Functional linkages for the pace of life, life-history, and environment in birds

For vertebrates, body mass underlies much of the variation in metabolism, but among animals of the same body mass, metabolism varies six-fold. Understanding how natural selection can influence variation in metabolism remains a central focus of Physiological Ecologists. Life-history theory postulates that many physiological traits, such as metabolism, may be understood in terms of key maturational and reproductive characteristics over an organism's life-span. Although it is widely acknowledged that physiological processes serve as a foundation for life-history trade-offs, the physiological mechanisms that underlie the diversification of life-histories remain elusive. Data show that tropical birds have a reduced basal metabolism (BMR), field metabolic rate, and peak metabolic rate compared with temperate counterparts, results consistent with the idea that a low mortality, and therefore increased longevity, and low productivity is associated with low mass-specific metabolic rate. Mass-adjusted BMR of tropical and temperate birds was associated with survival rate, in accordance with the view that animals with a slow pace of life tend to have increased life spans. To understand the mechanisms responsible for a reduced rate of metabolism in tropical birds compared with temperate species, we summarized an unpublished study, based on data from the literature, on organ masses for both groups. Tropical birds had smaller hearts, kidneys, livers, and pectoral muscles than did temperate species of the same body size, but they had a relatively larger skeletal mass. Direct measurements of organ masses for tropical and temperate birds showed that the heart, kidneys, and lungs were significantly smaller in tropical birds, although sample sizes were small. Also from an ongoing study, we summarized results to date on connections between whole-organism metabolism in tropical and temperate birds and attributes of their dermal fibroblasts grown in cell culture. Cells derived from tropical birds had a slower rate of growth, consistent with the hypothesis that these cells have a slower metabolism. We found that dermal fibroblasts from tropical birds resisted chemical agents that induce oxidative and non-oxidative stress better than do cells from temperate species, consistent with the hypothesis that birds that live longer invest more in self-maintenance such as antioxidant properties of cells.     

Monthly survival of African Sylvia warblers in a seasonally arid tropical environment

Latitudinal gradients of life-history traits in animals are thought to be shaped by environmental variables. For example, it has been suggested that the increase in avian clutch size from the tropics towards the northern temperate regions is caused by a reduced survival of adult birds in the north due to increasing environmental seasonality. However, the tropical savannahs of East Africa show pronounced seasonality in resources caused by distinct rainy and dry seasons. This raises the question of whether survival and other life-history traits of birds living in these tropical savannahs are influenced by this seasonality, making them more similar to northern temperate species. We used 2-year monthly resighting data, a multistate modelling approach and the program MARK to test whether survival, transition probabilities between breeding states and other life-history traits of two resident Kenyan Sylvia species (Aves: Passeriformes: Sylviidae) are shaped by seasonality of rainfall in their environment. Contradicting our hypotheses, the two species showed only very slight influence of seasonality of rainfall on their survival. Survival in the dry months was hardly lower than in the rainy months. The species in the more seasonal environment ( S. boehmi , annual survival 71%) survived as well as the one in the more constant environment ( S. lugens , 56%). The observed survival rates correspond well to other life-history traits of the two species and are of similar magnitude to survival rates of other tropical passerines. This implies that either seasonality is not the driving force behind the life-history traits of the two species or the birds do not experience their environment as seasonal, as might be suggested by fluctuations in rainfall.     

Metabolic rate is negatively linked to adult survival but does not explain latitudinal differences in songbirds

Survival rates vary dramatically among species and predictably across latitudes, but causes of this variation are unclear. The rate‐of‐living hypothesis posits that physiological damage from metabolism causes species with faster metabolic rates to exhibit lower survival rates. However, whether increased survival commonly observed in tropical and south temperate latitudes is associated with slower metabolic rate remains unclear. We compared metabolic rates and annual survival rates that we measured across 46 species, and from literature data across 147 species of birds in northern, southern and tropical latitudes. High metabolic rates were associated with lower survival but survival varied substantially among latitudinal regions independent of metabolism. The inability of metabolic rate to explain latitudinal variation in survival suggests (1) species may evolve physiological mechanisms that mitigate physiological damage from cellular metabolism and (2) extrinsic rather than intrinsic sources of mortality are the primary causes of latitudinal differences in survival.     

Seasonality of peak metabolic rate in non‐migrant tropical birds

Birds that are year‐round residents of temperate and tropical regions have divergent life histories. Tropical birds have a slower ‘pace of life’, one characteristic of which includes lower peak metabolic rate and daily activity levels. Temperate resident birds are faced with seasonal variation in thermogenic demand. This challenge is met with seasonally increased peak metabolic rate during winter. These thermogenic demands are much lower in birds that are year‐round tropical residents. By measuring peak (summit) metabolic rate in tropical and temperate resident bird species during summer and winter, we asked whether tropical birds exhibit seasonality in peak metabolic rate, and if the direction of seasonality differs between tropical and temperate species. We measured summit metabolism in seven tropical and one temperate species during the winter and during the summer breeding season to test the hypothesis that summit metabolism of tropical residents would change seasonally. We consider whether metabolic seasonality is associated with breeding season for tropical species. We found that summit metabolism was significantly greater during the summer for most tropical residents, while the temperate resident matched several previous reports with higher summit metabolism in winter. We conclude that metabolic seasonality occurs in tropical residents and differs from temperate residents, suggesting that breeding during the summer may be driving relatively higher metabolism as compared to winter thermogenesis in temperate birds.     

Life-history of two African Sylvia warblers: low annual fecundity and long post-fledging care

The investigation of factors that cause differences in life-history traits between temperate and tropical birds is often hampered by a lack of knowledge about tropical species. Even within the well-known warblers of the genus Sylvia , which include resident species from temperate and tropical regions as well as migrants, there are few data from tropical species. We investigated the breeding biology of the tropical species Sylvia lugens and S. boehmi in a 2-year study in Kenya. Both species had a clutch size of 2.0 and laid c.  3.7 clutches per year. Breeding was characterized by long incubation periods ( S. lugens 14.5 days, S. boehmi 15.0 days), long nestling periods (16.0 and 12.9 days, respectively) and high predation rates (Mayfield nest success S. lugens 33.2%, S. boehmi 19.4%). Annual fecundity was 2.3 fledglings in S. lugens and 1.4 fledglings in S. boehmi . After fledging, the young birds were fed for 37.5 days ( S. lugens ) and 58.5 days ( S. boehmi ) (time to independence) and they stayed in their parents' territory for days or weeks, even after feeding had stopped. Fledgling survival until independence was 55.4% in S. lugens and 69.2% in S. boehmi . In general, S. lugens and S. boehmi have smaller but more numerous clutches, longer developmental periods, higher nest predation rates, lower annual fecundity and longer post-fledging care than their temperate congenerics.     

Sperm competition in tropical versus temperate zone birds

Sperm competition represents an important component of post-copulatory sexual selection. It has been argued that the level of sperm competition declines in birds towards the equator. However, to date, sperm competition estimates have been available mainly for avian species inhabiting the northern temperate zone. Here we apply a novel approach, using the coefficient of between-male variation (CV_bm) in sperm size as an index for sperm competition risk, in a comparative analysis of 31 Afrotropical and 99 northern temperate zone passerine species. We found no difference in sperm competition risk between the two groups, nor any relationship with migration distance. However, a multivariate model indicated that sperm competition risk was highest in species with a combination of low body mass and few eggs per clutch. The effect of clutch size was most pronounced in tropical species, which indicates that sperm competition risk in tropical and temperate species is differently associated with particular life-history traits. Although tropical species had lower sperm competition risk than temperate zone species for overlapping clutch sizes, the idea of a generally reduced risk of sperm competition in tropical birds was not supported by our analysis.     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.     

Corticosterone,testosterone and life-history strategies of birds

Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.     

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.     

Does low daily energy expenditure drive low metabolic capacity in the tropical robin, Turdus grayi?

Temperate and tropical birds possess divergent life history strategies. Physiological parameters including energy metabolism correlate with the life history such that tropical species with a slower ‘pace of life’ have lower resting and maximal metabolic rates than temperate congeners. To better understand the physiological mechanisms underlying these differences, we investigated the relationship of metabolic capacity, muscle oxidative capacity and activity patterns to variation in life history patterns in American robins (Turdus migratorius), while resident in central North America and Clay-colored robins (Turdus grayi) resident in Panama. We measured summit metabolism $ \left( {\dot{V}{\text{O}_{2\text{summit}}}} \right) $ in birds from both tropical and temperate habitats and found that the temperate robins have a 60 % higher metabolic capacity. We also measured the field metabolic rate (FMR) of free-living birds using heart rate (HR) telemetry and found that temperate robins’ daily energy expenditure was also 60 % higher. Thus, $\dot{V}{\text{O}_{2\text{summit}}} $ and FMR both reflect life history differences between the species. Further, both species operate at a nearly identical ~50 % of their thermogenic capacity throughout a given day. As a potential mechanism to explain differences in activity and metabolic capacity, we ask whether oxidative properties of flight muscle are altered in accordance with life history variation and found minimal differences in oxidative capacity of skeletal muscle. These data demonstrate a close relationship between thermogenic capacity and daily activity in free-living birds. Further, they suggest that the slow pace of life in tropical birds may be related to the maintenance of low activity rather than functional capacity of the muscle tissue.     

Ecological factors affect the level and scaling of avian BMR

The basal rate of metabolism (BMR) in 533 species of birds, when examined with ANCOVA, principally correlates with body mass, most of the residual variation correlating with food habits, climate, habitat, a volant or flightless condition, use or not of torpor, and a highland or lowland distribution. Avian BMR also correlates with migratory habits, if climate and a montane distribution is excluded from the analysis, and with an occurrence on small islands if a flightless condition and migration are excluded. Residual variation correlates with membership in avian orders and families principally because these groups are behaviorally and ecologically distinctive. However, the distinction between passerines and other birds remains a significant correlate of avian BMR, even after six ecological factors are included, with other birds having BMRs that averaged 74% of the passerine mean. This combination of factors accounts for 97.7% of the variation in avian BMR. Yet, migratory species that belong to Anseriformes, Charadriiformes, Pelecaniformes, and Procellariiformes and breed in temperate or polar environments have mass-independent basal rates equal to those found in passerines. In contrast, penguins belong to an order of polar, aquatic birds that have basal rates lower than passerines because their flightless condition depresses basal rate. Passerines dominate temperate, terrestrial environments and the four orders of aquatic birds dominate temperate and polar aquatic environments because their high BMRs facilitate reproduction and migration. The low BMRs of tropical passerines may reflect a sedentary lifestyle as much as a life in a tropical climate. Birds have BMRs that are 30-40% greater than mammals because of the commitment of birds to an expensive and expansive form of flight.     

Faster Speciation and Reduced Extinction in the Tropics Contribute to the Mammalian Latitudinal Diversity Gradient

The increase in species richness from the poles to the tropics, referred to as the latitudinal diversity gradient, is one of the most ubiquitous biodiversity patterns in the natural world. Although understanding how rates of speciation and extinction vary with latitude is central to explaining this pattern, such analyses have been impeded by the difficulty of estimating diversification rates associated with specific geographic locations. Here, we use a powerful phylogenetic approach and a nearly complete phylogeny of mammals to estimate speciation, extinction, and dispersal rates associated with the tropical and temperate biomes. Overall, speciation rates are higher, and extinction rates lower, in the tropics than in temperate regions. The diversity of the eight most species-rich mammalian orders (covering 92% of all mammals) peaks in the tropics, except that of the Lagomorpha (hares, rabbits, and pikas) reaching a maxima in northern-temperate regions. Latitudinal patterns in diversification rates are strikingly consistent with these diversity patterns, with peaks in species richness associated with low extinction rates (Primates and Lagomorpha), high speciation rates (Diprotodontia, Artiodactyla, and Soricomorpha), or both (Chiroptera and Rodentia). Rates of range expansion were typically higher from the tropics to the temperate regions than in the other direction, supporting the “out of the tropics” hypothesis whereby species originate in the tropics and disperse into higher latitudes. Overall, these results suggest that differences in diversification rates have played a major role in shaping the modern latitudinal diversity gradient in mammals, and illustrate the usefulness of recently developed phylogenetic approaches for understanding this famous yet mysterious pattern.     

Nest survival rates of Red‐crested Cardinals increase with nest age in south‐temperate forests of Argentina

ABSTRACT The main cause of nest mortality for most bird species is predation and nest survival rates often vary in relation to time‐specific variables. Few investigators have examined time‐specific patterns of nest survival in Neotropical birds, and most such studies have focused on tropical and subtropical species. To better understand age‐related patterns of nest survival, we studied nest survival of Red‐crested Cardinals (Paroaria coronata, Thraupidae) in a south‐temperate forest in Argentina. We modeled daily nest survival rates (DSR) using program MARK. We examined the relationship between nest age and nest survival rate, controlling for the effects of physical characteristics of nest sites and progression of the breeding season. We monitored 367 nests for a total of 4018 exposure days. We found that DSR increased with nest age and was higher in small isolated patches than in large continuous patches of forests. The increase of DSR with nest age could be a consequence of more vulnerable nests being predated early in the nesting cycle or a result of parents defending nests more vigorously as nestlings age because of their increasing reproductive value. Open areas of grassland that surrounded the small isolated patches of forests in our study may have been a barrier to predator movements, possibly explaining the lower predation rates. Nest survival rates in our study were lower than those reported for tropical or Nearctic temperate birds, but similar to those reported in other studies of Neotropical temperate birds. Reasons for the low nest survival rates of Neotropical temperate birds remain unclear, and additional studies of predator communities are needed to help elucidate this topic.     

Minimal effects of latitude on present-day speciation rates in New World birds

The tropics contain far greater numbers of species than temperate regions, suggesting that rates of species formation might differ systematically between tropical and non-tropical areas. We tested this hypothesis by reconstructing the history of speciation in New World (NW) land birds using BAMM, a Bayesian framework for modelling complex evolutionary dynamics on phylogenetic trees. We estimated marginal distributions of present-day speciation rates for each of 2571 species of birds. The present-day rate of speciation varies approximately 30-fold across NW birds, but there is no difference in the rate distributions for tropical and temperate taxa. Using macroevolutionary cohort analysis, we demonstrate that clades with high tropical membership do not produce species more rapidly than temperate clades. For nearly any value of present-day speciation rate, there are far more species in the tropics than the temperate zone. Any effects of latitude on speciation rate are marginal in comparison to the dramatic variation in rates among clades.     

Testosterone is correlated with courtship but not aggression in the tropical buff-banded rail,Gallirallus philippensis

Past studies into the roles of testosterone in birds have focused on species that occur in temperate regions. In such species, plasma testosterone levels are high during the establishment of territories in the spring and are associated with increased aggression. In contrast to most temperate species, tropical birds frequently defend territories year-round, during which time territoriality often occurs in a nonsexual context. The few studies that have been carried out on tropical birds show lower levels of circulating testosterone than occur in their temperate counterparts. In some year-round territorial tropical species, testosterone and aggression are dissociated, while in other species testosterone still plays a role in regulating aggression. This study examined the relationship between aggression and plasma testosterone levels in a year-round territorial, subtropical population of the buff-banded rail with characteristics typical of tropical species. Peak testosterone levels were substantially lower than those found in temperate species. Males displayed a seasonal peak in plasma testosterone level when their partners were most likely to be fertile. At other times, testosterone levels were mostly undetectable, despite year-round territoriality. We found that T levels increased with courtship behavior but showed no relationship with aggression, supporting the hypothesis that dissociation between testosterone and territoriality may be widespread among tropical avian taxa.     

Extra‐pair paternity in a population of Chilean Swallows breeding at 54 degrees south

Geographic patterns of variation in life‐history traits have puzzled researchers for decades. However, the widely accepted idea that mating systems exhibit a tropical–temperate latitudinal trend, with extra‐pair mating systems being the norm among temperate species and genetic monogamy the norm among tropical species, is supported by sparse data, particularly for birds breeding in the tropics and even more so for birds that breed in the southern hemisphere temperate zone. Our objective was to examine the genetic mating system of Chilean Swallows (Tachycineta meyeni) breeding at 54°S in Tierra del Fuego, Argentina. From 2006 to 2009, we examined the paternity of young in 52 broods. Contrary to predictions based on their congeners that breed at equivalent latitudes in the northern hemisphere, Chilean Swallows in our study had low rates of extra‐pair paternity (EPP), with 13.5% of nests (N = 52) having at least one extra‐pair young and 6.8% of all nestlings (N = 161) fathered by extra‐pair males. These rates are also lower than those reported for species of Tachycineta swallows that breed at tropical latitudes. We found no support for a tropical–south temperate latitudinal cline in EPP rates. The highly unpredictable weather of the island of Tierra del Fuego might be influencing parental investment at this site; small clutch sizes and low EPP rates are expected if biparental attention is crucial for chick survival and reproductive success for these aerial insectivores. We argue that the sparse sampling of mating systems in birds worldwide may have contributed to a misapprehension of a global pattern. More studies of tropical and south‐temperate systems are needed to build on theories based on a wider set of taxa.     

Nesting success of understory forest birds in central Panama

Greater nest predation in tropical than temperate birds has been hypothesized to be a primary selective force generating latitudinal differences in avian life history traits. Few extensive data sets, however, have been available from tropical forests to compare with data from temperate forests. To increase the amount of empirical information available for addressing issues related to the evolution of life history traits of tropical birds, we measured the nesting success of understory birds in lowland forest of central Panama. We found and monitored the fates of 696 nests of 71 species over two breeding seasons. Daily nest predation rates for the ten species for which we obtained the largest samples ranged from 1.6 to 8.3%, equivalent to a loss of 43 to 92% of nests. These values overlapped extensively the range of daily predation rates experienced by ecologically similar species in North America. Proportion of nests fledging young, estimated with the Mayfield method, was significantly lower in tropical (range: 8 to 57%) than temperate (27 to 60%) species. Nesting success in Panama varied among years, however, being greater in 1996 than 1997. In 1996, nesting success was similar to that of species breeding in forest fragments of midwestern North America. When compared with success of nests in large, contiguous forest tracts of North America, however, tropical avian nesting success was consistently lower by approximately 23%. We conclude that nesting success in central Panama may be poor in most breeding seasons, but also may be punctuated by occasional years of relatively exceptional success, a possibility heretofore unappreciated because of a general paucity of data from the tropics. Furthermore, our results indicate substantial variation in levels of nesting success among species, and almost no variation in clutch size. Such large interspecific variation, as well as potentially large annual variation, in nesting success does not support the hypothesis that uniformly low levels of nesting success select for small tropical clutch sizes.     

Divergent patterns of telomere shortening in tropical compared to temperate stonechats

Telomeres have emerged as important biomarkers of health and senescence as they predict chances of survival in various species. Tropical birds live in more benign environments with lower extrinsic mortality and higher juvenile and adult survival than temperate birds. Therefore, telomere biology may play a more important role in tropical compared to temperate birds. We measured mean telomere length of male stonechats (Saxicola spp.) at four age classes from tropical African and temperate European breeding regions. Tropical and temperate stonechats had similarly long telomeres as nestlings. However, while in tropical stonechats pre‐breeding first‐years had longer telomeres than nestlings, in temperate stonechats pre‐breeding first‐years had shorter telomeres than nestlings. During their first breeding season, telomere length was again similar between tropical and temperate stonechats. These patterns may indicate differential survival of high‐quality juveniles in tropical environments. Alternatively, more favorable environmental conditions, that is, extended parental care, may enable tropical juveniles to minimize telomere shortening. As suggested by previous studies, our results imply that variation in life history and life span may be reflected in different patterns of telomere shortening rather than telomere length. Our data provide first evidence that distinct selective pressures in tropical and temperate environments may be reflected in diverging patterns of telomere loss in birds.     

Energetics and life-history of bats in comparison to small mammals

Mammals can be aligned along a slow-fast life-history continuum and a low–high metabolic rate continuum based on their traits. Small non-volant mammals occupy the fast/high end in both continua with high reproductive rates and short life spans linked with high mass-specific metabolic rates. Bats occupy the high end of the metabolic continuum, but the slow end of the life-history continuum with low reproductive rates and long life spans. Typically, both continua are linked, and similar life-history traits of species are reflected in more similar metabolic rates. We therefore hypothesized that metabolic rates are similar in species with similar life-history traits. Resting metabolic rates (RMR) were measured for three ecologically and morphologically similar sympatric bat species (Myotis nattereri, M. bechsteinii, and Plecotus auritus; Vespertilionidae) and compared to data from other similar-sized, temperate insectivorous mammals with other life-history strategies. The bat species share similar life-histories and RMRs, both of which differ from the remaining mammals and therefore supporting our hypothesis. To verify that bats are similar in RMR, two energetically contrasting periods were compared. RMRs in post-lactating females did not differ between bat species. It was, however, positively correlated with parasite load in both Myotis species. However, RMRs differed during energy-demanding pregnancy where M. nattereri had the significantly lowest RMR, suggesting metabolic compensation as an energy-saving strategy. We conclude that the energy requirements of bat species with similar life-history traits resemble each other during periods of low energetic demands and are more similar to each other than to other small temperate mammals.     

Life history strategies of cladocerans: comparisons of tropical and temperate taxa

We review recent works on different life history variables of cladoceran taxa in tropical and temperate freshwater bodies, comparing the strategies that cladocerans have evolved to adapt to contrasting environmental conditions in the two geographical regions. These life-history parameters relate to age and size at maturity, survival, fecundity, life-expectancy at birth, lifespan, gross, and net reproductive rates, generation time, the rate of population increase, peak population density and day of peak abundance. We also discuss the role of photoperiod and temperature on some of these life history parameters. We found a general paucity of experimental work and field data in tropics on cladocerans. There is very limited information on the few Daphnia species found in the tropics. The misconception of low species diversity of cladocerans in the tropics arose due to several reasons including lack of extensive and intensive field collections. Higher water temperatures apparently promote permanent infestation of tropical waters with toxic cyanobacteria, which reduce the zooplankton diversity. In addition to higher temperatures in the tropics, the year-round high predation pressure of planktivorous fish probably causes the tropical species, particularly in pelagic habitats, to reach maturity earlier (< 3 days) than in temperate regions. Species of Daphnia in temperate regions are particularly adapted to living at food concentrations that are much lower and seasonably more variable than those for tropical genera such as Diaphanosoma. This is further corroborated by the more than an order of magnitude higher threshold food concentration (TFC) for tropical Cladocera than for their temperate counterparts. Fecundity patterns differ between tropical and temperate cladoceran taxa: cultured under optimal temperature regimes, tropical taxa have fewer eggs than temperate species of a comparable body size. Predation pressure may act differently depending on the size of the cladoceran neonates and thus on their population size structure. Global warming and climate changes seem to affect the behaviour (migration), distribution, and abundance of cladocerans. Apparently, in direct response to these changes, the possibility of encountering the tropical cladocerans in the northern, temperate hemisphere (bioinvasions) is on the rise.