and female size becomes more evident. Sexual size dimorphism is negatively correlated with egg size, but the effect disappears when adult size is controlled for. Small egg size is related to long preoviposition. No relationship between egg laying mode and egg size appears. The results are interpreted as evidence for positive scaling of egg to female size, with superimposed effects of egg size radiation. The correlation of egg size with latitude suggests that temperature, or season length, is likely to have made an important contribution to egg size selection within this geographic area.     

Reproductive variation and the egg size-clutch size trade-off within and among populations of painted turtles (Chrysemys picta bellii)

Interpopulation variation in egg size, clutch size and clutch mass was studied 3 years in four populations of painted turtles (Chrysemys picta bellii) from western Nebraska. Body size varied among all populations and was larger in two large (56–110 ha), sandhills lake populations than in two populations in smaller habitats (1.5–3.6 ha) of the Platte River floodplain. Reproductive parameters (egg mass, clutch mass, and clutch size) generally increased with maternal body size within populations. Clutch wet and dry mass varied among populations but largely as a function of maternal body size. Clutch size was largest in the sandhills lake populations, both absolutely and relative to maternal body size. Egg mass was smallest in the sandhills lakes and varied annually in one population. Over all populations, an egg sizeclutch size trade-off was detected (a negative correlation between egg mass and clutch size) after statistically removing maternal body size effects. Egg wet mass and clutch size were negatively correlated over all years within the sandhills populations and in some years in three populations. Although egg size varied within populations, egg size and clutch size covaried as expected by optimal offspring size models. Thus, patterns of egg size variation should be interpreted in the context of proximate or adaptive maternal body size and temporal effects. Comparisons among populations suggest that large egg size relative to maternal body size may occur when juvenile growth potential is poor and mean maternal body size is small.     

Group Size in Animal Societies: The Potential Role of Social and Ecological Limitations in the Group‐Living Fish,Paragobiodon xanthosomus

The number of group members in an animal society can have a major influence on group members’ life history, survival, and reproductive success. Identifying the factors that limit group size is therefore fundamental for a complete understanding of social behavior. Here, I examined the relationships between resource availability, social conflict, and group size in the coral‐dwelling fish, Paragobiogon xanthosomus (Gobiidae). The size of the largest (breeding) female and the minimum size difference required for hierarchy stability strongly but not perfectly predicted maximum group size, suggesting that social conflicts and hierarchy structure set the upper limit on group size. Deviations in group size around the predicted maximum were explained by variation in average body size ratios but not by variation in coral size, suggesting that coral size does not directly influence group size. In contrast, coral size was a significant predictor of body size ratios, and possible explanations for this relationship are discussed. Group size may be limited by the social conflicts that characterize size‐based dominance hierarchies. Ecological factors, namely coral size, may in turn play an indirect role via an effect on body size ratios.     

Relationships between brood size and offspring body size in an ovoviviparous fish: maternal effects and genetic trade-off

Maternal effects can form an important source of variation in offspring fitness and have important evolutionary and ecological consequences. To explore genetic control for body size of newborn offspring, the present study examined maternal brood size and offspring body size in 14 strains and performed cross experiments in an ovoviviparous fish, guppy. Correlation analyses among the strains indicated that the strains with larger brood size have smaller offspring body size. Diallel and reciprocal crosses among four strains revealed a large maternal, but no paternal, contribution to offspring body size. To examine whether offspring body size is determined by maternal genotypes and whether offspring body size correlates with albinism, backcrosses were performed between a wild-type strain with large offspring body size and an albino strain. Offspring body size differed from both the parental strains in one generation of backcrossing but was not significantly different from the paternal strains after three generations. In the backcross generations, the offspring yielded by albino individuals showed significantly smaller body size than those yielded by normal individuals. These results indicated that offspring body size is determined by maternal genotypes, suggesting that offspring body size is influenced by the maternal albinism gene or the maternal locus linked with the albinism locus. The significant negative correlation between maternal brood size and offspring body size detected through our experiments showed that the maternal genetic character of brood size strongly correlated with offspring body size, indicating genetic trade-off between maternal brood size and offspring body size in the guppy.     

Fruit–frugivore interactions in two southern hemisphere forests: allometry,phylogeny and body size

The size of fleshy fruits spans several orders of magnitude. However, the evolution of fruit size diversity is poorly understood. Fruit size diversity is hypothesised to result from several potential processes. The frugivore hypothesis postulates that different‐sized animal fruit consumers select for different‐sized fruits. The correlated selection hypothesis postulates that fruit size is allometrically related to other plant traits (e.g. leaf size, plant height); therefore differences in fruit size result from correlated evolution with other plant traits. We tested the frugivore and correlated selection hypotheses as potential explanations for fruit size diversity in two New Zealand study sites. We observed birds foraging for fruits over two fruiting seasons at each site and measured fruit size, leaf size and plant height in a total of 32 plant species. Relationships between average fruit size, leaf size, plant size and the average size of birds consuming each fruit species were then evaluated using phylogenetically independent contrasts. Similar results were obtained in both study sites. Fruit size was correlated with the size of avian fruit consumers, but was unrelated to leaf size or plant height. Therefore, results falsified the correlated selection hypothesis but failed to falsify the frugivore hypothesis. Although results suggest that frugivores may have influenced the evolution of fruit size in New Zealand, further study is needed to generate a mechanistic understanding of how frugivores may have selected for interspecific variation in fruit size.     

A neutral model with fluctuating population size and its effective size

We consider a diffusion model with neutral alleles whose population size is fluctuating randomly. For this model, the effects of fluctuation of population size on the effective size are investigated. The effective size defined by the equilibrium average heterozygosity is larger than the harmonic mean of population size but smaller than the arithmetic mean of population size. To see explicitly the effects of fluctuation of population size on the effective size, we investigate a special case where population size fluctuates between two distinct states. In some cases, the effective size is very different from the harmonic mean. For this concrete model, we also obtain the stationary distribution of the average heterozygosity. Asymptotic behavior of the effective size is obtained when the population size is large and/or autocorrelation of the fluctuation is weak or strong.     

Mitotic chromosome length scales in response to both cell and nuclear size

Multicellular development requires that cells reduce in size as a result of consecutive cell divisions without increase in embryo volume. To maintain cellular integrity, organelle size adapts to cell size throughout development. During mitosis, the longest chromosome arm must be shorter than half of the mitotic spindle for proper chromosome segregation. Using high-resolution time-lapse microscopy of living Caenorhabditis elegans embryos, we have quantified the relation between cell size and chromosome length. In control embryos, chromosome length scaled to cell size. Artificial reduction of cell size resulted in a shortening of chromosome length, following a trend predicted by measurements from control embryos. Disturbing the RAN (Ras-related nuclear protein)-GTP gradient decoupled nuclear size from cell size and resulted in chromosome scaling to nuclear size rather than cell size; smaller nuclei contained shorter chromosomes independent of cell size. In sum, quantitative analysis relating cell, nuclear, and chromosome size predicts two levels of chromosome length regulation: one through cell size and a second in response to nuclear size.     

Geographical variation in egg size of the Great Tit Parus major: a new perspective

A recent study on geographical variation in egg size of Great Tits Parus major concluded that: (1) mean egg size tended to increase with increasing latitude; and (2) mean egg size was positively correlated with mean clutch size. Including new data on both egg and clutch size, we reanalysed the relationships between egg size, clutch size and latitude, and investigated the possible effects of habitat type, female body size and egg shape on these relationships. We found that (1) egg volume showed minimum values around 51°N, increasing both north and southwards; (2) female body size increased linearly with increasing latitude; (3) female body size was positively correlated with egg breadth, but not with egg length or egg volume; (4) the sphericity index of the eggs (breadth to length ratio) was largest at medium latitudes, and eggs were more elongated towards the north and the south; (5) the relationship between clutch size and latitude was curvilinear, with the largest clutch sizes at intermediate latitudes; (6) egg size was not correlated with clutch size when the complete latitudinal range was considered, but egg size was negatively correlated with clutch size between 40 and 51°N; and (7) egg size did not differ among habitat types. We suggest that female body size (which probably limits egg breadth), and the pressure for producing large eggs (which in turn increases the reproductive success) are the main determinants of geographical variation in egg size and shape. Populations of small-bodied Great Tits seem to escape from the limits of their size, producing relatively elongated eggs, so that from a certain latitude southwards, egg volume does not decrease in spite of a decrease in female body size. Moreover, the negative relationship between egg and clutch size at low latitudes suggests that energetic trade-offs may also contribute to determine egg size in the south.     

Ejaculate size in bushcrickets: the importance of being large

Sexual size dimorphism is common in animals. Females, which often experience increased fecundity with size, are generally the larger sex. In males, the pre-copulatory advantages of large size have been extensively investigated. However, in polyandrous species, large male size may also be advantageous after copulation due to sperm competition. Male size co-varies with ejaculate size and increased fertilization success in many insects. Large male size can also be important for production of nuptial gifts enhancing female fecundity. In this study, the relative importance of male size for ejaculate and nuptial gift production is assessed in 20 bushcricket species (Orthoptera: Tettigoniidae). Relative male size is found to correlate with increased ejaculate size across taxa. In contrast, no association is found between relative male size and the size or quality of the nuptial gift. These findings indicate that selection for increased ejaculate volumes as a response to sperm competition may act to promote large male size in insects.     

SIZE‐DEPENDENT MORTALITY AND COMPETITION INTERACTIVELY SHAPE COMMUNITY DIVERSITY

Body size is recognized as a major factor in evolutionary processes mediating sympatric diversification and community structuring. Life‐history types with distinct body sizes can result from two fundamental mechanisms, size‐dependent competition and size‐dependent mortality. While previous theoretical studies investigated these two processes in separation, the model analyzed here allows both selective forces to affect body‐size evolution interactively. Here we show for the first time that in the presence of size‐dependent competition, size‐dependent mortality can give rise to multiple, coexisting size morphs representing the final outcomes of evolution. Moreover, our results demonstrate that interactions between size‐dependent competition and mortality can create characteristic abrupt changes in size structure and nonmonotonic patterns of biological diversity along continuous and monotonic environmental gradients. We find that the two selective forces differentially affect the body‐size ratios of coexisting morphs: size‐dependent competition results in small and relatively constant ratios, whereas size‐dependent mortality can open niches for morphs that greatly differ in body size. We show that these differential effects result in characteristic distributions of size ratios across communities, which we suggest can help detect the concurrent action and relative influence of size‐dependent competition and mortality in nature.     

SEXUAL SIZE DIMORPHISM AND EGG-SIZE ALLOMETRY IN BIRDS

We tested the hypothesis that egg size should evolve in sexually dimorphic birds to reduce costs associated with more rapid growth by nestlings of the larger sex. Consistent with this hypothesis, we found that in species in which males were larger, females laid proportionately larger eggs as sexual size dimorphism increased. However, this result was also consistent with the hypothesis that egg size varied allometrically with both male and female body size. Furthermore we found that in species in which females were larger, relative egg size decreased as size dimorphism increased, which is consistent with the “allometry hypothesis” but not the “cost-reduction hypothesis. That male body size contributes to the allometric relationship between egg size and body size suggests that the basis for the allometric relationship is not wholly a mechanical one stemming from the physical requirements of developing, transporting, and laying an egg of a particular size. Rather, the relationship seems likely to be tied more directly to body size itself the tact that male body size influences a female trait suggests that egg size–body size relationships otter some scope for investigating the basis for allometric relationships in general.     

Patterning the size and number of tooth and its cusps

Mice and rats, two species of rodents, show some dental similarities such as tooth number and cusp number, and differences such as tooth size and cusp size. In this study, the tooth size, tooth number, cusp size and cusp number, which are four major factors of the tooth patterning, were investigated by the heterospecific recombinations of tissues from the molar tooth germs of mice and rats. Our results suggest that the dental epithelium and mesenchyme determine the cusp size and tooth size respectively and the cusp number is co-regulated by the tooth size and cusp size. It is also suggested that the mesenchymal cell number regulates not the tooth size but the tooth number. The relationships among these factors in tooth patterning including micropatterning (cusp size and cusp number) and macropatterning (tooth size and tooth number) were analyzed in a reaction diffusion mechanism. Key molecules determining the patterning of teeth remains to be elucidated for controlling the tooth size and cusp size of bioengineered tooth.     

Life history evolution in cichlids 2: directional evolution of the trade-off between egg number and egg size

The negative relationship between offspring number and offspring size provides a classic example of the role of trade-offs in life history theory. However, the evolutionary transitions in egg size and clutch size that have produced this negative relationship are still largely unknown. Since body size may affect both of these traits, it would be helpful to understand how evolutionary changes in body size may have facilitated or constrained shifts in clutch and egg size. By using comparative methods with a database of life histories and a phylogeny of 222 genera of cichlid fishes, we investigated the order of evolutionary transitions in these traits in relation to each other. We found that the ancestral large-bodied cichlids first increased egg size, followed by a decrease in both body size and clutch size resulting in the common current combination of a small-bodied cichlid with a small clutch of large eggs. Furthermore, lineages that deviated from the negative relationship between clutch and egg size underwent different transitions in these traits according to their body size (large bodied genera have moved towards the large clutch/small egg end of the continuum and small bodied genera towards the small clutch/large egg end of the continuum) to reach the negative relationship between clutch size and egg size. Our results show that body size is highly important in shaping the negative relationship between clutch size and egg size.     

Allometry and selection in a novel predator–prey system: Australian snakes and the invading cane toad

Because many organismal traits vary with body size, interactions between species can be affected by the respective body sizes of the participants. We focus on a novel predator–prey system involving an introduced, highly toxic anuran (the cane toad, Bufo marinus ) and native Australian snakes. The chance of a snake dying after ingesting a toad depends on the size of the snake and the size of the toad, and ultimately reflects the effect of four allometries: (1) physiological tolerance (the rate that physiological tolerance to toad toxin changes with snake size); (2) swallowing ability (the rate that maximal ingestible toad size (i.e. snake head size) increases with snake body size); (3) prey size (the rate that prey size taken by snakes increases with snake head size) and (4) toad toxicity (the rate that toxicity increases with toad size). We measured these allometries, and combined them to estimate the rate at which a snake's resistance changes with toad toxicity. The parotoid glands (and thus, toxicity) of toads increased disproportionately with toad size (i.e. relative to body size, larger toads were more toxic) but simultaneously, head size relative to body size (and thus, maximal ingestible prey size relative to predator size) declined with increasing body size in snakes. Thus, these two allometries tended to cancel each other out. Physiological tolerance to toxins did not vary with snake body size. The end result was that across snake species, mean adult body size did not affect vulnerability. Within species, however, smaller predators were more vulnerable, because the intraspecific rate of decrease in relative head size of snakes was steeper than the rate of increase in toxicity of toads. Thus, toad invasion may cause disproportionate mortality of juvenile snakes, and adults of the sex with smaller mean adult body sizes.     

How host size may constrain the evolution of parasite body size and clutch size. The parasitic isopod Ichthyoxenus fushanensis and its host fish, Varicorhinus bacbatulus, as an example

The flesh-burrowing parasitic isopod Ichthyoxenus fushanensis was found infecting the body cavity of a freshwater fish, Varicorhinus bacbatulus , in heterosexual pairs. Herein we investigate the question of how the host body size may constrain the parasite size and clutch size by analyzing the interactions among the body size and clutch size of the parasite, and host size. Due to the low transmission rate of I. fushanensis to its host and the positive relationship between clutch size and female size, selection may favor larger females with larger clutch sizes to compensate for massive losses of manca (the free-living juveniles). The path model reveals that clutch size depends not only directly on female size, but also on the sizes of her host and mate. Female size also depends on the sizes of the host and her mate. A negative correlation exists between the body sizes of the paired males and females. This negative correlation may be regarded as a consequence of competition for limited available space or other resources provided by the host. The effects of host size on parasite size, however, act on the total volume of both sexes as a whole, not specifically on either the female or the male. In this case, the available space/resources may not allow both individuals of different sexes to evolve toward a larger size simultaneously. Under the constraint of host size, a strategy of reducing the body size of the paired male may provide a way to increase the body size of the paired female and achieve a larger clutch size.     

Life histories as sampling devices: optimum egg size in pelagic fishes*

In teleosts, egg size is an important determinant of larval size. The latter determines where larvae start feeding in the size spectrum of biomass particles. Egg and larval size also determine the size range of predators exploiting these early ontogenetic stages. Thus, egg size is important in determining the particle size range sampled during ontogeny. However, egg size also largely determines how many can be produced by a female. Optimum egg size for marine, pelagic fishes was estimated using a modification of a model developed by Ware. A numerical analysis using empirical information on size and temperature dependent mortality and growth in pelagic, marine systems suggests that allometry of size-dependent mortality and temperature are significant factors in determining optimum egg size. Allometry of mortality will depend on the frequency of encounters between particles of different sizes. Both empirical and modelling studies of such frequencies in pelagic and other environments are required.     

Unit pricing: Economics and the evolution of litter size

Variability in litter size and the concept of optimality are central to our current understanding of parental investment patterns and life histories. A fundamental component of most models of optimum litter size is an apparently inescapable trade-off between litter size and size of offspring. Most previous models of litter size have focused on the evolution of an optimum litter size rather than variability in litter size. Because variability provides the raw material from which numerous optimal litter sizes are fashioned to meet prevailing conditions, approaches that specifically address adaptive patterns of variability might provide new insights into the evolution of litter size. A model based on a non-linear relationship between total parental effort and litter size reduces trade-offs between offspring size and offspring number to simple reproductive economics, and illustrates how litter size variability might be predictable under certain environmental conditions. This revised version was published online in July 2006 with corrections to the Cover Date.