Latitudinal and seasonal patterns in clutch size of some single‐brooded British birds

Capsule Daylength, rather than latitude, was found to be an important determinant of variation in clutch size.

Aims To describe the nature of spatial and temporal variation in clutch size, and explore the ecological correlates of these patterns.

Methods We tested the prediction that seasonal declines in clutch size will be greater at higher latitudes. The environmental variables focused on were the influence of daylength, plant productivity, seasonality (i.e. Ashmole's hypothesis) and physiological mechanisms that relate clutch size to ambient temperature. We used data from 1980 to 2003 on spatial variation in clutch size across Britain for single‐brooded species, in which clutch size can be taken as a measure of annual reproductive investment. We included all seven species, from five families, with sufficient data in the British Trust for Ornithology's Nest Record Scheme.

Results There are strong seasonal declines in clutch size but little evidence for latitudinal gradients in clutch size or in latitudinal gradients in the rate of seasonal clutch size decline. Of the environmental variables investigated, daylength had the most marked effect on clutch size; this was positive in diurnal species and negative in the one nocturnal species.

Conclusions Although this study was confined to a relatively small latitudinal range of 8°, we found marked latitudinal gradients in a number of factors thought to drive spatial patterns in clutch size. Moreover, such variation is of sufficient magnitude to generate spatial patterns in other ecological variables in Britain. There is thus no simple explanation for the lack of a latitudinal gradient in clutch size. The results concerning daylength indicate that the time available for foraging is an important determinant of variation in clutch size.     

Determinants of local abundance in a major radiation of Australian passerines (Aves: Meliphagoidea)

Aim  To identify the factors that contribute to variation in abundance (population density), and to investigate whether habitat breadth and diet breadth predict macroecological patterns in a suborder of passerine birds (Meliphagoidea).
Location  Australia (including Tasmania).
Methods  Mean abundance data were collated from site surveys of bird abundance (the Australian Bird Count); range size and latitudinal position data from published distribution maps; and body mass and diet breadth information from published accounts. A diversity index of habitats used (habitat breadth) was calculated from the bird census data. We used bivariate correlation and multiple regression techniques, employing two phylogenetic comparative methods: phylogenetic generalized least squares and independent contrasts.
Results  Body mass and latitude were the only strong predictors of abundance, with larger-bodied and lower-latitude species existing at lower densities. Together, however, body mass and latitude explained only 11.1% of the variation in mean abundance. Range size and habitat breadth were positively correlated, as were diet breadth and body mass. However, neither range size, nor habitat breadth and diet breadth, explained patterns in abundance either directly or indirectly.
Main conclusions  Levels of abundance (population density) in meliphagoid birds are most closely linked to body mass and latitudinal position, but not range size. As with many other macroecological analyses, we find little evidence for aspects of niche breadth having an effect on patterns of abundance. We hypothesize that evolutionary age may also have a determining effect on why species tend to be rarer (less abundant) in the tropics.     

What determines conformity to Bergmann's rule?

Aim  Bergmann's rule, the tendency of body size within species in bird and mammal populations to be positively correlated with latitude, is among the best known biogeographical generalizations. The factors behind such clines, however, are not well understood. Here we use a large data base of 79 mammalian carnivore species to examine the factors affecting latitudinal size clines.
Location  Worldwide.
Methods  We measured the skulls and teeth of carnivores in natural history museums, and calculated the amount of variation in size explained by latitude, supplementing our measurements with published data. We examined the effects of a number of variables on the tendency to show latitudinal clines.
Results  We found that geographical range and latitudinal extent are strongly related to size clines. Minimum temperatures across the range, net primary productivity and habitat diversity also have some, albeit much less, influence.
Main conclusions  We suggest that species with large geographical ranges are likely to encounter significant heterogeneity in those factors that influence body size, and are thus likely to exhibit size clines. However, the key factors that determine body size may not always operate along a latitudinal (or other geographical) cline, but be spatially linked to patches in the species range. One such important factor is likely to be food availability, which we show is a strong predictor of size in the brown bear ( Ursus arctos ) but is not associated with a latitudinal cline. We argue that the spatial distribution of key resources within the species range constitutes a significant predictor of carnivore body size.     

Spatial variation in bird species abundances: Environmental constraints across southern Neotropical regions

Climate and habitat type are frequently related with the abundance of individual species and have been hypothesized to be primary drivers of the spatial variation in species abundances at the regional scale. Our aim is to evaluate the relative roles of those environmental factors in determining spatial variation in bird species abundance. We surveyed birds and habitat-cover variables and compiled climatic data along a 1700-km latitudinal gradient in the southern Neotropics. To identify the primary environmental variable explaining spatial changes in species abundances we performed simple regressions; a goodness of fit test identified the environmental factor that most frequently acted as the primary predictor. Mantel tests and partial regressions were performed to account for the spatial structure of abundance and environmental factors and collinearity between them. Of the 88 species included, 70% responded primarily to habitat cover and the remaining to climate. Forest cover and annual thermal amplitude were the main habitat-cover and climatic variables, respectively, explaining spatial variation in bird abundances. Our results indicated that the considered environmental factors accounted for latitudinal changes in species abundances; however, habitat cover and climate together explained a higher proportion of the variation than each factor independently of each other. There was a primacy of habitat-cover type over climate to predict spatial changes in bird species abundances across the neotropical biogeographic regions studied, but the underlying causes are likely related with the interaction of both factors.     

Latitudinal variation in clutch size–lay date regressions in Tachycineta swallows: effects of food supply or demography?

In a study of almost 16 000 nest records from seven swallow species across the entire Western Hemisphere, clutch sizes decline with relative laying date in each population, but the slope of this decline grows steeper with increasing distance from the equator. Late‐laying birds at all latitudes lay clutches of similar sizes, suggesting that latitudinal differences may be driven primarily by earlier‐laying birds. Focused comparisons of site‐years in North America with qualitatively different food availability indicate that food supply significantly affects mean clutch size but not the clutch size–lay date regression. Other studies on the seasonality of swallow food also indicate that steeper clutch size–lay date declines in the North are not caused by steeper earlier food peaks there. The distribution of lay dates grows increasingly right‐skewed with increasing latitude. This variation in lay‐date distributions could be due to the predominance of higher quality, early‐laying (and large‐clutched) individuals among populations at higher latitudes, resulting from latitudinal variation in mortality rates and the intensity of sexual selection. Our results underscore the importance of studying clutch size and lay date in tandem and suggest new research into the causes of their joint geographic variation.     

A meta‐analysis of global avian survival across species and latitude

Tropical birds are purported to be longer lived than their temperate counterparts, but it has not been shown whether avian survival rates covary with latitude worldwide. Here, we perform a global‐scale meta‐analysis of 949 estimates from 204 studies of avian survival and demonstrate that a latitudinal survival gradient exists in the northern hemisphere, is dampened or absent for southern hemisphere species, and that differences between passerines and nonpasserines largely drive these trends. We also show that while extrinsic factors related to climate were poor predictors of apparent survival compared to latitude alone, the relationship between apparent survival and latitude is strongly mediated by intrinsic traits – large‐bodied species and species with smaller clutch size had the highest apparent survival. Our findings reveal that differences among intrinsic traits and whether species were passerines or nonpasserines surpass latitude and its underlying climatic factors in explaining global patterns of apparent avian survival.     

Interspecific pairwise relationships among body size, clutch size and latitude: deconstructing a macroecological triangle in birds

Aim Ecogeographical ‘rules’, large‐scale patterns in ecological variables across geographical space, can provide important insights into the mechanisms of evolution and ecological assembly. However, interactions between rules could obscure both the observation of large‐scale patterns and their interpretation. Here, we examine a system of three variables interrelated by ecogeographical rules – the latitudinal increase in body size within closely related homeotherms (Bergmann’s rule), the negative allometry of clutch size (Calder’s rule) and the latitudinal increase in clutch size (Lack’s rule) – in a global dataset of birds. Location Global. Methods We used linear regressions and meta‐analysis techniques to quantify the three rules across clades and through the taxonomic hierarchy. Path analysis was used to quantify interactions between rules at multiple taxonomic levels, as a function of both phylogenetic inheritance of traits and indirect feedbacks between the three rules. Independent contrasts analyses were performed on four clades with available phylogenies, and the taxonomic partitioning of variation in each trait was quantified. Results Standardizing across all clades, Lack’s and Bergmann’s rules were supported at all taxonomic levels, with Calder’s rule being supported at the order level. Lack’s rule was consistently stronger and more often detected than the other two rules. Path analysis showed that the indirect effects often outweighed the direct effects of Calder’s rule at the genus level and Bergmann’s rule at the order level. Strong interactions between Calder’s and Bergmann’s rules led to a trade‐off between the rules depending on taxonomic resolution. Main conclusions We found strong interactions between Bergmann’s, Lack’s and Calder’s rules in birds, and these interactions varied in strength and direction over the taxonomic hierarchy and among avian clades. Ecogeographical rules may be masked by feedbacks from other, correlated variables, even when the underlying selective mechanism is operating. The apparently conflicting pairwise relationships among clutch size, body size and latitude illustrate the difficulty of interpreting individual pairwise correlations without recognition of interdependence with other variables.     

A comparative study of egg mass and clutch size in the Anseriformes

The factors explaining interspecific differences in clutch investment in precocial birds are poorly understood. We investigated how variations in clutch characteristics are related to environmental factors in a comparative study of 151 extant species of ducks, geese and swans (Anseriformes). Egg mass was negatively related to clutch size in a phylogenetic regression, a relationship that was much stronger when controlling for female mass. Nest placement was related to both egg size and clutch size, with cavity-nesting species laying more but smaller eggs. Egg size was positively correlated with incubation period and with female mass, and also with sexual size dimorphism (i.e. male mass relative to that of the female). Clutch size was not related to female mass. Species with long term pair bonds laid smaller clutches and larger eggs. The size of the breeding range was strongly positively correlated with clutch size and clutch mass, and its inclusion in multivariate models made other biogeographical variables (hemisphere, breeding latitude or insularity) non-significant. The small clutches in insular species appear to be a product of small range size rather than insularity per se. Our results suggest there is an evolutionary trade-off between clutch and egg size, and lend support to Lack’s resource-limitation hypothesis for the waterfowl.     

Covariation between clutch size, egg weight and egg shape: comparative evidence for chelonians

The variation in reproductive variables is documented both within and across species of chelonians. At both the generic and family levels, egg weight and clutch size show positive, significant correlations with adult carapace length. There is a negative correlation across both genera and families between clutch size and egg weight after removing the effects of body size, suggesting an evolutionary trade-off between these two life-history characteristics. However, the trade-off is not complete, since clutch size is positively correlated with clutch weight after removing the effects of body size. Terrestrial species lay fewer and larger eggs for their size than freshwater or marine species, but this association is statistically confounded by the fact that chelonian families form ecological groups. There is no significant association between habitat and clutch weight after removing the effects of body size, nor between latitude and either egg weight or clutch size, but temperate species have a heavier clutch weight after removing the effects of body size. Larger species lay eggs that are more spherical, but after controlling for body size, egg shape is not associated with clutch size. The patterns of covariation between adult weight, egg weight and clutch size contrast with those reported for birds and mammals, and some reasons for these differences are discussed.     

Latitudinal patterns in phenotypic plasticity: the case of seasonal flexibility in lizards’ fat body size

Several studies published over the last years suggest that the ability of many species to cope with global change will be closely related to the current amount of plasticity for fitness-related traits. Thus, disentangling general patterns in phenotypic flexibility, which could be then included in models aimed to predict changes in species distribution, represent a central goal in the current ecological agenda. The climatic variability hypothesis (CVH) could be considered a timely and promising hypothesis since it provides an explicit link between climatic and geographic variables and phenotypic plasticity. Specifically, the CVH states that as the range of climatic fluctuation experienced by terrestrial animals increases with latitude, individuals at higher latitudes should present greater levels of phenotypic flexibility. Within this framework, here we evaluate the existence of latitudinal patterns in fat body size flexibility—estimated as the difference between maximum and minimum fat body size values observed throughout a year—for 59 lizard species, comprising the first evaluation of the CVH for a trait, other than thermic or metabolic characters, in ectothermic species. Conventional and phylogenetic analyses indicated a positive relationship between fat body size flexibility and latitude, and also between flexibility and temperature variability indexes. Together with previous findings our results suggest that: (1) latitudinal pattern for fitness-related traits, other than thermal characters, are beginning to emerge; (2) latitude is usually a better predictor of phenotypic plasticity than putative climatic variables; (3) hemispheric differences in climatic variability appears to be correlated with hemispheric differences in phenotypic plasticity.     

CLUTCH SIZES AND BREEDING STRATEGIES AMONG BERMUDAN AND NORTH AMERICAN PASSERINES

Evidence for reduction of clutch size in insular passerines is presented. Average clutch sizes for the Catbird, Cardinal and White-eyed Vireo were reduced 14.0, 18.2 and 30.7%, respectively as compared with coastal North American populations from similar latitudes. Differences are highly significant. Clutch size in the Eastern Bluebird was not reduced. Trends in latitudinal and longitudinal variations of clutch size in North America are examined. With the exception of a slight latitudinal gradient in coastal populations, no consistent trends existed. There is no evidence for reduced variability in clutch size or increased egg size in the Bermudan populations, but both these parameters may show some response to combined effects of latitude and longitude in continental populations.
Reduction of clutch size in insular nidicolous birds is consistent with both the Lack-Ashmole resource limitation hypothesis and Cody's energy allocation hypothesis. In the absence of experimental evidence for life history optimization in response to K-selection, the modified Lack hypothesis seems sufficient.     

SOME ASPECTS OF THE BEHAVIOUR OF THE WATTLED PLOVER AFRIBYX SENEGALLUS (LINNAEUS)

The question of how aridity might influence avian clutch size, through the influences of rainfall seasonality and environmental stochasticity (unpredictability), has received little attention. A marked east-west gradient in aridity across South Africa provides a unique opportunity to test for such influences. Using an extensive collection of nest records for 106 terrestrial bird species from the South African Nest Record Card Scheme, we tested three predictions related to rainfall seasonality and stochasticity. Analyses were conducted at two levels, the first examining each species independently, and the second grouping species into five dietary guilds. The first prediction, that clutch size should generally increase with higher rainfall seasonality (i.e. higher seasonal fluctuation of food availability), was supported, particularly in the most arid environments where food abundance is more closely linked to rainfall. Controlling for rainfall seasonality, the second prediction, that clutch size should generally decrease as a bet-hedging strategy in arid, stochastic environments, was also supported. Although the timing of the rainy season differs among regions in South Africa (winter, early summer, later summer, year-round), birds primarily nest during spring. The relative timing of rainfall and breeding is expected to have different consequences for seasonal variation in clutch size among rainfall regions. The third prediction, of different patterns of seasonal variation in clutch size between rainfall regions, was also supported. In the winter and early-summer rainfall regions, early-nesting birds (breeding with or soon after the rains) generally had a larger clutch size than late-nesting birds. In the late-summer rainfall region, early-nesting birds (breeding well before the rains) had a smaller clutch size than late-nesting birds.     

Biomass compensation: Behind the diversity gradients of tidepool fishes

Species richness is unevenly distributed on the Earth, with biodiversity gradients of various spatial scales supposedly being affected by abiotic as well as biotic factors including community traits such as body size spectra and relative abundance patterns. To explore large-scale spatial variation in species diversity and their processes, tidepool fish communities were investigated through an intensive field work conducted on 55 shore sites in south-western Japan. Multiple ecological measures were taken into account to assess changes in local community structures with changes in the number of species. Biomass (total fish wet weight) per unit area showed no systematic change with latitude, while taxa richness and number of individuals tended to increase toward lower latitudes. In addition, median fish body weight scaled positively with latitude, which was more conspicuous in Blenniidae than in Gobiidae. The latitudinal gradient of diversity in tidepool fish assemblages appears to be characterized by partitioning of total biomass that tends to stay constant across latitudes, suggesting the phenomenon of “biomass compensation” whereby body size and abundance/diversity change in opposite directions with latitude. Our study highlights that biomass compensation can be part of processes involved in generating gradients of species richness even without an apparent energy/resource gradient.     

South temperate birds have higher apparent adult survival than tropical birds in Africa

Life history theory predicts an inverse relationship between annual adult survival and fecundity. Globally, clutch size shows a latitudinal gradient among birds, with south temperate species laying smaller clutches than north temperate species, but larger clutches than tropical species. Tropical birds often have higher adult survival than north temperate birds associated with their smaller clutches. However, the prediction that tropical birds should also have higher adult survival than south temperate birds because of smaller clutch sizes remains largely untested. We measured clutch size and apparent annual breeding adult survival for 17 south temperate African species to test two main predictions. First, we found strong support for a predicted inverse relationship between adult survival and clutch size among the south temperate species, consistent with life‐history theory. Second, we compared our clutch size and survival estimates with published estimates for congeneric tropical African species to test the prediction of larger clutch size and lower adult survival among south temperate than related tropical species. We found that south‐temperate species laid larger clutches, as predicted, but had higher, rather than lower, apparent adult survival than related tropical species. The latter result may be an artefact of different approaches to measuring survival, but the results suggest that adult survival is generally high in the south temperate region and raises questions about the importance of the cost of reproduction to adult survival.     

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.     

Does daylength explain the latitudinal variation in clutch size of Pied Flycatchers Ficedula hypoleuca?

Geographical trends in breeding parameters were studied in the Pied Flycatcher Ficedula hypoleuca in the western Palaearctic. Predictions arising from the hypothesis that daylength and/or energy requirements of the brood explain latitudinal clutch size variation were tested. The nestling period decreased with latitude, but nestling mass on day 13 after hatching did not show a trend with latitude. The length of the daily activity period (working day) at the time of peak brood demand showed a quadratic relationship with latitude and did not increase linearly with daylight hours. The present study supports the hypothesis that latitudinal clutch size variation is influenced by the duration of the working day and the energy requirements of the brood. The balance between the energy requirements of the brood and the parents, in relation to the duration of working day and ambient temperature, are proposed to explain the latitudinal variation in clutch size in the Pied Flycatcher.     

Biogeography of parasitism in freshwater fish: spatial patterns in hot spots of infection

Contrary to species occurrence, little is known about the determinants of spatial patterns of intraspecific variation in abundance, particularly for parasitic organisms. In this study, we provide a multi‐faceted overview of spatial patterns in parasite abundance and examine several potential underlying processes. We first tested for a latitudinal gradient in local abundance of the regionally most common parasite species and whether these species achieve higher abundances at the same localities (shared hot spots of infection). Secondly, we tested whether intraspecific similarity in local abundance between sites follows a spatial distance decay pattern or is better explained by variation in extrinsic biotic and abiotic factors between localities related to local parasite transmission success. We examined the infection landscape of a model fish host system (common and upland bullies, genus Gobiomorphus: Eleotridae) across its entire distributional range. We applied general linear models to test the effect of latitude on each species local abundance independently, including the abundance of each co‐infecting species as another predictor. We computed multiple regressions on distance matrices among localities based on abundance of each of the four most common trematode species, as well as for geographic distance, biotic and abiotic distinctness of the localities. Our results showed that the most widely distributed parasites of bullies also achieve the highest mean local abundances, following the abundance – occupancy relationship. Variation in local abundance of any focal parasite species was independent of latitude, the abundance of co‐occurring species and spatial distance or disparity in biotic attributes between localities. For only one parasite species, similarity of abundance between sites covaried with the extent of abiotic differences between sites. The lack of association between hot spots of infection for co‐occurring species reinforces the geographic mosaic scenario in which hosts and parasites coevolve by suggesting non‐deterministic, species‐specific variation in parasite abundance across space.     

Latitudinal gradient of species diversity: multi-scale variability in rocky intertidal sessile assemblages along the Northwestern Pacific coast

This study examined the latitudinal gradient of species diversity of rocky intertidal sessile assemblages on the slopes of rocks along the Northwestern Pacific coast of Japan, located between 31°N and 43°N, by explicitly incorporating an hierarchical spatial scale into the monitoring design. The specific questions were to examine: (1) whether there is a latitudinal gradient of regional diversity, (2) how spatial components of the regional diversity (local diversity and turnover diversity) vary with latitude depending on spatial scale, and (3) whether the latitudinal gradient differs between different measures of species diversity, i.e. species richness and Simpsons diversity index. We measured coverage and the presence or absence of all sessile organisms in a total of 150 census plots established at five shores in each of six regions. The results showed that there were clear latitudinal gradients in regional species richness and in species turnover among shores. However, these patterns were not reflected in smaller-scale local species richness. For Simpsons diversity index, there was no evidence of latitudinal clines either in regional diversity or in spatial components. These results suggest that relative abundance of common species does not vary along latitude, while the number of rare species increases with decreasing latitude.

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An erratum to this article can be found at     

Disentangling environmental drivers of metabolic flexibility in birds: the importance of temperature extremes versus temperature variability

Examining physiological traits across large spatial scales can shed light on the environmental factors driving physiological variation. For endotherms, flexibility in aerobic metabolism is especially important for coping with thermally challenging environments and recent research has shown that aerobic metabolic scope [the difference between maximum thermogenic capacity (M_sum) and basal metabolic rate (BMR)] increases with latitude in mammals. One explanation for this pattern is the climatic variability hypothesis, which predicts that flexibility in aerobic metabolism should increase as a function of local temperature variability. An alternative explanation is the cold adaptation hypothesis, which predicts that cold temperature extremes may also be an important driver of variation in metabolic scope. To determine the thermal drivers of aerobic metabolic flexibility in birds, we combined data on metabolic scope from 40 bird species sampled across a range of environments with several indices of local ambient temperature. Using phylogenetically‐informed analyses, we found that minimum winter temperature was the best predictor of variation in avian metabolic scope, outperforming all other thermal variables. Additionally, M_sum was a better predictor of latitudinal patterns of metabolic scope than BMR, with species inhabiting colder environments exhibiting increased M_sum over their counterparts in warmer environments. Taken together, these results suggest that cold temperature extremes drive latitudinal patterns of metabolic scope via selection for enhanced thermogenic performance in cold environments, supporting the cold adaptation hypothesis. Temperature extremes may therefore be an important selective pressure driving macrophysiological trends of aerobic performance in endotherms.     

Evolution of life history traits in Leporidae: a test of nest predation and seasonality hypotheses

Recently, it has been proposed that adult mortality schedules may be responsible for latitudinal patterns of life history variation in passerine birds, whereas nest predation only could explain within latitude patterns. Unfortunately, no independent test has been performed regarding the importance of nest predation with different taxa. In the present study, seasonality and nest predation hypotheses explaining variations in gestation time and litter size in 17 lagomorph species were tested. Among latitude patterns were analysed using the phylogenetic independent contrast method of Felsenstein and within latitude patterns were analysed by the pairwise comparative method. The results obtained indicate that latitudinal patterns observed in both variables are explained by different factors: seasonality for litter size and nest predation for gestation time. Litter size variations within latitudes are also explained by differences in nest predation, supporting previous hypotheses. In conclusion, the present study suggests that, when compared among latitudes, different life history traits (e.g. litter size and gestation time) may be shaped by different selective forces and that the effects of nest predation may be high both within and between latitudes.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 603–610.