Resistance of flight feathers to mechanical fatigue covaries with moult strategy in two warbler species

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.     

Annual variation in fat storage by a migrant warbler overwintering in the Indian tropics

1. We studied fat storage in a population of greenish leaf warblers ( Phylloscopus trochiloides ) in southern India over four winters (1993–97). This species breeds in temperate regions and overwinters in India from October to April.
2. Diurnal variation in fat scores was comparable to that seen among temperate wintering passerines. Seasonal variation was slight, except for premigratory fattening. There was significant annual variation: in drier winters, which were also winters of low food supplies, fat scores were higher.
3. Energy metabolized overnight is unlikely to vary across winters, and annual variation in fat scores is considered a response to uncertainty of food intake on a given day. Fat scores varied inversely with short-term rainfall, which drives the abundance of arthropod prey.
4. Annual variation in fat scores was not accompanied by changes in total body mass. This suggests that protein reserves were being compromised by increased fat deposition in dry years and offers an explanation for the observed delay in moult during drier winters. These results suggest that strategies for maximizing short-term probability of survival translate into future fitness costs, and may shape long-term life-history strategies through simple physiological processes.     

Habitat Use and Body Mass Regulation among Warblers in the Sahel Region during the Non-Breeding Season

Migratory birds face significant challenges across their annual cycle, including occupying an appropriate non-breeding home range with sufficient foraging resources. This can affect demographic processes such as over-winter survival, migration mortality and subsequent breeding success. In the Sahel region of Africa, where millions of migratory songbirds attempt to survive the winter, some species of insectivorous warblers occupy both wetland and dry-scrubland habitats, whereas other species are wetland or dry-scrubland specialists. In this study we examine evidence for strategic regulation of body reserves and competition-driven habitat selection, by comparing invertebrate prey activity-density, warbler body size and extent of fat and pectoral muscle deposits, in each habitat type during the non-breeding season. Invertebrate activity-density was substantially higher in wetland habitats than in dry-scrubland. Eurasian reed warblers Acrocephalus scirpaceus occupying wetland habitats maintained lower body reserves than conspecifics occupying dry-scrub habitats, consistent with buffering of reserves against starvation in food-poor habitat. A similar, but smaller, difference in body reserves between wet and dry habitat was found among subalpine warblers Sylvia cantillans but not in chiffchaffs Phylloscopus collybita inhabiting dry-scrub and scrub fringing wetlands. Body reserves were relatively low among habitat specialist species; resident African reed warbler A. baeticatus and migratory sedge warbler A. schoenobaenus exclusively occupying wetland habitats, and Western olivaceous warblers Iduna opaca exclusively occupying dry habitats. These results suggest that specialists in preferred habitats and generalists occupying prey-rich habitats can reduce body reserves, whereas generalists occupying prey-poor habitats carry an increased level of body reserves as a strategic buffer against starvation.     

Call divergence is correlated with geographic and genetic distance in greenish warblers (Phylloscopus trochiloides): a strong role for stochasticity in signal evolution?

Divergence in signalling systems might play a central role in speciation. To assess the importance of possible causes of signal divergence, we examine two types of vocalizations within a geographically variable species complex, the greenish warblers (Phylloscopus trochiloides Sundevall). Calls, which are used by both sexes throughout the year, and songs, which are sung primarily by breeding males, differ distinctly between two distinct Siberian forms. Through a ring of southern populations that connect the northern forms, signal divergence is correlated with both geographic distance and genetic divergence. Calls and songs differ in their particular patterns of geographic variation, probably because of the larger influence of sexual selection on songs than on calls. These patterns are supportive of neither acoustic adaptation nor morphology being major drivers of divergence in vocalizations. Rather, these results support the importance of stochastic evolution of communication systems in the evolution of new species.     

Fecundity selection predicts Bergmann's rule in syngnathid fishes

The study of latitudinal increases in organismal body size (Bergmann's rule) predates even Darwin's evolutionary theory. While research has long concentrated on identifying general evolutionary explanations for this phenomenon, recent work suggests that different factors operating on local evolutionary timescales may be the cause of this widespread trend. Bergmann's rule explains body size variation in a diversity of warm-blooded organisms and there is increasing evidence that Bergmann's rule is also widespread in ectotherms. Bergmann's rule acts differentially in species of the Syngnathidae, a family of teleost fishes noted for extreme adaptations for male parental care. While variation in body size of polygamous Syngnathus pipefish is consistent with Bergmann's rule, body size is uncorrelated with latitude in monogamous Hippocampus seahorses. A study of populations of Syngnathus leptorhynchus along a natural latitudinal and thermal gradient indicates that increases in body size with latitude maintain the potential reproductive rate of males despite significant decreases in ambient temperatures. Polygyny is necessary in order to maximize male reproductive success in S. leptorhynchus , suggesting a possible a link between fecundity selection and Bergmann's rule in this species.     

Bergmann's idiosyncratic rule: a role for fecundity selection?

The negative relationship between temperature and geographical variation in body size, or Bergmann's rule, is among the most thoroughly studied ecogeographical rules, yet the pattern and process underlying it remain controversial. Bergmann's original observations were of body size clines among endotherms, but in the last 50 years there has been increasing recognition that both Bergmann's rule and its reverse occur in many ectotherm taxa. A new study of syngnathid fish by Wilson (2009 ) in this issue of Molecular Ecology sheds light on intriguing alternative mechanisms that may explain variation in the direction of body size clines across taxa. Wilson shows that Bergmann's rule is found in pipefish of the genus Syngnathus , but not in seahorses of the genus Hippocampus . His results suggest that polygamy in pipefish allows fecundity selection to favour large size at low temperatures, compensating for increases in brooding time.     

The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards

We tested for the occurrence of Bergmann's rule, the pattern of increasing body size with latitude, and Rapoport's rule, the positive relationship between geographical range size and latitude, in 34 lineages of Liolaemus lizards that occupy arid regions of the Andean foothills. We tested the climatic-variability hypothesis (CVH) by examining the relationship between thermal tolerance breadth and distribution. Each of these analyses was performed varying the level of phylogenetic inclusiveness. Bergmann's rule and the CVH were supported, but Rapoport's rule was not. More variance in the data for Bergmann's rule and the CVH was explained using species belonging to the L. boulengeri series rather than all species, and inclusion of multiple outgroups tended to obscure these macroecological patterns. Evidence for Bergmann's rule and the predicted patterns from the CVH remained after application of phylogenetic comparative methods, indicating a greater role of ecological processes rather than phylogeny in shaping the current species distributions of these lizards.     

Why do European stoats Mustela erminea not follow Bergmann's rule?

Stoat Mustela erminea body size shows remarkably great variation over the species' European range. The pattern of this variation is opposite to that suggested by Bergmann's rule, i.e., stoats in central Europe are significantly larger than those in southern Sweden which, in turn are considerably larger than their conspecifics in northern Sweden. Neither winter temperature nor the length of snow cover shows any consistent correlation with stoat body size variation, and the larger body size of the southern populations could not be related to a widening of the stoat's feeding niche. A positive correlation was found between the frequency distribution, by size, of available prey and stoat body size in the different areas examined. This supports the hypothesis that body size variation in the stoat, especially females, is an adjustment to regional variations in the sizes of their available prey.     

Bergmann's rule in nonavian reptiles: turtles follow it,lizards and snakes reverse it

Abstract Bergmann's rule is currently defined as a within-species tendency for increasing body size with increasing latitude or decreasing environmental temperature. This well-known ecogeographic pattern has been considered a general trend for all animals, yet support for Bergmann's rule has only been demonstrated for mammals and birds. Here we evaluate Bergmann's rule in two groups of reptiles: chelonians (turtles) and squamates (lizards and snakes). We perform both nonphylogenetic and phylogenetic analyses and show that chelonians follow Bergmann's rule (19 of 23 species increase in size with latitude; 14 of 15 species decrease in size with temperature), whereas squamates follow the converse to Bergmann's rule (61 of 83 species decrease in size with latitude; 40 of 56 species increase in size with temperature). Size patterns of chelonians are significant using both nonphylogenetic and phylogenetic methods, whereas only the nonphylogenetic analyses are significant for squamates. These trends are consistent among major groups of chelonians and squamates for which data are available. This is the first study to document the converse to Bergmann's rule in any major animal group as well as the first to show Bergmann's rule in a major group of ectotherms. The traditional explanation for Bergmann's rule is that larger endothermic individuals conserve heat better in cooler areas. However, our finding that at least one ectothermic group also follows Bergmann's rule suggests that additional factors may be important. Several alternative processes, such as selection for rapid heat gain in cooler areas, may be responsible for the converse to Bergmann's rule in squamates.     

Amphibians do not follow Bergmann's rule

The tendency for organisms to be larger in cooler climates (Bergmann's rule) is widely observed in endotherms, and has been reputed to apply to some ectotherms including amphibians. However, recent reports provide conflicting support for the pattern, questioning whether Bergmann's clines are generally present in amphibians. In this study, we measured 96,996 adult Plethodon from 3974 populations to test for the presence of Bergmann's clines in these salamanders. Only three Plethodon species exhibited a significant negative correlation between body size and temperature consistent with Bergmann's rule, whereas 37 of 40 species did not display a pattern consistent with this prediction. Further, a phylogenetic comparative analysis found no relationship between body size and temperature among species. A meta-analysis combining our data with the available data for other amphibian species revealed no support for Bergmann's rule at the genus (Plethodon), order (Caudata), or class (Amphibia) levels. Our findings strongly suggest that negative thermal body size clines are not common in amphibians, and we conclude that Bergmann's rule is not generally applicable to these taxa. Thus, evolutionary explanations of Bergmann's clines in other tetrapods need not account for unique life-history attributes of amphibians.     

Testing mechanisms of Bergmann's rule: phenotypic decline but no genetic change in body size in three passerine bird populations

Bergmann's rule predicts a decrease in body size with increasing temperature and has much empirical support. Surprisingly, we know very little about whether "Bergmann size clines" are due to a genetic response or are a consequence of phenotypic plasticity. Here, we use data on body size (mass and tarsus length) from three long-term (1979-2008) study populations of great tits (Parus major) that experienced a temperature increase to examine mechanisms behind Bergmann's rule. We show that adult body mass decreased over the study period in all populations and that tarsus length increased in one population. Both body mass and tarsus length were heritable and under weak positive directional selection, predicting an increase, rather than a decrease, in body mass. There was no support for microevolutionary change, and thus the observed declines in body mass were likely a result of phenotypic plasticity. Interestingly, this plasticity was not in direct response to temperature changes but seemed to be due to changes in prey dynamics. Our results caution against interpreting recent phenotypic body size declines as adaptive evolutionary responses to temperature changes and highlight the importance of considering alternative environmental factors when testing size clines.     

A rare study from the wintering grounds provides insight into the costs of malaria infection for migratory birds

Malaria parasites can have strong effects on the population dynamics and evolution of migratory bird species. In many species, parasite transmission occurs on the wintering grounds, but studies to determine the consequences of infection have taken place during the breeding season, when malaria parasites circulate at chronic levels. We examined the predictors of malarial infections for great reed warblers during the northern winter in Africa, where active parasite transmission is thought to occur and naïve individuals experience acute infections. Counter to expectations, we found that winter infection intensities were lower than those encountered on the breeding grounds. One potential explanation is that reduced immune function during breeding allows parasites to persist at higher chronic intensities. We found no relationships between the incidence or intensity of infection on condition (as measured by scaled mass index, plasma metabolites, and feather corticosterone), spring migration departure dates, or home range sizes. We also tested a prediction of the Hamilton–Zuk hypothesis and found that male ornament (song) quality was unrelated to parasitic infection status. Overall, our results provide the first evidence that long‐distance migrants captured on their wintering grounds are in the chronic stage of infection, and suggest that winter studies may fare no better than breeding studies at determining the costs of acute malarial infection for great reed warblers.     

甘肃莲花山暗绿柳莺的孵卵行为

2001年5～7月,在甘肃省莲花山自然保护区对暗绿柳莺Phylloscopus trochiloides的孵卵行为进行了初步研究.结果 表明,孵化期内雌鸟日活动期长度平均为(848.5±14.8) min (n=17),每天离巢(16.0±3.0)次(n=15),每次离巢时间为(12.3±5.0) min (n=251),每次在巢时间为(43.6±21.9) min (n=236),在巢率为(78.8±2.4)%.雌鸟在巢时卵温平均为(31.3±3.5)℃ (n=10646),离巢时卵温平均为(26.6±4.8)℃ (n=2876);夜晚的平均卵温为(30.6±3.5)℃ (n=9239).孵卵温度在孵卵期有逐渐上升的趋势.     

Phylogeographic breaks without geographic barriers to gene flow

Abstract.— The spatial distribution of genetic markers can be useful both in estimating patterns of gene flow and in reconstructing biogeographic history, particularly when gene genealogies can be estimated. Genealogies based on nonrecombining genetic units such as mitochondrial and chloroplast DNA often consist of geographically separated clades that come into contact in narrow regions. Such phylogeographic breaks are usually assumed to be the result of long-term barriers to gene flow. Here I show that deep phylogeographic breaks can form within a continuously distributed species even when there are no barriers to gene flow. The likelihood of observing phylogeographic breaks increases as the average individual dispersal distance and population size decrease. Those molecular markers that are most likely to show evidence of real geographic barriers are also most likely to show phylogeographic breaks that formed without any barrier to gene flow. These results might provide an explanation as to why some species, such as the greenish warblers ( Phylloscopus trochiloides ), have phylogeographic breaks in mitochondrial or chloroplast DNA that do not coincide with sudden changes in other traits.     

Conflicting patterns of mitochondrial and nuclear DNA diversity in Phylloscopus warblers

Molecular variation is often used to infer the demographic history of species, but sometimes the complexity of species history can make such inference difficult. The willow warbler, Phylloscopus trochilus, shows substantially less geographical variation than the chiffchaff, Phylloscopus collybita, both in morphology and in mitochondrial DNA (mtDNA) divergence. We therefore predicted that the willow warbler should harbour less nuclear DNA diversity than the chiffchaff. We analysed sequence data obtained from multiple samples of willow warblers and chiffchaffs for the mtDNA cytochrome b gene and four nuclear genes. We confirmed that the mtDNA diversity among willow warblers is low (pi = 0.0021). Sequence data from three nuclear genes (CHD-Z, AFLP-WW1 and MC1R) not linked to the mitochondria demonstrated unexpectedly high nucleotide diversity (pi values of 0.0172, 0.0141 and 0.0038) in the willow warbler, on average higher than the nucleotide diversity for the chiffchaff (pi values of 0.0025, 0.0017 and 0.0139). In willow warblers, Tajima's D analyses showed that the mtDNA diversity, but not the nuclear DNA diversity, has been reduced relative to the neutral expectation of molecular evolution, suggesting the action of a selective sweep affecting the maternally inherited genes. The large nuclear diversity seen within willow warblers is not compatible with processes of neutral evolution occurring in a population with a constant population size, unless the long-term effective population size has been very large (N(e) > 10(6)). We suggest that the contrasting patterns of genetic diversity in the willow warbler may reflect a more complex evolutionary history, possibly including historical demographic fluctuations or historical male-biased introgression of nuclear genes from a differentiated population of Phylloscopus warblers.     

Migratory stopover conditions affect the developmental state of male gonads in garden warblers (Sylvia borin)

Long-distance migrants face the challenge of a short window for reproduction that requires optimal timing and full functional gonads. Male garden warblers (Sylvia borin) meet these demands by initiating testicular recrudescence during spring migration, enabling them to reproduce immediately after arrival at the breeding grounds. In a combined field and laboratory study, we investigated testicular size, plasma levels of luteinizing hormone (LH), androstenedione (AE), 5α-dihydrotestosterone (DHT), testosterone and nocturnal migratory restlessness (Zugunruhe) under different stopover conditions. We manipulated food availability, the duration of stopover and simulated migration by food deprivation. Garden warblers showed significantly retarded testicular development after nine days of stopover under limited food conditions compared to birds that had ad libitum access to food. However, there was no significant difference in Zugunruhe between the two groups. Thus, the degree of Zugunruhe was unaffected by the quality of the stopover site and migration continued independent of the developmental state of the testis. We suggest that male garden warblers face the necessity to either compensate for slowed testicular recrudescence during the subsequent leg of migration and delay arrival at the breeding grounds, or arrive with less developed testes. Either of these may reduce annual reproductive success.     

Geographical variation in body size of the Wood Mouse Apodemus sylvaticus L.

This paper analyses the geographical variation in body size of the Wood Mouse. A size increase is observed from north to south, contrary to Bergmann's rule. This increase is related to regions of sympatry and allopatry with the Yellow-necked Mouse, a closely related, potentially competing species. The possibility of character displacement playing an important role in determining the observed variation in body size is discussed.     

WHEN RENSCH MEETS BERGMANN: DOES SEXUAL SIZE DIMORPHISM CHANGE SYSTEMATICALLY WITH LATITUDE?

Bergmann's and Rensch's rules describe common large-scale patterns of body size variation, but their underlying causes remain elusive. Bergmann's rule states that organisms are larger at higher latitudes (or in colder climates). Rensch's rule states that male body size varies (or evolutionarily diverges) more than female body size among species, resulting in slopes greater than one when male size is regressed on female size. We use published studies of sex-specific latitudinal body size clines in vertebrates and invertebrates to investigate patterns equivalent to Rensch's rule among populations within species and to evaluate their possible relation to Bergmann's rule. Consistent with previous studies, we found a continuum of Bergmann (larger at higher latitudes: 58 species) and converse Bergmann body size clines (larger at lower latitudes: 40 species). Ignoring latitude, male size was more variable than female size in only 55 of 98 species, suggesting that intraspecific variation in sexual size dimorphism does not generally conform to Rensch's rule. In contrast, in a significant majority of species (66 of 98) male latitudinal body size clines were steeper than those of females. This pattern is consistent with a latitudinal version of Rensch's rule, and suggests that some factor that varies systematically with latitude is responsible for producing Rensch's rule among populations within species. Identifying the underlying mechanisms will require studies quantifying latitudinal variation in sex-specific natural and sexual selection on body size.     

Bergmann's rule in larval ant lions: testing the starvation resistance hypothesis

Abstract.  1. Body size of the ant lion Myrmeleon immaculatus follows Bergmann's rule – an increase in body size towards higher latitudes. The hypothesis that ant lion body size is larger in the north as an adaptation for starvation resistance was tested.
2. In a laboratory experiment testing starvation resistance, survivorship curves differed among 10 ant lion populations for both a starved and a fed treatment.
3. The average number of months survived by each population was correlated positively with latitude for both treatments. Across both treatments and all populations, large individuals survived longer than small individuals; however individuals from high latitudes had higher survivorship, even after factoring out variation due to initial body size.
4. These results suggest that starvation resistance may be an adaptation for coping with reduced prey availability in high latitudes. Starvation resistance may contribute to latitudinal gradients in body size of ant lions and other ectotherms.     

Bergmann's rule and the mammal fauna of northern North America

The observation that "on the whole…  larger species live farther north and the smaller ones farther south" was first published by Carl Bergmann in 1847. However, why animal body mass might show such spatial variation, and indeed whether it is a general feature of animal assemblages, is currently unclear. We discuss reasons for this uncertainty, and use our conclusions to direct an analysis of Bergmann's rule in the mammals in northern North America, in the communities of species occupying areas that were covered by ice at the last glacial maximum. First, we test for the existence of Bergmann's rule in this assemblage, and investigate whether small- and large-bodied species show different spatial patterns of body size variation. We then attempt to explain the spatial variation in terms of environmental variation, and evaluate the adequacy of our analyses to account for the spatial pattern using the residuals arising from our environmental models. Finally, we use the results of these models to test predictions of different hypotheses proposed to account for Bergmann's rule. Bergmann's rule is strongly supported. Both small- and large-bodied species exhibit the rule. Our environmental models account for most of the spatial variation in mean, minimum and maximum body mass in this assemblage. Our results falsify predictions of hypotheses relating to migration ability and random colonisation and diversification, but support predictions of hypotheses relating to both heat conservation and starvation resistance.