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The decision to provide parental care is often associated with trade-offs, because resources allocated to parental care typically cannot be invested in self-maintenance or mating. In most animals, females provide more parental care than males, but the reason for this pattern is still debated in evolutionary ecology. To better understand sex differences in parental care and its consequences, we need to study closely related species where the sexes differ in offspring care. We investigated parental care in relation to offspring growth in two closely related coucal species that fundamentally differ in sex roles and parental care, but live in the same food-rich habitat with a benign climate and have a similar breeding phenology. Incubation patterns differed and uniparental male black coucals fed their offspring two times more often than female and male white-browed coucals combined. Also, white-browed coucals had more 'off-times' than male black coucals, during which they perched and preened. However, these differences in parental care were not reflected in offspring growth, probably because white-browed coucals fed their nestlings a larger proportion of frogs than insects. A food-rich habitat with a benign climate may be a necessary, but-perhaps unsurprisingly-is not a sufficient factor for the evolution of uniparental care. In combination with previous results (Goymann et al . 2015 J. Evol. Biol . 28 , 1335-1353 (doi:10.1111/jeb.12657)), these data suggest that white-browed coucals may cooperate in parental care, because they lack opportunities to become polygamous rather than because both parents were needed to successfully raise all offspring. Our case study supports recent theory suggesting that permissive environmental conditions in combination with a particular life history may induce sexual selection in females. A positive feedback loop among sexual selection, body size and adult sex-ratio may then stabilize reversed sex roles in competition and parental care.

Most organisms in temperate or tropic regions employ the light-dark (LD) cycle as the primary Zeitgeber to synchronize circadian rhythms. At higher latitudes (>66°33'), continuous illumination during the summer presents a significant time-keeping dilemma for polar-adapted species. Lapland longspurs (Calcarius lapponicus), arctic-breeding migratory songbirds, are one of the few recorded species maintaining an intact diel rhythm in activity and plasma melatonin titers during polar summer. However, it is unknown whether rhythms are endogenous and entrain to low-amplitude polar Zeitgeber signals, such as daily variations in light intensity and the spectral composition of the sun (as measured by color temperature). Wild-caught male and female longspurs were brought into captivity, and locomotor activity was assessed using infrared detection. To examine if rhythms were endogenous, birds were exposed to constant bright light (LL; 1300 lux) or constant darkness (DD; 0.1 lux). All birds exhibited free-running activity rhythms in LL and DD, suggesting the presence of a functional circadian clock. Mean periods in LL (22.86 h) were significantly shorter than those in DD (23.5 h), in accordance with Aschoff's rule. No birds entrained to diel changes in light intensity, color temperature, or both. To examine endogenous molecular clock function, the Per2 gene was partially cloned in longspurs (llPer2) and transcripts were measured in hypothalamic tissue punches, eye, and liver using competitive polymerase chain reaction. Ocular llPer2 gene expression was periodic in LL and elevated at ZT24 (CT24) for LD or constant conditions (LL and DD), but llPer2 rhythmicity was not detected in hypothalamus or liver. Plasma melatonin was significantly lower in LL compared with LD or DD. In conclusion, rhythmic ocular Per2 expression and melatonin secretion may maintain the circadian activity rhythm across the polar day., (© 2014 The Author(s).)

Florida scrub-jays (Aphelocoma coerulescens) in the suburbs breed earlier than jays in native habitat. Amongst the possible factors that influence this advance (e.g., food availability, microclimate, predator regime, etc.), is exposure to artificial lights at night (LAN). LAN could stimulate the reproductive axis of the suburban jays. Alternatively, LAN could inhibit pineal melatonin (MEL), thus removing its inhibitory influence on the reproductive axis. Because Florida scrub-jays are a threatened species, we used western scrub-jays (Aphelocoma californica) to investigate the effects of LAN upon reproductive hormones and melatonin. Jays were held under conditions in which the dark-phase of the light:dark cycle was without illumination and then under low levels of LAN. Under both conditions, birds were exposed first to short-days (9.5L:14.5D) that were gradually increased to long-days (14.5L:9.5D). At various times, blood samples were collected during the light part of the cycle to measure reproductive hormones (luteinizing hormone, LH; testosterone, T; and estradiol, E2 ). Similarly, samples to assess melatonin were collected during the dark. In males, LAN caused a depression in LH levels and levels were ∼4× greater under long- than short-days. In females, there was no effect of LAN or photoperiod upon LH. LAN resulted in depressed T levels in females, although there was no effect on T in males. E2 levels in both sexes were lower under LAN than under an unlighted dark-phase. Paradoxically, MEL was higher in jays under LAN, and under long-days. MEL did not differ by sex. LAN disrupted the extraordinarily strong correlation between T and E2 that existed under unlighted nocturnal conditions. Overall, our findings fail to support the hypothesis that LAN stimulates the reproductive axis. Rather, the data demonstrate that LAN tends to inhibit reproductive hormone secretion, although not in a consistent fashion between the sexes., (© 2013 Wiley Periodicals, Inc.)

Polar environments are characterized by discrete periods of continuous light or darkness during the summer and winter months, respectively. Because the light/dark cycle serves as the primary Zeitgeber to synchronize rhythms of most organisms, its seasonal absence in polar regions poses challenges to the circadian organization of organisms that reside in these environments. Although some species become arrhythmic, others, such as migratory songbirds, are able to maintain an intact diurnal rhythm during polar summer. This suggests that birds may switch to alternative environmental cues, such as daily changes in light intensity and ambient temperature, which may have the potential to reset the biological clock. However, identifying the low-amplitude Zeitgeber that synchronizes rhythms in free-living polar-dwelling animals has been difficult to demonstrate. In this study, we measured behavioral and melatonin profiles of free-living Lapland longspurs (Calcarius lapponicus) near Barrow, Alaska (71°N) during the continuous daylight of summer in the Arctic. Diel cycles in activity and male singing were apparent throughout the polar day with a quiescence period of 4-5 hr starting around 24:00 Alaska Daylight Time. This inactivity corresponded with elevated melatonin profiles. In contrast, territorial aggression of males in response to a conspecific intruder was not dependent upon time-of-day. Diel changes in light intensity and ambient temperature were negatively associated with daily melatonin profiles after taking into account time-of-day effects. These results suggest that photic and thermal cues may act either as alternative Zeitgeber cues, or possibly masking agents. Distinguishing between these two possibilities will require further study., (Copyright © 2012 Wiley Periodicals, Inc.)

At higher latitudes, most organisms use the periodic changes in day length to time their annual life cycle. At the equator, changes in day length are minimal, and it is unknown which cues organisms use to synchronize their underlying circannual rhythms to environmental conditions. Here, we demonstrate that the African stonechat (Saxicola torquatus axillaris)-an equatorial songbird-can use subtle solar cues for the annual timing of postnuptial moult, a reliable marker of the circannual cycle. We compared four groups that were kept over more than 3 years: (i) a control group maintained under constant equatorial day length, (ii) a 12-month solar time group maintained under equatorial day length, but including a simulation of the annual periodic change in sunrise and sunset times (solar time), (iii) a 14-month solar time group similar to the previous group but with an extended solar time cycle and (iv) a group maintained under a European temperate photoperiod. Within all 3 years, 12-month solar time birds were significantly more synchronized than controls and 14-month solar time birds. Furthermore, the moult of 12-month solar time birds occurred during the same time of the year as that of free-living Kenyan conspecifics. Thus, our data indicate that stonechats may use the subtle periodic pattern of sunrise and sunset at the equator to synchronize their circannual clock.

Zebra Finches (Taeniopygia guttata) are highly social and monogamous birds that display relatively low levels of aggression and coordinate group life mainly by means of vocal communication. In the wild, small groups may congregate to larger flocks of up to 150-350 birds. Little is known, however, about possible effects of population density on development in captivity. Investigating density effects on physiology and behaviour might be helpful in identifying optimal group size, in order to optimise Zebra Finch wellbeing. A direct effect of population density on development and reproduction was found: birds in lower density conditions produced significantly more and larger (body mass, tarsus length) surviving offspring than birds in high density conditions. Furthermore, offspring in low density aviaries produced slightly longer song motifs and more different syllables than their tutors, whereas offspring in high density aviaries produced shorter motifs and a smaller or similar number of different syllables than their tutors. Aggression levels within the populations were low throughout the experiment, but the number of aggressive interactions was significantly higher in high density aviaries. Baseline corticosterone levels did not differ significantly between high- and low density aviaries for either adult or offspring birds. On day 15 post hatching, brood size and baseline corticosterone levels were positively correlated. On days 60 and 100 post hatching this correlation was no longer present. The results of this study prove that population density affects various aspects of Zebra Finch development, with birds living in low population density conditions having an advantage over those living under higher population density conditions., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Timing "in the real world" must cope with the temporal complexity of natural environments. Extreme examples for the resultant "multitasking" are migratory birds, which precisely time movements to remote areas. New field technologies highlight temporal accuracy, while captivity studies emphasize underlying programs and plasticity of schedules. After reviewing these findings, we focus on waders, which undertake spectacular long-distance migrations, have robust circannual clocks, and cope with diel, tidal, and polar environments. To explore features that may facilitate such multitasking, we speculated that melatonin amplitudes are low and damped during seasons when entrainment to subtle Zeitgebers occurs. We measured melatonin profiles under European daylength in two species with different ecologies and found low-amplitude melatonin cycles that changed over the year. Annual patterns neither fully supported our hypothesis, nor simply reflected daylight availability. While migratory birds are inspiring models for chronobiology, mechanistic understanding of their multitasking is still poor., (Copyright © 2012 Elsevier B.V. All rights reserved.)

A large number of passerine species migrate at night, although most of them are diurnal outside the migratory seasons. This diurnal-to-nocturnal transition is a major life-history event, yet little is known about its physiological control. Previous work showed that during the migratory periods captive birds showing nocturnal migratory restlessness (Zugunruhe) have reduced concentrations of circulating melatonin at night compared to non-migratory periods. This suggested that the hormone melatonin, a main component of the avian circadian system, is involved in the expression of Zugunruhe. Other studies demonstrated that the relationship between low melatonin levels and Zugunruhe is not a seasonal correlation. When Zugunruhe was interrupted by exposing birds to a fasting-and-refeeding protocol, melatonin levels increased. Here we studied whether melatonin and food availability influence the intensity of Zugunruhe in wild migrating garden warblers (Sylvia borin) at a stopover site. Birds were held in recording cages overnight, with or without food available, and either bled to determine melatonin concentrations or treated transdermally with melatonin. We found that melatonin levels at night were correlated with the intensity of diurnal locomotor activity and with condition, but were not correlated with Zugunruhe. Similarly, the melatonin treatment did not have effects on Zugunruhe, whereas food availability increased it. Our study shows that the nocturnal melatonin levels in migrating warblers depend on food availability and are correlated with condition. In addition, it suggests that melatonin does not control Zugunruhe and might rather be involved in energy conservation and/or clock synchronization during migration., (Copyright © 2010 Elsevier Inc. All rights reserved.)

Plasma testosterone increases during breeding in many male vertebrates and has long been implicated in the promotion of aggressive behaviors relating to territory and mate defense. Males of some species also defend territories outside of the breeding period. For example, the European nuthatch (Sitta europaea) defends an all-purpose territory throughout the year. To contribute to the growing literature regarding the hormonal correlates of non-breeding territoriality, we investigated the seasonal testosterone and corticosterone profile of male (and female) nuthatches and determined how observed hormone patterns relate to expression of territorial aggression. Given that non-breeding territoriality in the nuthatch relates to the reproductive context (i.e., defense of a future breeding site), we predicted that males would exhibit surges in plasma testosterone throughout the year. However, we found that males showed elevated testosterone levels only during breeding. Thus, testosterone of gonadal origin does not appear to be involved in the expression of non-breeding territoriality. Interestingly, territorial behaviors of male nuthatches were stronger in spring than in autumn, suggesting that in year-round territorial species, breeding-related testosterone elevations may upregulate male-male aggression above non-breeding levels. In females, plasma testosterone was largely undetectable. We also examined effects of simulated territorial intrusions (STIs) on testosterone and corticosterone levels of breeding males. We found that STIs did not elicit a testosterone response, but caused a dramatic increase in plasma corticosterone. These data support the hypothesis that corticosterone rather than testosterone may play a role in the support of behavior and/or physiology during acute territorial encounters in single-brooded species., (Copyright 2010 Elsevier Inc. All rights reserved.)

Arctic environments are challenging for circadian systems. Around the solstices, the most important zeitgeber, the change between night and day, is reduced to minor fluctuations in light intensities. However, many species including songbirds nonetheless show clear diel activity patterns. Here we examine the possible physiological basis underlying diel rhythmicity under continuous Arctic summer light. Rhythmic secretion of the hormone melatonin constitutes an important part of the songbird circadian system and its experimental suppression, e.g., by constant light, usually leads to behavioral arrhythmia. We therefore studied melatonin patterns in a free-living migratory songbird, the willow warbler (Phylloscopus trochilus), that maintains diel activity during the Arctic summer. We compared melatonin profiles during late spring and summer solstice in two Swedish populations from the south (58 degrees N) and near the Arctic circle (66 degrees N). We found the northern Swedish population maintained clear diel changes in melatonin secretion during the summer solstice, although peak concentrations were lower than in southern Sweden. Melatonin levels were highest before midnight and in good accordance with periods of reduced activity. The maintenance of diel melatonin rhythmicity under conditions of continuous light may be one of the physiological mechanisms that enables continued functioning of the circadian system.

To anticipate seasonal change, organisms schedule their annual activities by using calendrical cues like photoperiod. The use of cues must be fitted to local conditions because schedules differ between species and habitats. In complete absence of temporal information, many species show persistent circannual cycles that are synchronised, but not driven, by photoperiod. The contribution of circannual rhythms to timing under natural photoperiodic conditions is still unclear. In a suite of experiments, we examined timing in two closely related songbirds (Siberian and European stonechats) that inhabit similar latitudes but differ in seasonal behaviour. Under a more continental climate, Siberian stonechats breed later, moult faster and migrate further than European stonechats. We tested hypotheses for seasonal timing mechanisms by comparing the birds under constant and naturally changing daylengths. The taxa retained characteristic reproductive and moult schedules and hybrids behaved roughly intermediately. Based on their distinct circannual cycles, we expected European and Siberian stonechats to differ in photoperiodic responses at a given time of year. We found that the taxa responded, as predicted, in opposite ways to photoperiodic simulations as experienced on different migration routes. The findings indicate that circannual rhythms reflect geographically distinct periodic changes in seasonal disposition and cue-response mechanisms. Under natural daylengths, the phase relationship of the underlying circannual rhythm to the external year determines the action of photoperiod. Circannual rhythms are widespread among long-lived species. Accordingly, responses to environmental change, range expansion and novel migration patterns may depend on the particulars of a species' underlying circannual programming.

The winter immunoenhancement hypothesis associates long nights and increased exposure to melatonin with enhanced immune function in winter when resource availability is low and the chances of becoming ill are high. Thus, increased exposure to melatonin in the winter could be adaptive for species facing difficult winter conditions. This idea has found some support in studies of resident mammals. In birds, the link between day length and melatonin over the annual cycle is weaker, and contributions of melatonin to seasonal timing are unclear. Furthermore, many species, especially migrants, do not experience the most difficult conditions of their annual cycle in winter. In this study, we tested whether the winter immunoenhancement hypothesis holds in an avian species, the red knot Calidris canutus. We found that melatonin duration and amplitude varied significantly over the annual cycle with the highest values occurring in winter. However, peaks did not correspond to the winter solstice or with annual variation in immune function. Our findings do not support the winter immunoenhancement hypothesis in knots and question whether the idea that immune function should be bolstered in winter can be generalized to systems where winter is not the most difficult time of the year.

Ecological immunologists are interested in how immune function changes during different seasons and under different environmental conditions. However, an obstacle to answering such questions is discerning the effects of biological factors of interest and investigation artifacts such as handling stress. Here we examined handling stress and its effects on constitutive (noninduced) immune function via two protocols on captive red knots (Calidris canutus). We investigated how constitutive immunity responds to handling stress, how quickly these changes take place, and the practical implications for researchers interested in sampling baseline immune levels. We found that Staphylococcus aureus and Candida albicans killing increased with handling stress while total leukocyte and lymphocyte concentrations decreased. However, although corticosterone increased significantly and rapidly in response to handling stress, none of the 10 measures of constitutive immunity that we tested differed significantly from baseline within 20 or 30 min of capture. Thus, researchers interested in baseline immune function should sample animals as soon as possible after capture, but studies in species not easily sampled in less than 3 min (such as red knots) could still yield useful results.

Testosterone is assumed to be the key hormone related to resource-defence aggression. While this role has been confirmed mostly in the context of reproduction in male vertebrates, the effect of testosterone on the expression of resource-defence aggression in female vertebrates is not so well established. Furthermore, laboratory work suggests that progesterone inhibits aggressive behaviour in females. In this study, we investigated the hormonal changes underlying territorial aggression in free-living female African black coucals, Centropus grillii (Aves; Cuculidae). Females of this sex-role reversed polyandrous bird species should be particularly prone to be affected by testosterone because they aggressively defend territories similar to males of other species. We show, however, that territorial aggression in female black coucals is modulated by progesterone. After aggressive territorial challenges female black coucals expressed lower levels of progesterone than unchallenged territorial females and females without territories, suggesting that progesterone may suppress territorial aggression and is downregulated during aggressive encounters. Indeed, females treated with physiological concentrations of progesterone were less aggressive than females with placebo implants. This is one of the first demonstrations of a corresponding hormone-behaviour interaction under challenged and experimental conditions in free-living females. We anticipate that our observation in a sex-role reversed species may provide a more general mechanism, by which progesterone--in interaction with testosterone--may regulate resource-defence aggression in female vertebrates.

Daytime light intensity can affect the photoperiodic regulation of the reproductive cycle in birds. The actual way by which light intensity information is transduced is, however, unknown. We postulate that transduction of the light intensity information is mediated by changes in the pattern of melatonin secretion. This study, therefore, investigated the effects of high and low daytime light intensities on the daily melatonin rhythm of Afro-tropical stonechats (Saxicola torquata axillaris) in which seasonal changes in daytime light intensity act as a zeitgeber of the circannual rhythms controlling annual reproduction and molt. Stonechats were subjected to light conditions simulated as closely as possible to native conditions near the equator. Photoperiod was held constant at 12.25 h of light and 11.75 h of darkness per day. At intervals of 2.5 to 3.5 weeks, daytime light intensity was changed from bright (12,000 lux at one and 2,000 lux at the other perch) to dim (1,600 lux at one and 250 lux at the other perch) and back to the original bright light. Daily plasma melatonin profiles showed that they were linked with changes in daytime light intensity: Nighttime peak and total nocturnal levels were altered when transitions between light conditions were made, and these changes were significant when light intensity was changed from dim to bright. We suggest that daytime light intensity could affect seasonal timing via changes in melatonin profiles.

Endocrinological research on wild animals inhabiting remote areas has been hampered by the need to store plasma samples at subzero temperatures. In an attempt to remedy this logistical issue, we here investigate the use of ethanol as an alternative to freezing for the preservation of steroid and indoleamine hormones in avian plasma. Known quantities of the steroids 5alpha-dihydrotestosterone (DHT), testosterone, 17beta-estradiol, corticosterone, and the indoleamine melatonin were added to a stripped pool of chicken plasma. Samples were either immediately frozen at -40 degrees C, or treated with pure ethanol. Ethanol-treated samples were either immediately frozen, or-to simulate storage conditions at various field locations-left sitting at room temperature for one to two months, or incubated at 36 degrees C for one month before all treatment groups were frozen at -40 degrees C. All samples were then analyzed by radioimmunoassay. For DHT and estradiol there were no differences among treatment groups suggesting that ethanol-treatment is as effective as immediate freezing in preserving plasma steroid concentrations. For testosterone, corticosterone and melatonin ethanol-treated samples differed significantly from immediately frozen samples suggesting that caution is needed when comparing absolute concentrations of hormones between samples preserved in different ways. However, differences among ethanol-treated samples in general were small, demonstrating the feasibility of this preservation method in the field at remote locations.

Northern-temperate male birds show seasonal changes in testosterone concentrations with a peak during the breeding season. Many tropical birds express much lower concentrations of testosterone with slight elevations during breeding. Here we describe testosterone and corticosterone concentrations of male stonechats from equatorial Kenya during different substages of breeding and molt. This tropical species has a short breeding season of approximately 3 months. We compare their hormone concentrations to previously published data of males of a northern-temperate relative, the European stonechat, also a seasonal breeder but with a breeding season of approximately 5 months. Equatorial stonechats show a pronounced peak of testosterone during the nest-building and laying stage. During all other stages, testosterone concentrations are low, similar to other year-round territorial tropical bird species. Corticosterone concentrations peak also during the nest-building and laying stage suggesting that this period of maximum female fecundity is a demanding period for the male. Equatorial stonechats have significantly lower concentrations of testosterone than European stonechats during all stages, except during the nest-building and laying stage. During this stage of maximum female fertility, testosterone levels tend to be higher in equatorial than in European stonechats. Our results suggest that equatorial stonechats belong to a group of tropical bird species that are characterized by a short breeding season and a brief high peak of testosterone during the female's fertile period. Such brief, but substantial peaks of testosterone may be common in tropical birds, but they may easily be missed if the exact breeding stage of individual birds is not known.

Animals colonizing cities are exposed to many novel and potentially stressful situations. There is evidence that chronic stress can cause deleterious effects. Hence, wild animals would suffer from city life unless they adjusted their stress response to the conditions in a city. Here we show that European Blackbirds born in a city have a lower stress response than their forest conspecifics. We hand-raised urban and forest-living individuals of that species under identical conditions and tested their corticosterone stress response at an age of 5, 8, and 11 months. The results suggest that the difference is genetically determined, although early developmental effects cannot be excluded. Either way, the results support the idea that urbanization creates a shift in coping styles by changing the stress physiology of animals. The reduced stress response could be ubiquitous and, presumably, necessary for all animals that thrive in ecosystems exposed to frequent anthropogenic disturbances, such as those in urban areas.

Non-invasive methods to measure steroid hormone metabolites in bird droppings or mammalian feces have become very popular. However, the accuracy of these measurements may be affected by many factors. Here, we use the stonechat (Saxicola torquata) as a passerine bird model to test whether differences in ambient temperature affect food intake and dropping production and whether these changes lead to measurement artefacts in hormone metabolite concentrations. In addition, we tested for diurnal patterns in hormone metabolites. We held European stonechats in climate chambers and subjected them to two different long-term ambient temperature regimes, +5 degrees C and +22 degrees C. As expected, food intake and dropping production was higher at +5 degrees C than at +22 degrees C. Plasma concentrations of corticosterone and testosterone did not differ between different ambient temperature regimes. However, corticosterone and testosterone metabolite concentrations (in ng/g) were significantly lower at +5 degrees C than at +22 degrees C. When we measured the rate of hormone metabolite excretion (in picogram per hour) instead of the concentration, there was no difference between treatment groups. Thus, the measurement of hormone metabolite concentrations can be flawed because, depending on the treatment, similar amounts of hormone metabolites can be excreted into very different amounts of droppings. In conclusion, hormone metabolite concentration measurements are sensitive to changes in ambient temperature and probably any other factor that alters metabolic rates. Any study involving systematic changes in metabolism--i.e., during molt, migration, hibernation, egg production, or seasonal comparisons--needs to take these caveats into account.

Many songbirds in the temperate zones have comparatively short breeding seasons and are migratory. This often implies high breeding synchrony and competition for food, space, and mates. Tropical birds, in contrast, tend to be resident and react more flexibly to a more variable onset of environmental conditions conducive for breeding. Therefore, it can be expected that androgen metabolite concentrations in breeding males vary among birds from different latitudes. Studies on tropical birds, however, have revealed inconsistent results suggesting that additional factors not directly related to latitude, modify hormone levels. Furthermore, there is a potential for a phylogenetic bias when comparing birds of different taxonomic groups. In the present study, we compared androgen metabolite concentrations among closely related taxa of hand-raised male stonechats originating from tropical and temperate-zone populations in Kenya (0 degrees latitude), Kazakhstan (52 degrees N), and Ireland (52 degrees N), which differ partly in latitude of their provenance but also in the length of their breeding season (birds from Kazakhstan vs. Ireland). Hand-raised birds were housed in their natural photoperiod under otherwise identical environmental conditions. Androgen levels were determined from excrements to reduce disturbance. All three groups demonstrated a seasonal cycle of androgen metabolite concentrations concomitant with testis growth. Peak androgen metabolite concentrations were significantly lower in Kenyan stonechats compared to Kazakh stonechats. Irish birds had intermediate concentrations. Differences between Kazakh and Kenyan stonechats correlate with latitude, but data from the Irish population suggest the involvement of factors not directly related to latitude.