Mixed biodiversity benefits of agri-environment schemes in five European countries

Agri-environment schemes are an increasingly important tool for the maintenance and restoration of farmland biodiversity in Europe but their ecological effects are poorly known. Scheme design is partly based on non-ecological considerations and poses important restrictions on evaluation studies. We describe a robust approach to evaluate agri-environment schemes and use it to evaluate the biodiversity effects of agri-environment schemes in five European countries. We compared species density of vascular plants, birds, bees, grasshoppers and crickets, and spiders on 202 paired fields, one with an agri-environment scheme, the other conventionally managed. In all countries, agri-environment schemes had marginal to moderately positive effects on biodiversity. However, uncommon species benefited in only two of five countries and species listed in Red Data Books rarely benefited from agri-environment schemes. Scheme objectives may need to differentiate between biodiversity of common species that can be enhanced with relatively simple modifications in farming practices and diversity or abundance of endangered species which require more elaborate conservation measures.     

Energy availability, abundance, energy-use and species richness in forest bird communities: a test of the species–energy theory

Aim To test the ‘more individuals hypothesis’ as a mechanism for the positive association between energy availability and species richness. This hypothesis predicts that total density and energy use in communities is linearly related to energy availability, and that species richness is a positive function of increased density. We also evaluate whether similar energy–density patterns apply to different migratory groups (residents, short‐distance migrants and tropical migrants) separately. Location European and North American forest bird communities. Methods We collected published breeding bird census data from Europe and North America (n = 187). From each census data we calculated bird density (pairs 10 ha^?1), energy use by the community (the sum of metabolic needs of individuals, Watts 10 ha^?1) and geographical location with an accuracy of 0.5°. For each bird census data coordinate we extracted the corresponding monthly values of actual evapotranspiration (AET). From these values we calculated corresponding AET values that we expected to explain the density energy use of forest birds: total annual, breeding season (June) and winter AET. We used general linear modelling to analyse these data controlling for the area of census plots, forest type and census method. Results Total density and energy use in European and North American forest bird communities were linear functions of annual productivity, and increased density and energy use then translated into more species. Also resident bird density and energy consumption were positive functions of annual productivity, but the relationship between productivity and density as well as between productivity and energy use was weaker for migrants. Main conclusions Our results are consistent with the more individuals hypothesis that density and energy use in breeding forest bird communities is coupled tightly with the productivity of the environment, and that increased density and energy consumption results in more species. However, not all community members (migratory groups) are limited by productivity on the breeding grounds.     

Annual and seasonal trends in the use of garden feeders by birds in winter

Provision of bird food in gardens is a common activity that may provide an alternative food source to birds in winter. Long-term survey data recording the weekly presence of all bird species using garden feeders in the winter were analysed to see if there was any evidence of trends in feeder use between 1970 and 2000 and whether these trends were correlated with breeding bird population trends. Of 41 species analysed, 21 showed significant increases in occurrence at garden feeders between 1970 and 2000. Many of these increases were evident only in the last 10 years. Several species showed significant positive correlations between trends in winter garden use and trends in relative population size in the previous breeding season. This was especially the case for species with population change (either increase or decrease) of the greatest magnitude. There was no evidence that seasonal shifts in garden feeder use were associated with population change in any species. Temperature was an important predictor of garden use but could not explain the year-to-year trends. However, as the number of feeding stations had increased over time the response of birds to the greater food availability in gardens may have been responsible for the widespread increases in occurrence of several species.     

Traditional management of forests: plant and bird community responses to alternative restoration of oak–hazel woodland in Sweden

Species richness of vascular plants and birds were examined in relation to five types of management (including abandonment) of oak-hazel woodland in south-central Sweden. The biodiversity of this type of woodland is affected by lengthy management in contrast to present emphasis on the fragmentation of more or less pristine forests. The woodlands derive from old deciduous forests but were a source of agricultural commodities from medieval time to end of the 19th century. Vascular plants and birds were censused during spring–summer for four years. Total number of plant species, species of field layer forbs, breeding birds and migrant birds were more numerous at simulated original management with mowing of small interior grasslands than at mechanical clearing or abandonment. Summer grazing led to intermediate numbers. Exotic plants were more common at long-time abandonment. Species richness of trees and shrubs, graminoids, all spring birds, resident birds and non-passerine birds did not exhibit any difference between treatments. Inter-site variation in species richness was usually smaller at abandonment than at true management. Species richness of plants and birds showed positive correlation in the particularly large sites examined. The general findings were in accordance with the intermediate disturbance hypothesis. Original management would secure most potential plant and bird species but extensive cattle grazing would also retain many species.     

Comparing population trend estimates of migratory birds from breeding censuses and capture data at a spring migration bottleneck

Europe has a well‐established network of breeding bird monitoring that is used to produce supranational indices of population trends for many species. However, a comparison of breeding bird censuses with other methods may be beneficial to confirm the validity of such indices. The aim of this study was to assess the value of standardized capture data of migratory birds at migration bottlenecks as an indicator of the effective breeding populations. One limitation to this method is that several populations are co‐occurring at these bottlenecks and their catchment areas need to be clearly identified to allow extrapolation of population indices. Here, we used standardized trends in capture numbers of 30 species on the island of Ponza, a migration bottleneck in the central Mediterranean, and compared them to population trends estimated in the putative catchment breeding areas between 2005 and 2016. The catchment areas were identified through the analysis of ring recoveries during the breeding season of birds passing through Ponza. Our results show an agreement between the population trends observed on Ponza and those in the breeding areas in 15 out of 30 species. The correlations were strongest in species with a more robust definition of the catchment areas, that is, species with more than 10 recoveries, and for which the recoveries were most likely of breeding birds. The main reason for disagreement between the two indices in the remaining species might be related to different intensity of sampling in different areas. This issue can be solved by further developing monitoring projects in underrepresented countries, as well as by intensifying monitoring through ringing, both in the breeding grounds and at migration bottlenecks. These results show that spring migration monitoring at bottlenecks has the potential to provide a valuable complement and an independent control of breeding bird surveys, allowing raising early warnings of population declines and contributing to their conservation.     

A COMPLETE SPECIES CENSUS AND EVIDENCE FOR REGIONAL DECLINES IN PIPING PLOVERS

Abstract: Complete population estimates for widely distributed species are rarely possible. However, for the third time in 10 years, an International Piping Plover (Charadrius melodus) Breeding and Winter Census was conducted throughout the species range in 2001. Nearly 1,400 participants from 32 U.S. states and Puerto Rico; 9 Canadian provinces; St. Pierre and Miquelon, France; Cuba; and the Bahamas visited 2,244 sites covering 11,836 km of shoreline habitat. During the winter census, 2,389 piping plovers were observed at 33.5% of potentially occupied sites (n = 352). Of these, 56.8% had ≤10 birds present. The breeding census recorded 5,945 adults at 777 of 1,892 sites surveyed. More than 80% of sites with piping plovers present had ≤10 birds. Results indicated an 8.4% increase from 1991 but only a 0.2% increase since 1996. Regional trends suggest that since 1991, number of breeding birds increased on the Atlantic Coast by 78% (2,920 birds; 12.4% increase since 1996) and by 80% in the Great Lakes (72 birds; 50% increase since 1996). However, plovers declined 15% (2,953 birds; 10% decline since 1996) in Prairie Canada/U.S. northern Great Plains. Subregional trends since 1991 reflect a 32.4% decline in Prairie Canada (972 birds; 42.4% decline since 1996), a 2.5% decline in the U.S. northern Great Plains (1,981 birds; 24% increase since 1996), 5.5% decline in eastern Canada (481 birds; 14% increase since 1996), although a 66.2% increase on the U.S. Atlantic Coast (2,430 birds; 12% since 1996). While numbers were down in much of the U.S. northern Great Plains since 1996, an increase (460%, 1,048 birds; 67.7% increase since 1991) was detected on the Missouri River. Results from 3 complete species census efforts provide essential data for conservation planning and assessment and illustrate the utility of global censuses for species of concern.     

Using simple species lists to monitor trends in animal populations: new methods and a comparison with independent data

There is an urgent need to develop simple and inexpensive methods for monitoring wildlife populations in resource-poor countries. List-based methods have been advocated as simple yet potentially useful biodiversity monitoring tools, and systems have recently been launched in a number of countries to collect species lists. We attempt to advance the use of systematic list-based monitoring by (1) suggesting improvements to the way in which list reporting rates are calculated; (2) assessing the extent to which degrading effort-corrected measures of abundance into simple species lists results in loss of information on population trends; (3) comparing long-term trends in list reporting rates with population trends from a wholly independent monitoring scheme. Daily species lists of birds were derived from regular trapping at a nature reserve in southern England. Most species showed a strong correlation across years between the proportion of lists on which they occurred, adjusted for list length (adjusted list reporting rate; ALRR), and an effort-corrected measure of abundance (captures per unit effort; CPUE). ALRR revealed almost as much about annual variation in abundance as CPUE for all but the most frequently captured species. Long-term (>20 years) trends in ALRRs at the nature reserve were positively correlated with UK national population trends recorded over the same period by an independent, labour-intensive monitoring scheme that counted birds at a large number of widely spread sites. Our results support previous claims that simple species lists could generate data useful for monitoring long-term population trends, particularly where such lists are collected systematically. However, further research on the efficiency of list reporting rates relative to more sophisticated methods is necessary, before list-based methods can be advocated for dedicated monitoring schemes in resource-poor regions.     

Explanations of commonness and rarity in British breeding birds: separating resource use and resource availability

Two of the most prominent explanations for a positive interspecific relationship between local abundance and regional distribution are the resource breadth and the resource availability hypotheses. Here we test these hypotheses by characterising habitat use of British breeding birds using extensive census and environmental data. A group of 85 bird species was considered for study along with 34 land use or environmental variables, which were used to generate four ordination axes by Canonical Correspondence Analysis. Measures of niche breadth and position were derived from these synthetic environmental axes using standard procedures. Across species, none of five measures of abundance and distribution chosen were correlated with niche breadth, whereas four out of five of these measures were correlated negatively with niche position. Repeating the analyses using a method designed to control for phylogenetic non-independence confirmed these general patterns. Birds that tended to use resources that were more atypical of the environment tended to be rarer and thinly distributed, while those using typical resources were common and widely distributed. Performing the analyses on subsets of the data based on species sample sizes did not alter the conclusions. On the assumption that our analyses properly capture patterns of niche use by birds, we find little support for the resource breadth hypothesis, but considerable support for the resource availability hypothesis.     

Bird-habitat associations predict population trends in central European forest and farmland birds

Recent studies show differences in population trends between groups of species occupying different habitats. In Czech birds, as well as in many other European countries, populations of forest species have increased, whereas populations of farmland species have declined. The aim of our study was to test whether population trends of particular species were related to finer bird-habitat associations within farmland and forest birds. We assessed bird-habitat associations using canonical correspondence analysis based on data from a 400 km long transect across the Czech Republic. We calculated population trends of 62 bird species using log-linear models based on data from a large-scale annual monitoring scheme, which covers the time series from 1982 to 2005. Within forest birds, species with a closer association with lowland broad-leaved forest have had more positive population trends, whereas species with a closer association with montane and coniferous forest revealed more negative population trends. We attribute these opposite trends to the gradual replacement of coniferous forests by deciduous ones, which took place in the Czech Republic during recent decades. Our analyses revealed a hump-shaped relationship within farmland birds, species most closely associated with farmland habitat revealing the most negative trends, whereas species with intermediate association to farmland habitat showed the most positive population trends. Such a pattern can be explained by the abandonment of previously cultivated areas followed by the spread of unmanaged meadows and scrubland. Changes in quantity or quality of preferred habitats may thus represent major drivers of observed bird population changes.     

Abundance patterns of European breeding birds

The frequency distribution of national population sizes of breeding birds was examined for each of 48 European countries, including autonomous regions, using data from the European Bird Database. The frequency distribution of logarithmically transformed abundance was left skewed in 40 countries: there was significant left skew in 10, and positive skew, in 2 countries. The inclusion of data classed as of low quality did not alter the conclusion that abundance patterns were predominantly left skewed. Abundance patterns of resident and migrant breeding birds differed; residents were left skewed, migrants were lognormal. Skewness was correlated negatively the number of species within a country. At low sample size, both left and right skews were evident, but left skews predominated at larger samples sizes. This pattern was driven partly by a difference between breeding populations on islands and on continental countries. Resampling procedures with fixed sample size showed that resident and migrant birds differed consistently in their patterns of skew, independent of sample size. If the birds of Europe were treated as a single assemblage, their abundance pattern was left skewed but this was driven by a very small number of vagrant breeders.     

Using fledged brood counts of hedgerow birds to assess the effect of summer agri-environment scheme options

A key aim of many European agri-environment schemes (AES) is to improve biodiversity on farmland. In recent years, several countries have been looking at long term trends in the spring adult population size of a target group of farmland birds as an indicator of this. The overall trend in these indicators is however not upwards. While this might suggest we need to look at the design and deployment of habitat management options within schemes, there is an increasing view that we also need to refine and improve our indicators, or the way we monitor them.Relating spring adult bird population size to AES options is problematic not least because of the time lag between the deployment of those designed to enhance bird breeding success in summer, and the spring surveys the following year. At the other end of the scale detailed studies of breeding success in farmland birds have practical/cost restraints. We argue that to understand the impact on farmland birds of particular summer options within AESs we need to be able to estimate the breeding success of local bird populations quickly and cheaply. This would enable us to relate particular AES options to the breeding performance of the birds actually using them.Complementing a previous study of woodland birds, we assess the likelihood of encountering fledged broods of hedgerow nesting bird species during transect surveys without finding nests, and then apply a simple mark-recapture analysis technique to provide an index of breeding success for those species. Following spring adult assessments, counts of fledged broods were undertaken four times a week during April, May, June and July, in four 2.5 km hedgerow transects, at four sites in southern England in 2010. Mean daily detection probabilities of fledged broods of 16 common hedgerow birds were calculated from these counts using the software Presence. For 15 out of these 16 species these detection probabilities were sufficiently high for a programme of fledged brood surveys, involving just two or three visits per week from mid-May to mid-July, to provide a useful estimate of breeding success.The survey technique and associated analyses make certain assumptions when providing estimates of breeding success and these are discussed. Little is known about initial dispersal in passerine fledglings and a study in hedgerows may be useful here. However our pilot study suggests that the method could have application as a relatively easily derived productivity index for hedgerow birds, and hence an additional method available to study the impact of certain AES options on indicator species, or for research studies.     

Geographic variation in the population trends of common breeding birds across central Europe

Recent declines of many European bird species have been linked with various environmental changes, especially land-use change and climate change. Since the intensity of these environmental changes varies among different countries, we can expect geographic variation in bird population trends. Here, we compared the population trends of bird species among neighbouring countries within central Europe (Czech Republic, Denmark, Germany, Switzerland) between 1990 and 2016 and examined trait-associations with population trends at both national and international scales. We found that Denmark had the highest proportion of declining species while Switzerland had the lowest. Species associated with farmland had negative trends, but the effect size tended to differ among countries. A preference for higher temperature was positively associated with population trends and its effect size was similar among countries. Species that were increasing across all four countries were associated with forest; while species that were decreasing across all countries were long-distance migrants or farmland birds. Our results suggest that land-use change tends to be a more regionally variable driver of common bird population trends than climate change in central Europe. For species declining across all countries, international action plans could provide a framework for more efficient conservation. However, farmland birds likely need both, coordinated international action (e.g. through a green agricultural policy) to tackle their widespread declines as well as regionally different approaches to address varying national effect trajectories.     

The hunting of migratory birds in Europe: a ringing recovery analysis

The hunting of 20 species of migratory birds in Europe and countries bordering the Mediterranean Sea was investigated using ringing recovery data. The intensity of the hunting of birds in each country was measured by the calculation of an index which controls for ringing effort and reporting rates. The hunting of birds was shown to be consistently high in western Mediterranean countries, particularly in southern France, northern and southwestern Iberia, northern Italy and in northwest Africa. Geographical patterns in the relative magnitude of hunting indices for different breeding populations were species-specific. In Europe most birds are taken during autumn and winter but in North Africa hunting is almost equally prevalent in spring. All species investigated, other than those that remain legitimate quarry species, showed a general reduction in index values after 1980. Analysis of long-term trends in index values since 1950 indicated a statistically significant overall decrease in the hunting of the majority of species. These changes are thought to be at least partly attributable to a real decline in the taking of birds but they may also reflect changed attitudes to reporting the hunting of species which are now protected. This analysis provides the first quantitative Europe-wide assessment of geographical and temporal trends in the hunting of migratory birds.     

Winter availability of cereal stubbles attracts declining farmland birds and positively influences breeding population trends

Many studies have demonstrated the selection of stubble fields by farmland birds in winter, but none have shown whether provisioning of this key habitat positively influences national population trends for widespread farmland birds. We use two complementary extensive bird surveys undertaken at the same localities in summer and winter and show that the area of stubble in winter attracts increased numbers of several bird species of conservation concern. Moreover, for several farmland specialists, the availability of stubble fields in winter positively influenced the 10 year breeding population trend (1994-2003) whereas hedgerow bird species were less affected. For skylarks and yellowhammers, initially negative trends showed recovery with 10-20 ha of stubble per 1 km square. Thus, agri-environment schemes that promote retention of over-winter stubbles will attract birds locally and are capable of reversing current population declines if stubbles are available in sufficient quantity.     

Long-term population declines in Danish trans-Saharan migrant birds

Capsule Long-distance migrant birds show less favourable trends than sedentary/short-distance species.

Aims To use breeding bird surveys to contrast population trends amongst common species according to their migration pattern.

Methods Changes in abundance of 62 Danish breeding sedentary, short-distance (Europe/North Africa) or long-distance (trans-Saharan) migrants were described by fitting log linear regression models to point-count census data gathered during 1976–2005.

Results Trans-Saharan migrants declined by 1.3% per annum during this period, while short-distance migrants and sedentary species increased by 1.4% and 1.0% per annum, respectively. There were no significant decadal declines amongst species using different summer breeding habitats, except for wetlands, and there was no consistent variation in trends associated with wintering regions or habitats or diet.

Conclusions More information is urgently needed on diet, feeding ecology, habitat requirements, winter distribution and intra-African movements of the commoner European summer visitors to identify causes of the declines and highlight when in the annual cycle detrimental effects occur. Studies linking individuals on their breeding, staging and wintering grounds are especially needed. Danish trends resemble those from elsewhere in Europe, confirming that restoration to favourable conservation status requires inter-continental action to meet European and global targets to reduce or halt biodiversity loss.     

Abundance,biomass and energy use of native and alien breeding birds in Britain

We quantify the contribution of alien species to the total breeding population numbers, biomass and energy use of an entire taxonomic assemblage at a large spatial scale, using data on British birds from 1997 and 2013. A total of 216 native and 16 alien bird species were recorded as breeding in Great Britain across the two census years. Only 2.8–3.7% of British breeding bird individuals were alien, but alien species co-opted 11.9–13.8% of the energy used by the assemblage, and contributed 19.1–21.1% of assemblage biomass. Neither the population sizes nor biomasses of native and alien species differed, on average, in either census, but alien species biomass is higher than native species biomass for a given population size. Species richness underestimates the potential effects of alien bird species in Britain, which have disproportionately high overall biomass and population energy use. The main driver of these effects is the ring-necked pheasant (Phasianus colchicus), which comprised 74–81% of alien biomass, yet the breeding population of this species is still only a small fraction of the estimated 35 million birds released in the UK in autumn. The biomass of this release exceeds that of the entire breeding avifauna, and suggests that the pheasant should have an important role in structuring the communities in which it is embedded.     

When and where to count? Implications of migratory connectivity and nonbreeding distribution to population censuses in a migratory bird population

Migratory connectivity is a metric of the co-occurrence of migratory animals originating from different breeding sites, and like their spatial dispersion, can vary substantially during the annual cycle. Together, both these properties affect the optimal times and sites of population censusing. We tracked taiga bean geese (Anser fabalis fabalis) during 2014–2021 to study their migratory connectivity and nonbreeding movements and determine optimal periods to assess the size of their main flyway population. We also compared available census data with tracking data, to examine how well two existing censuses covered the population. Daily Mantel's correlation between breeding and nonbreeding sites lay between 0 and 0.5 during most of the nonbreeding season, implying birds from different breeding areas were not strongly separated at other times in the annual cycle. However, the connectivity was higher among birds from the westernmost breeding areas compared to the birds breeding elsewhere. Daily Minimum Convex Polygons showed tracked birds were highly aggregated at census times, confirming their utility. The number of tracked birds absent at count sites during the censuses however exceeded numbers double-counted at several sites, indicating that censuses might have underestimated the true population size. Our results show that connectivity can vary in different times during the nonbreeding period, and should be studied throughout the annual cycle. Our results also confirm previous studies, which have found that estimates using marked individuals usually produce higher population size estimates than total counts. This should be considered when using total counts to assess population sizes in the future.     

Long-term breeding phenology shift in royal penguins

The Earth's climate is undergoing rapid warming, unprecedented in recent times, which is driving shifts in the distribution and phenology of many plants and animals. Quantifying changes in breeding phenology is important for understanding how populations respond to these changes. While data on shifts in phenology are common for Northern Hemisphere species (especially birds), there is a dearth of evidence from the Southern Hemisphere, and even fewer data available from the marine environment. Surface air temperatures at Macquarie Island have increased by 0.62°C during the 30-year study period (0.21°C decade(-1)) and royal penguins (Eudyptes schlegeli) commenced egg laying on average three days earlier in the 1990s than during the 1960s. This contrasts with other studies of Southern Ocean seabirds; five of nine species are now breeding on average 2.1 days later than during the 1950s. Despite the different direction of these trends, they can be explained by a single underlying mechanism: resource availability. There was a negative relationship between the Southern Annular Mode (SAM) and median laying date of royal penguins, such that low-productivity (low SAM) years delayed laying date. This accords with the observations of other seabird species from the Antarctic, where later laying dates were associated with lower sea ice and lower spring productivity. The unifying factor underpinning phenological trends in eastern Antarctica is therefore resource availability; as food becomes scarcer, birds breed later. These changes are not uniform across the region, however, with resource increases in the subantarctic and decreases in eastern Antarctica.     

Climate as a driver of phenological change in southern seabirds

Seabirds are one of the most threatened groups of birds globally and, overall, their conservation status is deteriorating rapidly. Southern hemisphere countries are over-represented in the number of species of conservation concern yet long-term phenological data on seabirds in the southern hemisphere is limited. A better understanding of the implications of changes in the marine and terrestrial environments to seabird species is required in order to improve their management and conservation status. Here we conducted a meta-analysis of the phenological drivers and trends among southern hemisphere seabirds. Overall there was a general trend towards later phenological events over time (34 % of all data series, N?=?47; 67 % of all significant trends), though this varied by taxa and location. The strongest trends towards later events were for seabirds breeding in Australia, the Laridae (gulls, noddies, terns) and migratory southern polar seabirds. In contrast, earlier phenologies were more often observed for the Spheniscidae (penguins) and for other seabirds breeding in the Antarctic and subantarctic. Phenological changes were most often associated with changes in oceanographic conditions, with sea-ice playing an important role for more southerly species. For some species in some locations, such as the Little Penguin Eudyptula minor in south-eastern Australia, warmer oceans projected under various climate change scenarios are expected to correspond to increased seabird productivity, manifested through earlier breeding, heavier chicks, an increased chance of double brooding, at least in the short-term.     

Investigating connectivity and seasonal differences in wind assistance in the migration of Common Sandpipers

Many migratory bird species have undergone recent population declines, but there is considerable variation in trends between species and between populations employing different migratory routes. Understanding species-specific migratory behaviours is therefore of critical importance for their conservation. The Common Sandpiper Actitis hypoleucos is an Afro-Palaearctic migratory bird species whose European populations are in decline. We fitted geolocators to individuals breeding in England or wintering in Senegal to determine their migration routes and breeding or non-breeding locations. We used these geolocator data in combination with previously published data from Scottish breeding birds to determine the distributions and migratory connectivity of breeding (English and Scottish) and wintering (Senegalese) populations of the Common Sandpiper, and used simulated random migrations to investigate wind assistance during autumn and spring migration. We revealed that the Common Sandpipers tagged in England spent the winter in West Africa, and that at least some birds wintering in Senegal bred in Scandinavia; this provides insights into the links between European breeding populations and their wintering grounds. Furthermore, birds tagged in England, Scotland and Senegal overlapped considerably in their migration routes and wintering locations, meaning that local breeding populations could be buffered against habitat change, but susceptible to large-scale environmental changes. These findings also suggest that contrasting population trends in England and Scotland are unlikely to be the result of population-specific migration routes and wintering regions. Finally, we found that birds used wind to facilitate their migration in autumn, but less so in spring, when the wind costs associated with their migrations were higher than expected at random. This was despite the wind costs of simulated migrations being significantly lower in spring than in autumn. Indeed, theory suggests that individuals are under greater time pressures in spring than in autumn because of the time constraints associated with reproduction.