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.     

Effect of salmon carcasses on egg production of a freshwater amphipod, Jesogammarus jesoensis: a field observation

The effect of salmon carcasses on egg production of a freshwater amphipod, Jesogammarus jesoensis, was examined by comparing populations with and without a supply of carcasses in Naibetsu River in Hokkaido, northern Japan. The female body length in the population with carcasses became larger in the years when carcasses were abundant, being significantly larger than that in the population without carcasses. No evidence, however, was found that carcasses influenced the ratio of brooding females to mature females. Every year, the clutch size in the population with carcasses was larger than the clutch size without carcasses. Because clutch size showed a significant correlation with female size, the larger clutch size was considered to be a result of the larger female size. In addition, however, comparison of clutch size using analysis of cavariance revealed that clutch size in the population with carcasses was larger than that without carcasses, even for the same body size. Egg size showed no difference between populations and for different years. These results indicate that salmon carcasses influence egg production of J. jesoensis by increasing female body size and by increasing the investment in each clutch per unit of body size.     

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.     

Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle

Body size of many animals varies with latitude: body size is either larger at higher latitudes (Bergmann's rule) or smaller at higher latitudes (converse Bergmann's rule). However, the causes underlying these patterns are poorly understood. Also, studies rarely explore how sexual size dimorphism varies with latitude. Here we investigate geographic variation in body size and sexual size dimorphism of the seed-feeding beetle Stator limbatus, collected from 95 locations along a 38 degrees range in latitude. We examine 14 variables to test whether clines in environmental factors are adequate to explain geographic patterns of body size. We found that body size and sexual size dimorphism of S. limbatus varied considerably with latitude; beetles were smaller but more dimorphic at lower latitudes. Body size was not correlated with a gradient in mean temperature, contrary to the commonly accepted hypothesis that clines are produced by latitudinal gradients in temperature. Instead, we found that three factors were adequate to explain the cline in body size: clinal variation in host plant seed size, moisture (humidity), and seasonality (variance in humidity, precipitation, and temperature). We also found that the cline in sexual size dimorphism was partially explainable by a gradient in moisture, though moisture alone was not sufficient to explain the cline. Other ecological or environmental variables must necessarily contribute to differences in selection on male versus female body size. The main implications of our study are that the sexes differ in the magnitude of clinal variation in body size, creating latitudinal variation in sexual size dimorphism, and that clines in body size of seed beetles are likely influenced by variation in host seed size, water availability, and seasonality.     

Sex ratio, life-history invariants, and patterns of sex change in a family of protandrous gastropods

Application of optimality theory to the evolution of life histories has been broadly successful in predicting the conditions favoring sex change, the type of change, and the timing of such changes. The size advantage hypothesis predicts that the optimal size at which an individual should change sex is a function of its size and the size and sex of its potential mates. I collected data on the size, sex, and grouping of individuals of 27 populations of 19 species of the calyptraeids, a family of protandrous marine gastropods that includes Crepidula. These data are used to test the following predictions about variation in size at sex change: (1) sex ratio is biased toward the first sex; (2) the ratio of the size at sex change to the maximum size is a life-history invariant; and (3) species that form variable-sized groups or stacks have more variation in size at sex change than species that show less variation in stack formation. Across all 19 species, sex ratio was not significantly more often biased toward the first sex than it was toward the second sex, although sex ratios were significantly male biased more often than they were significantly female biased. Sex ratios ranged from 0.05 to 0.91, and this variation was related to mode of development, skew in size distribution, and frequency of stacking, but not with maximum body size. There was little evidence that the ratio of size at sex change and maximum size is invariant. There is evidence that one of the main underlying assumptions of this life-history invariant, that male fertility increases with the same function of size in all species, is invalid for calyptraeids and probably for other animals. Finally, species that form larger stacks or mating groups had more variation in size at sex change within a population than species that were generally solitary. These results suggest that information about individual groupings should be included in predictions of life-history theory and that more information about the relationship between male fitness and size is also needed.     

Allocation of offspring size and sex by female black ratsnakes

How females allocate resources to each offspring and how they allocate the sex of their offspring are two powerful potential avenues by which mothers can affect offspring fitness. Previous research has focussed extensively on mean offspring size, with much less attention given to variance in offspring size. Here we focussed on variation in offspring size in black ratsnakes, Elaphe obsoleta . We collected and hatched 105 clutches (1283 eggs) over 9 years. We predicted that females should lay larger eggs, or more variable eggs, when the environment is less predictable. We also predicted that females laying early or laying larger eggs should produce mostly sons because adult males are larger than adult female ratsnakes. The largest hatchling was more than twice the length and almost four times the mass of the smallest hatchling. Variation in offspring size was itself highly variable, with CVs in offspring mass among clutches ranging from 1% to 25%. With one exception, the variables we expected should influence variation in offspring size had little effect. We found that clutch size increased with maternal size and that egg size decreased with clutch size, but we found no evidence that variance in egg size among clutches increased as the season progressed or that females increased the mean size of their offspring the later in the season they laid their eggs. Females in better condition after they finish laying their eggs did produce larger eggs. There was no relationship between within-clutch variation in egg size and laying date or mean egg size. Finally, sex ratio did not vary with mean egg size or hatching date. Given evidence that offspring size in snakes affects survival, selection should reduce variation in offspring size unless that variance enhances maternal fitness and yet we found little support for hypothesized advantages of varying offspring size.     

The effect of egg size and habitat on starling nestling growth and survival

In spite of the fact that hatchling size and energy reserves in birds are affected by egg size, many studies have failed to find an effect of egg size on offspring fitness. One possibility is that this is because they have been performed in areas with high food availability and that effects of egg size on offspring fitness are most apparent in areas of low food availability. To investigate this, egg size,␣offspring mass and survival of European starlings (Sturnus vulgaris) were measured in an agricultural landscape with a low but variable amount of pasture, the preferred foraging habitat of parent starlings. Offspring mass was related to egg size, but egg size explained a declining proportion of the variation in nestling mean mass as nestlings grew older. Offspring survival during the early, but not during the late nestling period was related to egg size. Throughout the nestling␣period, survival was related to the mass of the nestlings. It is suggested that the effect of egg size on␣offspring survival is through the effect of egg size on offspring mass, this effect declining as offspring grow older. Offspring survival during the early part of the nestling period was related to egg size when availability of pasture was low, but not when it was high. However, the interaction was not significant. Selection for␣larger egg size is discussed in relation to the structuring␣of starling populations into sources and sinks. Received: 22 September 1997 / Accepted: 22 January 1998     

The evolution of large brain size in birds is related to social,not genetic,monogamy

There are several hypotheses suggesting that social complexity, including pair bonding, is important in the evolution of increased brain size. I examined whether genetic or social monogamy was related to large brain size in birds. Recent work has indicated that the length and strength of pair bonds are associated with large brain size. I tested several hypotheses for the evolution of large brain size in 42 species of bird by including life history variables in a regression model. A test on 100 phylogenetic trees revealed no phylogenetic signal in brain size. Controlling for body size, a principal components analysis was run on the life history variables and degrees of extra‐pair paternity. The main principal component (PC1) was regressed on brain size revealing a strong, positive association. Social, but not genetic, monogamy was positively related to brain size. Large brain size is related to the selective pressures of procuring extra‐pair copulations whilst maintaining a social partnership. However, other life history variables also loaded positively and significantly on brain size. These results indicate that the evolution of large brain size in birds was driven by several important selective pressures. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 668–678.     

Relationship between prey consumption and colony size in an orb spider

Summary I studied the relationship between prey consumption and colony size in the orb spiderPhiloponella semiplumosa. Observations of unmanipulated colonies showed that prey biomass per juvenile spider was positively correlated with colony size, indicating that prey consumption was highest in the largest colonies observed. In contrast, the relationship between prey biomass per adult female and colony size was curvilinear; prey consumption tended to be highest in intermediatesized colonies. Adult female cephalothorax width was positively correlated with colony size. Number of egg sacs per adult female tended to be highest in intermediate-sized colonies. Prey biomass per juvenile was lower in experimentally reduced colonies than in large control colonies. Aerial-arthropod abundance was not correlated with colony size, and experimental prey supplementation did not affect colony size. Thus, the relationship between prey consumption and colony size was influenced by coloniality directly, rather than by a correlation between prey abundance at a site and colony size.     

Influence of genetic and environmental factors on egg and clutch sizes among populations of Paratya australiensis Kemp (Decapoda: Atyidae) in␣upland rainforest streams, south-east Queensland

Headwater populations of the common shrimp Paratya australiensis were sampled to examine the factors influencing egg and clutch size. Much of the spatial variation in these reproductive traits was associated with differences in altitude, with upper sites having larger eggs and smaller clutches. Mean egg size at high-altitude sites was higher than that previously reported for this species. Temporal variation in egg and clutch size was also observed, with significant increases in egg size during the breeding season. At most sites, this increase was accompanied by a decrease in clutch size. The combined effect of the reciprocal patterns in egg volume and clutch size resulted in relatively little spatial and temporal variation in reproductive effort. In a field experiment, using a unique genetic marker, shrimps were translocated between two sites with significantly different egg sizes. After one generation (18 months), the mean egg size of translocated females was the same as that of females from the `source' population. In contrast, clutch size changed towards that of the resident females. This suggests that egg size is under strong genetic control, while clutch size is influenced by the environment. Received: 14 July 1997 / Accepted: 9 March 1998     

Number and size of clutch in Ptomascopus morio (Coleoptera: Silphidae: Nicrophorinae), with comparisons to Nicrophorus

Ptomascopus morio displays simpler parental care than Nicrophorus species. The effects of carcass size and clutch number on clutch size in P. morio were examined. Clutch size was related to carcass size. There was a negative correlation between number of clutch and clutch size for most sizes of carcass. Longevity of females was shorter when the carcass size was larger, such that total lifetime fecundity did not differ among carcasses of different sizes. The clutch size of P. morio was larger than that of Nicrophorus quadripunctatus. The clutch size of P. morio declined rapidly with repeated clutch production, but that of N. quadripunctatus was rather constant. This indicates that N. quadripunctatus maintains a more constant clutch size than P. morio over several reproductive attempts, although the former displays more complex parental care.     

堆肥处理对污泥腐殖物质形态及其重金属分配的影响

采用透析、凝胶色谱 (SephadexG 75 )研究了污泥堆肥前后腐殖质分子大小的变化及重金属Cu和Zn在各级组分中的分配。透析结果表明 ,污泥经过堆腐以后 ,腐殖质中小分子物质 (<10 0 0Da)组分的含量下降 6 4 % ,而相对高分子组分 (>2 5 0 0 0Da)却增加了 6 8%。凝胶色谱进一步证实 ,污泥经过 4 9d堆腐后 ,腐殖质中大于 2 0 0 0KDa的大分子组分是堆肥起始时的2 3倍。而小分子组分明显减少 ,表现在小分子组分的凝胶洗脱体积明显减少。堆肥腐熟以后 ,腐殖质吸附的Cu、Zn元素含量增加 ,其中Cu主要被吸附在大分子物质上 ,而Zn主要与小分子物质结合     

Visual Assessment of Relative Body Size in a Cichlid Fish,the Oscar,Astronotus ocellatus

In cichlid fishes, fighting ability, success at nest defense, and fecundity are all positively correlated with body size. Selection should therefore favor the evolution of strategies for the assessment of relative body size during intraspecific interactions. The effect of relative body size on agonistic behavior in the oscar (Astronotus ocellatus) was tested using dummies. As predicted by game theory models, attack was strongly correlated with the size of the dummy relative to the subject. The most intense aggression occurred when dummy size was approximately 75% of subject size. These data support the prediciton that perceived body size asymmetries are important determinants of agonistic behavior in these fishes. Further, the results suggest that oscars can visually assess relative body size, and may estimate relative fighting ability without tests of strength. Subject size was positively correlated with intensity of aggressive behavior. Attack and display frequencies were positively correlated with a subject's body size, while latencies to approach and attack dummies were negatively correlated with subject size.     

蜡皮蜥的两性异形和繁殖输出

为研究蜡皮蜥(Leiolepis reevesii)两性异形和繁殖输出,于2002、2003年4月下旬从海南乐东一种群捕获423头蜡皮蜥。经检测得到繁殖雌体的最小体长为89．0mm,据此判定≥89．0mm的个体为性成熟。研究结果表明：①蜡皮蜥具有两性异形,雄性大于雌性且具有较大的头部。成体雄性头长和头宽随体长的增长速率大于雌性,幼体头长和头宽随体长的增长速率无显著的两性差异。以性别和年龄(成、幼体)为因子的双因子ANOVA比较两性头长和头宽与体长的回归剩余值发现,雄性头部大于雌性,幼体头部相对大于成体。②饲养于实验室的母体中有42头于2002、2003年5月22日～7月16日产出正常卵,这些繁殖雌体具有年产多窝卵的潜力。窝卵数和卵重的变异系数分别为0．18和0．13,前者变异度大于后者。窝卵数、窝卵重和卵重均与母体体长无关。卵重与相对生育力之间无显著的负相关性,表明蜡皮蜥缺乏卵数量与卵大小之间的权衡。相对窝卵重与母体体长呈显著的负相关,表明较小的母体具有相对较大的繁殖输出。因雌体繁殖会滞缓其生长,小母体具有相对较大的繁殖输出,至少部分地解释了雌性蜡皮蜥的成体为什么个体较小。     

Relationships between oviposition date, hatch date, and offspring size in the grasshopper Chorthippus brunneus

Abstract 1. The grasshopper Chorthippus brunneus has been shown to increase egg size with maternal age under constant laboratory conditions, such that late-laid eggs are larger than early laid eggs. In this study, an increase in the size of eggs (and hatchlings from those eggs) was recorded with date of oviposition in a field cage population. This suggests that the relationship between maternal age and egg size, observed previously in the laboratory, also occurs in the field. There was, however, some evidence that the behaviour of the maternal females in field cages was modified by the onset of autumnal weather at the end of the breeding season. Only a small number of eggs was laid in the last 3 weeks of the summer but these yielded relatively small hatchlings.
2. The date on which eggs were laid was correlated positively with their date of hatch in the following year. A consequence of this relationship, and that between oviposition date and egg size, was that size at hatch increased with date of hatch through most of the spring but then declined as the last few eggs hatched. Temporal variation in size at hatch parallelled temporal variation in the maternal females' investment in egg size in the previous year.
3. An undisturbed field population was monitored to assess whether the temporal variation in size at hatch resulted in size variation at eclosion in subsequent developmental stages. There were negative correlations between size at eclosion and date of eclosion at the third and fourth instars, suggesting that despite their smaller size at hatch, early hatchers experienced more favourable conditions for juvenile growth than late hatchers. The larger size at hatch of late hatchers may enhance their survival and compensate (albeit incompletely) for their reduced opportunity for growth. Female reproductive behaviour may represent an adaptive response to a predictable seasonal decline in offspring fitness at hatch.     

Regulation of brood size in a burying beetle,Nicrophorus tomentosus (Silphidae)

Regulation of brood size in a biparental burying beetle, Nicrophorus tomentosusWeber, was studied by providing pairs with one of two sizes of mouse carcasses in the laboratory. For a given carcass size, there was an inverse relationship between number and mass offspring in a brood. The requirement for regulation was that brood size was adjusted such that mean mass of individual larvae was constant for carcasses of different size. Brood size was regulated if parents were present but regulation did not occur if parents were removed prior to hatching of larvae. Pairs bred in quick succession on two carcasses raised fewer than the regulated number of young in the second reproductive attempt. Reasons for regulation of brood size in this genus are discussed.     

Patterns in body size and melanism along a latitudinal cline in the wingless grasshopper,Phaulacridium vittatum

Aim We explore geographic variation in body size within the wingless grasshopper, Phaulacridium vittatum, along a latitudinal gradient, and ask whether melanism can help explain the existence of clinal variation. We test the hypotheses that both male and female grasshoppers will be larger and lighter in colour at lower latitudes, and that reflectance and size will be positively correlated, as predicted by biophysical theory. We then test the hypothesis that variability in size and reflectance is thermally driven, by assessing correlations with temperature and other climatic variables. Location Sixty‐one populations were sampled along the east coast of Australia between latitudes 27.63° S and 43.10° S, at elevations ranging from 10 to 2000 m a.s.l. Methods Average reflectance was used as a measure of melanism and femur length as an index of body size for 198 adult grasshoppers. Climate variables were generated by BIOCLIM for each collection locality. Hierarchical partitioning was used to identify those variables with the most independent influence on grasshopper size and reflectance. Results Overall, there was no simple relationship between size and latitude in P. vittatum. Female body size decreased significantly with latitude, while male body size was largest at intermediate latitudes. Rainfall was the most important climatic variable associated with body size of both males and females. Female body size was also associated with radiation seasonality and male body size with reflectance. The reflectance of females was not correlated with latitude or body size, while male reflectance was significantly higher at intermediate latitudes and positively correlated with body size. Analyses of climate variables showed no significant association with male reflectance, while female reflectance was significantly related to the mean temperature of the driest quarter. Main conclusions Geographic variation in the body size of the wingless grasshopper is best explained in terms of rainfall and radiation seasonality, rather than temperature. However, melanism is also a significant influence on body size in male grasshoppers, suggesting that thermal fitness does play a role in determining adaptive responses to local conditions in this sex.     

The coadaptation of female morphology and offspring size: a comparative analysis in crickets

Few studies of invertebrates have considered combinations of morphological and life history traits in the context of the evolution of reproductive strategies. Cricket species that exploit habitats harsh with respect to egg survival have evolved a long ovipositor, presumably because laying deep in the soil reduces egg mortality. Yet hatchling mortality increases with laying depth, and the ability of hatchlings to climb through the soil increases with egg size. Thus a conflict may exist between survival of the egg and that of the hatchling, inducing a positive covariation between egg size and ovipositor length across species evolving under contrasting selective habitats. We used the phylogenetic autocorrelation method and a path analysis to assess whether egg size coevolved with ovipositor length across 40 species of crickets, and whether egg size was affected by body size or ecological factors that influence egg mortality. Body size and ovipositor length were affected by taxonomic association, whereas common ancestry had no significant effect on egg size, diapausing strategy, and oviposition preference for soil types. The path model indicated that 29.11% of the variance in egg size was explained by independent evolution. As expected, ovipositor length was positively correlated with egg size, and species diapausing in the egg stage produced larger eggs than crickets diapausing in the nymphal stage or with no diapause. Ovipositor length and diapausing strategy were the first and second most important traits, respectively, in term of the proportion of variance in egg size explained by specific values. These results support the hypothesis that the ability of hatchlings to climb through the soil, and variation in diapause strategies, are general selective factors affecting the evolution of egg size in crickets. Phylogeny explained 51.01% of the variance in egg size. Egg size in a current cricket species, however, was not directly determined by egg size in its ancestor. Instead, it was strongly related to the phylogenetic values of body size and ovipositor length. Such indirect phylogenetic effects of body size and ovipositor length may have arisen because clades originating from ancestors with different ovipositor lengths experienced different selective pressures on egg size. Recelived: 13 October 1995 / Accepted: 30 September 1996     

Litter size at birth in purebred dogs--a retrospective study of 224 breeds

Despite the long history of purebred dogs and the large number of existing breeds, few studies of canine litter size based upon a large number of breeds exist. Previous studies are either old or include only one or a few selected breeds. The aim of this large-scale retrospective study was to estimate the mean litter size in a large population of purebred dogs and to describe some factors that might influence the litter size. A total of 10,810 litters of 224 breeds registered in the Norwegian Kennel Club from 2006 to 2007 were included in the study. The overall mean litter size at birth was 5.4 (± 0.025). A generalized linear mixed model with a random intercept for breed revealed that the litter size was significantly influenced by the size of the breed, the method of mating and the age of the bitch. A significant interaction between breed size and age was detected, in that the expected number of puppies born decreased more for older bitches of large breeds. Mean litter size increased with breed size, from 3.5 (± 0.04) puppies in miniature breeds to 7.1 (± 0.13) puppies in giant breeds. No effect on litter size was found for the season of birth or the parity of the bitch. The large number of breeds and the detail of the registered information on the litters in this study are unique. In conclusion, the size of the breed, the age of the bitch and the method of mating were found to influence litter size in purebred dogs when controlling for breed, with the size of the breed as the strongest determinant.