Convariation of Spider Egg and Clutch Size: The Influence of Foraging and Parental Care

Reproduction in a Northern Population of Clemmys guttata

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.

Early and late season clutch parameters were examined over a three year period in the Florida scrub lizard, Sceloporus woodi. Precipitation levels were monitored throughout the study. In the early and late season of 1984 and the early season of 1986 precipitation levels approximated long-term mean levels of precipitation. In 1985 a severe winter drought occurred. Clutch size was positively related to body size in all samples in every year. In 1984 and 1986, egg size was not related to clutch size, whereas, in 1985 egg size was negatively related to clutch size. In 1985, females produced large clutches of small eggs early in the season and small clutches of large eggs late in the season. In 1984, no seasonal changes in egg or clutch size occurred. In the late season of 1986, females produced the largest clutches and the smallest eggs of all the samples, but egg and clutch size were not statistically different from the early season egg and clutch size of 1986. Total clutch dry weight, an estimate of total clutch energy, was not different in any of the six sampling periods. These data do not support current adaptationist models that attempt to explain the control of clutch and egg size in lizards. It is argued in this paper that egg and clutch size may vary in response to past environments that affect a female's physical condition, as well as, current resources that may be important for maintenance and reproduction. Egg and clutch size appear to be plastic traits selected to respond to proximal environmental variation, whereas, the investment of total dry matter/clutch has been optimized.

Variation in egg and clutch size of the Black Redstart (Phoenicurus ochruros) at the northeastern edge of the Qinghai-Tibetan Plateau

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.

Reproductive and body size data from 169 populations of 146 species (56% of those recognized), 65 genera (75%), and 11 families (92%) of turtles were tabulated to test for latitudinal variation in egg and clutch size. Body-size-adjusted correlation analysis of all populations (as well as within most families) revealed (i) a significant negative relationship (r2 = 0.26) between latitude and egg size, (ii) a significant positive relationship (r2 = 0.21) between latitude and clutch size, and (iii) no relationship between latitude and clutch mass. Phylogenetic contrast analyses corroborated these patterns. Clutch size was also negatively correlated with egg size across all populations as well as within most families. We evaluate the applicability to turtles of hypotheses postulated to explain such latitudinal patterns for other vertebrate groups. The observed pattern may be the result of latitudinal variation in selection on egg size and (or) clutch size, as well as on the optimal trade-off between these two traits.

BackgroundEgg size and clutch size are key life history traits. During the breeding period, it is possible for females to increase their reproductive output either by increasing the number of eggs if the optimal egg size (OES) is maintained, or by increasing the allocation of energy to each egg. However, the strategies adopted are often influenced by animals’ morphology and environment.MethodsHere, we examined variation in female morphological and reproductive traits, tested for trade-offs between egg size and clutch size, and evaluated the relationship between egg size and female morphology in three populations of Phrynocephalus helioscopus.ResultsFemale body size, egg size, and clutch size were larger in the Yi Ning (YN) and Fu Yun (FY) populations than in the Bei Tun (BT) population (the FY and YN populations laid more, and rounder eggs). Egg size was independent of female body size in two populations (BT and FY), even though both populations had an egg-size/clutch size trade-off. In the YN population, egg size and clutch size were independent, but egg size was correlated with female body size, consistent with the hypothesis of morphological constraint.ConclusionsOur study found geographical variation in body size and reproductive strategies of P. helioscopus. Egg size was correlated with morphology in the larger-bodied females of the YN population, but not in the smaller-bodied females of the BT population, illustrating that constraints on female body size and egg size are not consistent between populations.

BackgroundThe evolution of sexual size dimorphism (SSD) is likely constrained by life history. Using phylogenetic comparative methods, we examined correlations between SSD among anurans and their life history traits, including egg size, clutch size, mating combat, and parental care behaviour. We used sexual dimorphism index (SDI = Body-sizefemale /Body-sizemale –1) as the measurement for SSD. Body size, life history and phylogenetic data were collected from published literature. Data were analysed at two levels: all anuran species and within individual families.ResultsFemale-biased SSD is the predominant form in anurans. SSD decreases along with the body size increase, following the prediction of Rensch’s rule, but the magnitude of decrease is very small. More importantly, female body size is positively correlated with both fecundity related traits, egg size and clutch size, and SDI is also positively correlated with clutch size, suggesting fecundity advantage may have driven the evolution of female body size and consequently leads to the evolution of female-biased SSD. Furthermore, the presence of parental care, male parental care in particular, is negatively correlated with SDI, indicating that species with parental care tend to have a smaller SDI. A negative correlation between clutch size and parental care further suggests that parental care likely reduces the fecundity selection pressure on female body size. On the other hand, there is a general lack of significant correlation between SDI and the presence of male combat behaviour, which is surprising and contradictory to previous studies.ConclusionsWe find clear evidence to support the ‘fecundity advantage hypothesis’ and the ‘parental care hypothesis’ in shaping SSD in anurans. Nevertheless, the relationships of both parental care and combat behaviour to the evolution of SSD are complex in anurans and the extreme diversity of life history traits may have masked some potential interesting relationships. Our study represents the most comprehensive study of SSD in anurans to date.

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.

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.

Environmental variation along altitudinal gradients can promote life‐history trait differentiation in ectothermic animals. Life‐history theory predicts that increased environmental stress results in a shift in reproductive allocation from offspring quantity to quality and a stronger trade‐off between egg size and clutch size. To test this prediction, we investigated patterns of variation in life‐history traits (i.e. age, body size, clutch size and egg size) among four populations of Bufo andrewsi from Baoxing County, western China, at different altitudes. We found that body size, age, egg size and total reproductive output, but not clutch size, differed between populations. Clutch size and total reproductive output increased with female size and age. However, egg size decreased with female size and did not change with female age. The egg size and clutch size trade‐off was evident for all populations except at lowest altitude, and the strength of trade‐off between egg size and clutch size increased with altitude. Our findings suggest that environmental constraints at high altitude select for investment in larger eggs at a cost of offspring number.

Variation in avian clutch and egg sizes across geographical gradients has long been debated, particularly regarding the mechanisms underlying latitudinal clines in reproductive investment. These patterns are primarily driven by climatic conditions, with latitude acting as a proxy for environmental gradients. Here, we assembled a robust dataset of eggs and clutches of eight species of kingbird flycatchers (Tyrannus) covering a broad geographic range to test the hypothesis that clutch and egg size vary systematically along environmental gradients, and predict that egg and clutch size would (a) increase with latitude, (b) be larger in more seasonal climate categories, (c) be larger at sites with cooler winters, and (d) be larger at sites with warmer and wetter long-term climatic conditions. We analysed clutch and egg data from 35 scientific egg collections representing the geographically widespread genus Tyrannus. After applying spatial, temporal and taxonomic data quality controls, we analysed the relationships between clutch and egg size and climatic variables. The analyses of 1332 clutches and 4680 eggs of eight species collected during 158 years (1858-2016) showed that clutch and egg sizes increased toward higher latitudes. Both traits varied according to major climate types, regional subclimates, and local temperature and precipitation conditions. More seasonal regions had the largest clutches, although sites with colder winters did not have the largest clutches. Egg size was larger at sites with less severe dry periods, and larger eggs were also associated with lower temperatures. Overall, residual variation in clutch and egg size is strongly associated with climatic conditions at both regional and local scales. These findings contribute to understanding how avian reproductive traits respond to climatic gradients and may help anticipate how species will respond to future climate scenarios.

In Algonquin Park, Ontario, body size and clutch characteristics were recorded for 51 female painted turtles (Chrysemyspicta) in 1983, 61 in 1984, and 24 in 1985. Clutch size, clutch mass, and egg width correlated significantly with body size (carapace length) in all 3 years. Egg length and egg mass were significantly related to body size in 1984 and 1985, but not in 1983. There were no significant correlations of egg width or egg mass to clutch size. For a group of the same individuals compared by repeated-measures ANOVA, mean clutch mass and mean egg size, but not mean clutch size, varied significantly among years. Correlation of egg size with body size, lack of correlation between egg size and clutch size, and annual variation in egg size, but not clutch size, all fail to support current versions of optimal egg size theory. Twenty-six females nested in both 1983 and 1984 and 11 females nested in both 1984 and 1985. Fourteen females nested twice in 1 year: six in 1983 and eight in 1984. Between 43 and 73% of adult females nested in a given year and 12–13% nested twice in a single season. These estimates are similar to those reported for other populations of this species. It appears that variations in both clutch size (frequency) and egg size are important sources of variation in reproductive output.

Intraspecific variation in embryonic and larval traits of the dorid nudibranch mollusc Adalaria proxima (Alder and Hancock) around the northern coasts of the British Isles

Interspecific patterns for egg and clutch sizes of African Bufonidae (Amphibia: Anura)