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.     

Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone

Summary Previous studies have suggested that tropical and temperate-zone lizards may differ fundamentally in life histories. We tested the applicability of this idea to Australian species by comparing temperate-zone species of agamid and scincid lizards with their congeners from the seasonal tropics. Data were derived from dissection of 1,941 specimens and from published information. Clutch size and egg size were positively correlated with mean maternal body size in most lizard species from both climatic zones. Mean body size of the lizards studies did not differ between the tropics and the temperate zone, nor did egg or hatchling size. However, tropical skinks showed considerably (approximately 20%) lower clutch size and relative clutch mass than did temperate-zone skinks. This difference was partly due to the higher incidence of species with low, invariant clutch size in the tropical lizard fauna (as seen in other continents as well), but primarily due to a trend for lineages (especially genera) with relatively high fecundity to be more common in the temperate zone than in the tropics. In contrast to studies on African lizards, our data suggested that modification of clutch size between areas has not occurred within genera: congeneric species from the tropics and temperate zone did not differ in clutch size. Production of more than one clutch per annum by individual females was common in both climatic zones. Tropical lizards may differ from temperate-zone species in showing higher reproductive frequencies, more rapid growth and earlier maturation. However, most of these effects may be due to phenotypic responses to environmental conditions (especially longer annual activity season), rather than to genetically based lifehistory adaptations.     

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.     

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.     

Sperm competition and the evolution of testes size in birds

Comparative analyses suggest that a variety of ecological and behavioural factors contribute to the tremendous variability in extrapair mating among birds. In an analysis of 1010 species of birds, we examined several ecological and behavioural factors in relation to testes size; an index of sperm competition and the extent of extrapair mating. In univariate and multivariate analyses, testes size was significantly larger in species that breed colonially than in species that breed solitarily, suggesting that higher breeding density is associated with greater sperm competition. After controlling for phylogenetic effects and other ecological variables, testes size was also larger in taxa that did not participate in feeding their offspring. In analyses of both the raw species data and phylogenetically independent contrasts, monogamous taxa had smaller testes than taxa with multiple social mates, and testes size tended to increase with clutch size, which suggests that sperm depletion may play a role in the evolution of testes size. Our results suggest that traditional ecological and behavioural variables, such as social mating system, breeding density and male parental care can account for a significant portion of the variation in sperm competition in birds.     

Sperm morphology and evidence for sperm competition among parrots

Sperm competition is an important component of post‐copulatory sexual selection that has shaped the evolution of sperm morphology. Previous studies have reported that sperm competition has a concurrently directional and stabilizing effect on sperm size. For example, bird species that show higher levels of extrapair paternity and larger testes (proxies for the intensity of sperm competition) have longer sperm and lower coefficients of variation in sperm length, both within and between males. For this reason, these sperm traits have been proposed as indexes to estimate the level of sperm competition in species for which other measures are not available. The relationship between sperm competition and sperm morphology has been explored mostly for bird species that breed in temperate zones, with the main focus on passerine birds. We measured sperm morphology in 62 parrot species that breed mainly in the tropics and related variation in sperm length to life‐history traits potentially indicative of the level of sperm competition. We showed that sperm length negatively correlated with the within‐male coefficient of variation in sperm length and positively with testes mass. We also showed that sperm is longer in sexually dichromatic and in gregarious species. Our results support the general validity of the hypothesis that sperm competition drives variation in sperm morphology. Our analyses suggest that post‐copulatory sexual selection is also important in tropical species, with more intense sperm competition among sexually dichromatic species and among species that breed at higher densities.     

Cellular Metabolic Rate Is Influenced by Life-History Traits in Tropical and Temperate Birds

In general, tropical birds have a “slow pace of life,” lower rates of whole-animal metabolism and higher survival rates, than temperate species. A fundamental challenge facing physiological ecologists is the understanding of how variation in life-history at the whole-organism level might be linked to cellular function. Because tropical birds have lower rates of whole-animal metabolism, we hypothesized that cells from tropical species would also have lower rates of cellular metabolism than cells from temperate species of similar body size and common phylogenetic history. We cultured primary dermal fibroblasts from 17 tropical and 17 temperate phylogenetically-paired species of birds in a common nutritive and thermal environment and then examined basal, uncoupled, and non-mitochondrial cellular O₂ consumption (OCR), proton leak, and anaerobic glycolysis (extracellular acidification rates [ECAR]), using an XF24 Seahorse Analyzer. We found that multiple measures of metabolism in cells from tropical birds were significantly lower than their temperate counterparts. Basal and uncoupled cellular metabolism were 29% and 35% lower in cells from tropical birds, respectively, a decrease closely aligned with differences in whole-animal metabolism between tropical and temperate birds. Proton leak was significantly lower in cells from tropical birds compared with cells from temperate birds. Our results offer compelling evidence that whole-animal metabolism is linked to cellular respiration as a function of an animal’s life-history evolution. These findings are consistent with the idea that natural selection has uniquely fashioned cells of long-lived tropical bird species to have lower rates of metabolism than cells from shorter-lived temperate species.     

Sperm competition selects for increased testes mass in Australian frogs

Game theory predicts that investment in spermatogenesis will increase with the risk and intensity of sperm competition. Widespread support for this prediction has come from comparative studies of internal fertilizing species reporting positive associations between testes mass and the probability that females mate with more than one male. Data for external fertilizers have generated conflicting results. We investigated how risk of sperm competition affects testes size in two families of Australian frogs: the Myobatrachidae and the Hylidae. We also examined effects of clutch size, egg size and oviposition location as alternative factors that might influence sperm production. Species were ranked according to probability of group spawning, and hence risk of sperm competition. Controlling for body size and phylogenetic relationships, we demonstrated that within the Myobatrachidae, the risk of sperm competition explained a significant amount of variation in testes mass. Oviposition location had a weak influence, with species ovipositing into foam having smaller testes. No significant effects of clutch size or egg size were detected. In hylids, the relationship between testes mass and risk of sperm competition was positive but not significant, again with no predictable effects related to egg size or number. These data provide an important test of sperm competition theory for externally fertilizing 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.     

Differences between temperate and tropical birds in seasonal acclimatization of thermoregulatory traits

Phenotypic flexibility can be an important determinant of fitness in variable environments. The climatic variability hypothesis (CVH) predicts that phenotypic flexibility in thermoregulatory traits will be greater in temperate species than tropical species as a means of coping with increased temperature seasonality at higher latitudes. However, support for the CVH has been mixed, and recent studies suggest that tropical birds are capable of substantial phenotypic flexibility. To test the generality of the CVH, we used flow‐through respirometry to quantify seasonal acclimatization in thermoregulatory traits in suites of temperate (n = 6) and tropical (n = 41) birds. We used W/S ratios (winter/summer trait values) to quantify the direction and magnitude of seasonal change (W/S ratio of 1 means no seasonal change). Temperate species exhibited coordinated changes in thermoregulatory traits in winter, including large increases in thermoneutral zone (TNZ) breadth and reductions in heat loss below the lower limit of the TNZ. Conversely, tropical species exhibited idiosyncratic seasonal thermoregulatory responses, and mean W/S ratios were close to 1 for all traits, indicative of little seasonal change and consistent with predictions of the CVH. Nevertheless, mean W/S ratios did not differ significantly between temperate and tropical species for either M_b or BMR, demonstrating that tropical birds can also exhibit substantial thermoregulatory flexibility. Our results highlight the need for complementary acclimation experiments to determine if latitudinal differences in seasonal acclimatization are due to inherent differences in capacity for flexibility.     

COMPARATIVE EVIDENCE FOR THE EVOLUTION OF SPERM SWIMMING SPEED BY SPERM COMPETITION AND FEMALE SPERM STORAGE DURATION IN PASSERINE BIRDS

Sperm swimming speed is an important determinant of male fertility and sperm competitiveness. Despite its fundamental biological importance, the underlying evolutionary processes affecting this male reproductive trait are poorly understood. Using a comparative approach in a phylogenetic framework, we tested the predictions that sperm swim faster with (1) increased risk of sperm competition, (2) shorter duration of female sperm storage, and (3) increased sperm length. We recorded sperm swimming speed in 42 North American and European free-living passerine bird species, representing 35 genera and 16 families. We found that sperm swimming speed was positively related to the frequency of extrapair paternity (a proxy for the risk of sperm competition) and negatively associated with clutch size (a proxy for the duration of female sperm storage). Sperm swimming speed was unrelated to sperm length, although sperm length also increased with the frequency of extrapair paternity. These results suggest that sperm swimming speed and sperm length are not closely associated traits and evolve independently in response to sperm competition in passerine birds. Our findings emphasize the significance of both sperm competition and female sperm storage duration as evolutionary forces driving sperm swimming speed.     

No association between sperm competition and sperm length variation across dung flies (Scathophagidae)

Sperm length is extremely variable across species, but a general explanation for this variation is lacking. However, when the risk of sperm competition is high, sperm length is predicted to be less variable within species, and there is some evidence for this in birds and social insects. Here, we examined intraspecific variation in sperm length, both within and between males, and its potential associations with sperm competition risk and variation in female reproductive tract morphology across dung flies. We used two measures of variation in sperm size, and testis size was employed as our index of sperm competition risk. We found no evidence of associations between sperm length variation and sperm competition or female reproductive tract variation. These results suggest that variation in sperm competition risk may not always be associated with variation in sperm morphology, and the cause(s) of sperm length variation in dung flies remains unclear.     

Assessing the reproductive consequences of mate retention and pair bond duration in Thorn-tailed Rayadito (Aphrastura spinicauda), a short-lived,socially monogamous neotropical bird

The adaptive value of mate retention has been studied in several socially monogamous birds but evidence of reproductive benefits for short-lived species is inconclusive. Most studies come from northern latitudes but more research on tropical birds is needed, as these species typically show higher survival rates and longer pair bonds than those from temperate regions. We gathered data on the reproductive biology of a subtropical, isolated population of Thorn-tailed Rayadito Aphrastura spinicaudaduring 2008–2017 to evaluate the reproductive consequences of mate retention. We examined data from 243 breeding attempts made by 159 breeding pairs. We found that ~30% of all breeding pairs bred together during at least two consecutive years, and some were mated for 6 years. The main cause of pair dissolution was mate loss, not divorce. Mixed-effects models provided moderate evidence for positive effects of mate retention and successive remating on reproductive success. Newly formed pairs laid eggs later and had slightly smaller clutches than remated pairs. Furthermore, clutch size seemed to increase with successive remating. Overall, our results suggest that newly formed pairs are less efficient in reproduction and that minor yearly reproductive benefits of mate retention might accumulate for birds that are able to breed with the same partner over many years. Because breeding habitat is limited in our study population, Thorn-tailed Rayaditos could benefit from remating if the number of individuals that can breed exceeds the number of available breeding positions. Profitable long-term pair bonds might be more frequent in tropical birds and therefore more studies are needed to assess the prevalence of remating-mediated effects on reproduction in relatively short-lived monogamous species breeding in tropical regions.     

Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Dissimilar bill shapes in new world tropical versus temperate forest foliage-gleaning birds

Summary The bill shape of foliage-gleaning birds in temperate and tropical new world forests is dissimilar. Tropical species have longer and narrower bills than their temperate zone counterparts. In addition, their bills are longer for a given body size. These differences cannot be readily explained as phylogenetic artifacts. I suggest that the distinct bill morphology of the two assemblages is determined by the type of insects that comprise the largest size classes of potential prey. These large insects are particularly important since they generally comprise the bulk of the nestling diet for insectivorous birds. In tropical forests Orthoptera are probably the most abundant large soft-bodied arthropods; they form an important resource for foliage-gleaning birds during the breeding (rainy) seasons. Most temperate zone foliage-gleaning birds rely almost entirely upon caterpillars when breeding. Long, narrow bills are thought to close more rapidly than shorter, broader bills. These long, fast bills may be required to efficiently harvest active Orthoptera. Migrant warblers may face morphological constraints from breeding successfully in lowland tropical forests. While the short-billed temperate zone birds can survive the tropical dry season by foraging on small arthropods, they may be inefficient at handling large Orthoptera to feed to nestlings.     

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 length variation as a predictor of extrapair paternity in passerine birds

Background

The rate of extrapair paternity is a commonly used index for the risk of sperm competition in birds, but paternity data exist for only a few percent of the approximately 10400 extant species. As paternity analyses require extensive field sampling and costly lab work, species coverage in this field will probably not improve much in the foreseeable future. Recent findings from passerine birds, which constitute the largest avian order (∼5 900 species), suggest that sperm phenotypes carry a signature of sperm competition. Here we examine how well standardized measures of sperm length variation can predict the rate of extrapair paternity in passerine birds.

Methodology/Principal Findings

We collected sperm samples from 55 passerine species in Canada and Europe for which extrapair paternity rates were already available from either the same (n = 24) or a different (n = 31) study population. We measured the total length of individual spermatozoa and found that both the coefficient of between-male variation (CV_bm) and within-male variation (CV_wm) in sperm length were strong predictors of the rate of extrapair paternity, explaining as much as 65% and 58%, respectively, of the variation in extrapair paternity among species. However, only the CV_bm predictor was independent of phylogeny, which implies that it can readily be converted into a currency of extrapair paternity without the need for phylogenetic correction.

Conclusion/Significance

We propose the CV_bm index as an alternative measure to extrapair paternity for passerine birds. Given the ease of sperm extraction from male birds in breeding condition, and a modest number of sampled males required for a robust estimate, this new index holds a great potential for mapping the risk of sperm competition across a wide range of passerine birds.     

Annual cycle of the Helmeted Honeyeater Lichenostomus melanops cassidix, a sedentary inhabitant of a predictable environment

The annual cycle of breeding, moult and weight variation in the Helmeted Honeyeater Lichenostomus melanops cassidix , a sedentary bird of temperate southeast Australia, is documented. Breeding and moult were sequential unimodal annual events, whose timing was highly consistent between years. However, overlap of breeding and moult was frequent, and some individuals even commenced primary moult before laying their final clutch. The timing of the post-juvenile moult was coincident with that of adults. Early-hatched young moulted within a few months of hatching, but late-hatched young deferred moult for a year. Helmeted Honeyeaters were heaviest in autumn and early winter, and lightest in spring and early summer, a cycle most consistent with the redirection of all available resources to reproduction. The long breeding season (seven-and-a-half months) of the Helmeted Honeyeater, extensive overlap of breeding and moult, and other life-history attributes including small clutch size, are more consistent with the described bio-rhythmic patterns for birds in the humid tropics than the temperate zone. However, the Helmeted Honeyeater has a fairly rapid primary moult rate, unusual amongst species that overlap moult and breeding. This combination of attributes reflects the stable, somewhat seasonal environment occupied and the resource monopoly established by this tightly territorial subspecies. We speculate that extension of the breeding season, by overlapping breeding and moult, is one of the few options available to vary life-history strategies amongst 'old-endemic' Australian birds of the temperate zone.     

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.     

Patterns of sperm damage in Chernobyl passerine birds suggest a trade-off between sperm length and integrity

Interspecific variation in sperm size is enigmatic, but generally assumed to reflect species-specific trade-offs in selection pressures. Among passerine birds, sperm length varies sevenfold, and sperm competition risk seems to drive the evolution of longer sperm. However, little is known about factors favouring short sperm or constraining the evolution of longer sperm. Here, we report a comparative analysis of sperm head abnormalities among 11 species of passerine bird in Chernobyl, presumably resulting from chronic irradiation following the 1986 accident. Frequencies of sperm abnormalities varied between 15.7 and 77.3% among species, more than fourfold higher than in uncontaminated areas. Nonetheless, species ranked similarly in sperm abnormalities in unpolluted areas as in Chernobyl, pointing to intrinsic factors underlying variation in sperm damage among species. Scanning electron microscopy of abnormal spermatozoa revealed patterns of acrosome damage consistent with premature acrosome reaction. Sperm length, but not sperm competition risk explained variation in sperm damage among species. This suggests that longer spermatozoa are more susceptible to premature acrosome reaction. Therefore, we hypothesize a trade-off between sperm length and sperm integrity affecting sperm evolution in passerine birds.