Waterbird responses to regular passage of a birdwatching tour boat: Implications for wetland management

Participation in outdoor recreation can increase support for wildlife conservation, but may also disturb wildlife. We examined responses of wintering waterbirds to the regular passage of a small boat specifically dedicated to birdwatching tours in a coastal Ramsar site in northern Spain. Disturbances were measured during two separate periods: 2006–2008 and 2012–2015. Incidence and magnitude of disturbance events were compared by grouping species based on their interest to birders (target vs. non-target) and compared across sectors of the tour route. Flight-initiation distances (FID) were used to estimate species-specific buffer zones, which can be used to manage recreational disturbance to waterbirds. We further examined relationships between species-specific traits and FID, time flying, and distance flying following disturbance. A single boat tour disturbed on average 0.3% of non-target and 2.8% of target wintering bird populations within the wetland, with the effect being more pronounced on target species due to their smaller populations. Wing loading was positively associated with distance flying after disturbance. Based on measured FIDs, we calculated an overall buffer zone for all species of 100 m, and species-specific buffer zones ranging from 41 to 211 m. Disturbance incidence and the number of birds disturbed per tour were both greatest in narrow tidal channels (<200 m), where boats were forced to pass within 100 m of waterbirds. We urge caution in allowing boat passage through tidal channels in which boat operators cannot effectively maintain recommended buffer zones between their boat and waterbirds.     

Relationship between avian range limits and plant transition zones in Maine

Aim To determine if vegetation complexity associated with transition zones may be a contributing factor affecting bird species distributions in Maine, USA, and in increased numbers of bird species at about 45° north latitude in northeastern North America. Location Maine, USA; North America north of Mexico. Methods We delineated the ranges within Maine (86,156 km²) of 186 bird species and 240 woody plants using literature and expert review. Maps showing species richness and numbers of range limits, at 324 km² resolution, were developed for woody plants and groups of breeding birds: forest specialists, forest generalists, and those that used barren and urban habitats, early successional areas, and wetlands or open water. Two plant transition zones for Maine were identified previously, with the north–south transition zone mapped across eastern North America. Patterns in bird distribution maps were compared to woody plant maps and to transition zones. Results When the distributions of forest specialists were compared to the north–south vegetation transition zone in Maine, they were spatially coincident, but were not for other groups. Forest specialists had more species with range limits in the state (61%) than generalists (13%) or any other group. At a continental‐scale, the vegetation transition zone within eastern North America agreed fairly well with the areas of highest bird richness. Main conclusions A bird transition zone occurs in Maine and across eastern North America, akin to and overlapping the vegetation transition zone. Seasonality is likely the primary source of the inverse gradient in bird richness in the eastern USA, as reported by others. However, vegetation structure and habitat selection at very broad spatial scales appear to contribute to the reversed gradient. North of the vegetation transition zone, forest structure is simpler and coniferous forests more dominant, and this may contribute to reduced bird species richness. However, the northern (> 49°) typical gradient in bird species richness has been related to many hypotheses, and several are likely involved in the genesis of the gradient.     

Bird pollination syndrome is the plant's adaptation to ornithophily,but nectarivorous birds are not so selective

Many tropical plants are pollinated by birds and several bird phylogenetical lineages have specialised to a nectar diet. The long-assumed, intimate ecological and evolutionary relationship between ornithophilous plants and phenotypically specialised nectarivorous birds has nevertheless been questioned in recent decades, where such plant–pollinator interactions have been shown to be highly generalised. In our study, we analysed two extensive interaction datasets: bird–flower and insect–flower interactions, both collected on Mt Cameroon, west-central Africa. We tested if: 1) insects and birds interact with distinct groups of plants; 2) plants with a typical set of ornithophilous floral traits (i.e. bird pollination syndrome) interact mainly with birds; 3) birds favour plants with bird pollination syndrome and; 4) if and how the individual floral traits and plant level nectar production predict bird visitation. Bird-visited plants were typically also visited by insects, while approximately half of the plants were visited by insects only. We confirmed the validity of the bird pollination syndrome hypothesis, as plants with bird-pollination syndrome traits were visited by birds at a higher rate and mostly hosted a lower frequency of visiting insects. However, these ornithophilous plants were not more attractive than the other plants for nectar-feeding birds. Nectar production per plant individual was a better predictor of bird visitation than any other floral trait traditionally related to the bird pollination syndrome. Our study thus demonstrated the highly asymmetrical relationship between ornithophilous plants and nectarivorous birds.     

Elevation‐dependent effects of forest fragmentation on plant–bird interaction networks in the tropical Andes

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human‐induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species‐poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.     

Ostrich,Southern Ostrich or Common Ostrich? The ‘eternal vexed question’ of English bird names and name changes in southern Africa through eight editions of Roberts field guides, 1940–2016

A set of stable simple common bird names helps non-ornithologist birders, who contribute to conservation by visiting protected areas and participating in citizen science projects. Changes in English bird names have caused discomfort in the local birding community, especially those that followed international standardisation of common bird names between 2000 and 2005. To understand the extent and nature of English bird name changes, an analysis was done of all southern African bird names through the eight editions of Roberts Birds of South/Southern Africa field guides published from 1940 to 2016. Of 813 species listed in both the first and the latest of the field guides, 453 (55.7%) had their names changed, among which 108 (13.3%) had changes in both the group name and the species epithet. The greatest single wave of changes (31.4%) occurred in the first ‘Roberts bird guide’ (the seventh field guide) in 2007, following international standardisation. Mean word and syllable counts of bird names also increased significantly in that edition. Name changes were associated with new authorships, taxonomic changes and use of geographic species epithets. There was a trend towards name stability for southern African endemic species. Further name changes should be kept to a minimum, shortening and simplifying wherever possible.     

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.     

Exploring intraspecific variation in migratory destinations to investigate the drivers of migration

Various benefits (e.g. tracking of resources and of climate niche) and costs (e.g. distance travelled) are hypothesized to drive seasonal animal migrations. Until now, these potential factors have been investigated together at the species level, but migratory movements are made at the individual level, leading to intraspecific variability. Here, we use ringing/recovery data from 1308 individuals belonging to thirteen North American bird species to analyse patterns in intraspecific variability of migratory destinations in order to investigate which factors underpin bird migration and how individuals trade-off among multiple factors. Our results suggest that migratory destinations have been shaped by access to resources (most important during the breeding season) and climatic niche tracking (during winter, mostly). However, the benefits of resource surpluses and climate niche tracking appear to be traded off against the cost of migratory distance, which seems to strongly constrain where individuals migrate.     

杭州西溪国家湿地公园生态旅游环境容量

综合旅游生态容量、旅游空间容量、旅游设施容量、旅游管理容量、旅游心理容量5个因素,对杭州西溪国家湿地公园的旅游环境容量进行了测算.结果表明:其旅游环境容量为4145～6450人次.d-1.本研究首次将人鸟间的合理距离应用于湿地生态容量的确定,有效地解决了在充分保证鸟类活动与栖息安全的前提下适度开发湿地旅游的问题.基于游客满意度的旅游心理容量的估算较客观地反映了旅游区(点)在规划建设和经营管理方面满足旅游者需求的程度.这种基于人鸟距离和游客满意度的综合测算方法对于湿地公园的规划和管理具有较实际的指导意义.     

A spatialized digital database for all bird species in China

The lack of refined spatial detail on bird distribution in China has hindered further research due to the large geographic unit(provincial level) in existing national bird distribution data. Based on multi-source bird distribution data, we built a more spatially detailed distribution database for every bird species(1,371 species) in China, covering 2,908 counties. The sources on bird distribution are grouped into six categories: Handbooks, Literature; Fauna, Avifauna; Paper; Citizen Science data by ornithologists or birders; GPS tracing; and Website data. The database contains the following records: taxonomy, distribution data, suspicious species information, and data sources. Our database recorded 835 species(61%) appearing outside the distribution range previously known. The use of provincial boundaries as the smallest geographical unit has created misleading distribution results due to geographic aggregation for most species. The new database was built based on increased observational frequency and individuals observed in previously undetected areas particularly in Western China and towards higher altitudes and latitudes. They coincided with the discovery of the range expansion of some waterfowls into Xinjiang. The dataset provides a new base for Chinese and international ornithology studies, especially for those requiring more detailed distribution information for many taxa and large-scale regional research.     

Guidelines for cross‐cultural Participatory Action Research partnerships: A case study of a customary seabird harvest in New Zealand

Abstract

Adaptive co‐management and Participatory Action Research (PAR) promotes social ecological resilience by simultaneously protecting wildlife and its habitat and promoting capacity and motivation for sustainable harvest management by communities. We report here on a case study of learning through a partnership (1994–2009) between science and Traditional Ecological Knowledge (TEK) to determine the sustainability of titi (sooty shearwater, Puffinus griseus) harvests by Rakiura Maori in southern New Zealand. Testimony of Maori elders and titi harvesters (birders), members of the Rakiura Titi Islands Administering Body, researchers and participants in workshops and meetings were recorded throughout the 14‐year research project to identify critical determinants of success of the partnership. A large majority of participants supported the research, mainly because it expanded their knowledge by investigating the reasons for declining bird numbers and the means of ensuring the continuation of their muttonbirding heritage. Initial concerns about the research included fear that prohibition or quota would be imposed through political pressure from external groups; the intrusion of strangers on the islands; the misconception that the research was being promulgated by government regulatory agencies; and scepticism about research findings. Research also precipitated conflict and division within the Rakiura community, and some birders feared that science might displace matauranga Maori (TEK) of the Rakiura people for guiding harvest management. Core conditions for community engagement included trust between parties, effective communication of the science, equitable decision‐making responsibility, and building scientific capability and monetary support to enable meaningful participation. The most fundamental requirement is mutual respect for each party's knowledge. Attention to this inclusive, equitable, slow and prolonged process makes it more likely that the community will uptake results to improve sustainability of harvesting. The research has heightened awareness within the harvesting community of conservation issues facing the titi and of potential options to mitigate them. Eradication or control of weka (Gallirallus australis), and reducing titi harvest levels from around a quarter of the manu (family birding territories), are the main practical ways of increasing sustainability, but the magnitude and direction of climate change impacts on the shearwater population remains uncertain.     

Comparison of audio recording system performance for detecting and monitoring songbirds

ABSTRACT Acoustic recording systems are being used more frequently to estimate habitat occupancy or relative abundance, and to monitor population trends over time. A potential concern with digital recording systems is that changes in technology could affect detectability of birds and cause bias in trend estimates based on counts of birds detected. We evaluated several currently available commercial recording systems ranging from low‐cost multipurpose digital recorders to custom‐designed wildlife recorders (US$250–$7000 price range) to examine possible differences among systems in species detection. We made recordings during Breeding Bird Surveys (BBS) counts using several units concurrently, and asked several expert birders to listen to the recordings in a factorial design. We found that birders detected, on average, 10% fewer species on some units compared to others, though there was high variance. Analysis of a subset of recordings, using spectrograms and repeated listening, suggested that ～90% of species on each BBS stop could be clearly detected on all units. The remaining species could be identified on at least one unit, but were hard or impossible to detect on others. We found that the recording unit with the lowest empirical signal‐to‐noise‐ratio (SNR) had the lowest number of birds detected on the BBS recordings, and that frequency‐specific SNR differed among units. Missed detections were likely related to variation in internal noise and frequency‐dependent sensitivity of the units, and were an issue for all systems regardless of price. We caution that researchers using recorders need to consider variation among recording systems in their study design, particularly for long‐term monitoring programs.     

Local parasite lineage sharing in temperate grassland birds provides clues about potential origins of Galapagos avian Plasmodium

Oceanic archipelagos are vulnerable to natural introduction of parasites via migratory birds. Our aim was to characterize the geographic origins of two Plasmodium parasite lineages detected in the Galapagos Islands and in North American breeding bobolinks (Dolichonyx oryzivorus) that regularly stop in Galapagos during migration to their South American overwintering sites. We used samples from a grassland breeding bird assemblage in Nebraska, United States, and parasite DNA sequences from the Galapagos Islands, Ecuador, to compare to global data in a DNA sequence registry. Homologous DNA sequences from parasites detected in bobolinks and more sedentary birds (e.g., brown‐headed cowbirds Molothrus ater, and other co‐occurring bird species resident on the North American breeding grounds) were compared to those recovered in previous studies from global sites. One parasite lineage that matched between Galapagos birds and the migratory bobolink, Plasmodium lineage B, was the most common lineage detected in the global MalAvi database, matching 49 sequences from unique host/site combinations, 41 of which were of South American origin. We did not detect lineage B in brown‐headed cowbirds. The other Galapagos‐bobolink match, Plasmodium lineage C, was identical to two other sequences from birds sampled in California. We detected a close variant of lineage C in brown‐headed cowbirds. Taken together, this pattern suggests that bobolinks became infected with lineage B on the South American end of their migratory range, and with lineage C on the North American breeding grounds. Overall, we detected more parasite lineages in bobolinks than in cowbirds. Galapagos Plasmodium had similar host breadth compared to the non‐Galapagos haemosporidian lineages detected in bobolinks, brown‐headed cowbirds, and other grassland species. This study highlights the utility of global haemosporidian data in the context of migratory bird–parasite connectivity. It is possible that migratory bobolinks bring parasites to the Galapagos and that these parasites originate from different biogeographic regions representing both their breeding and overwintering sites.     

The impact of roads on the avifauna of páramo grasslands in Cajas National Park,Ecuador

National parks are an important tool for conserving biodiversity, particularly in areas of high biodiversity and endemism such as the tropical Andes. However, national parks often face a variety of stressors related to recreation, road construction and illegal extraction of natural resources. Unfortunately, the influence of these stressors for biodiversity is rarely well documented. Cajas National Park in Ecuador is no exception. Despite being traversed by the Cuenca-Molleturo-Naranjal road, effects of the road construction on biodiversity have not been determined. We therefore assessed the influence of road proximity on bird species richness and abundance as well as composition of bird habitat groups in Cajas National Park using transect walks at 25 m and 250 m distance to the road (overall 18 transects, each 1 km length). In total, we recorded 1110 individuals of 28 páramo bird species. Overall species richness did not differ between transects near and far from the road. Nevertheless, the average abundance of shrubby páramo species was significantly higher far from the road than near the road (Far = 36, Near = 25). Moreover, we found a tendency towards differences in the composition of bird habitat groups between transects near and far from the road. One aspect potentially driving the observed patterns was the increasing proportion of planted non-native woody tree species within páramo grassland near the road, which may have caused reduced abundances of shrubby páramo bird species there. While roads represented a clear impact on the composition of bird species in the páramo, the major effect seems to be driven by the introduction of non-native plant species along the roadside. In order to reduce the impact of roads to a minimum, we suggest that park managers should control the introduction of such plant species.     

Predicting species distributions across the Amazonian and Andean regions using remote sensing data

Aim We explore the utility of newly available optical and microwave remote sensing data from the Moderate Resolution Imaging Spectroradiometer (MODIS) and QuikSCAT (QSCAT) instruments for species distribution modelling at regional to continental scales. Using eight Neotropical species from three taxonomic groups, we assess the extent to which remote sensing data can improve predictions of their geographic distributions. For two bird species, we investigate the specific contributions of different types of remote sensing variables to the predictions and model accuracy at the regional scale, where the benefits of the MODIS and QSCAT satellite data are expected to be most significant. Location South America, with a focus on the tropical and subtropical Andes and the Amazon Basin. Methods Potential geographic distributions of eight species, namely two birds, two mammals and four trees, were modelled with the maxent algorithm at 1‐km resolution over the South American continent using climatic and remote sensing data separately and combined. For each species and model scenario, we assess model performance by testing the agreement between observed and simulated distributions across all thresholds and, in the case of the two focal bird species, at selected thresholds. Results Quantitative performance tests showed that models built with remote sensing and climatic layers in isolation performed well in predicting species distributions, suggesting that each of these data sets contains useful information. However, predictions created with a combination of remote sensing and climatic layers generally resulted in the best model performance across the three taxonomic groups. In Ecuador, the inclusion of remote sensing data was critical in resolving the known geographically isolated populations of the two focal bird species along the steep Amazonian–Andean elevational gradients. Within remote sensing subsets, microwave‐based data were more important than optical data in the predictions of the two bird species. Main conclusions Our results suggest that the newly available remote sensing data (MODIS and QSCAT) have considerable utility in modelling the contemporary geographical distributions of species at both regional and continental scales and in predicting range shifts as a result of large‐scale land‐use change.     

Where the wild birds go: explaining the differences in migratory destinations across terrestrial bird species

Given their large movement capacities, migratory birds have in principle a wide range of possible geographical locations for their breeding and non‐breeding destinations, yet each species migrates between consistent breeding and non‐breeding ranges. In this study, we use a macroecological approach to search for the general factors explaining the location of the seasonal ranges of migratory bird species across the globe. We develop a null model to test the hypotheses that access to resources, geographical distance, tracking of temperature, and habitat conditions (separately as well as considered together) have a major influence in the location of species’ migratory destinations, once each species’ geographical constraints are taken into account. Our results provide evidence for a trade‐off between costs associated with distance travelled and gains in terms of better access to resources. We also provide strong support to the hypotheses that all factors tested, with the exception of habitat, have a strong and additive effect on the global geography of bird migration. Indeed, our results indicate that species’ contemporary migratory destinations (i.e. the combination of their breeding and non‐breeding ranges) are such that they allow species to track a temperature regime throughout the year, to escape local competition and reach areas with better access to resources, and to minimise the spatial distance travelled, within the limitations imposed by the geographical location of each species. Our study thus sheds light on the mechanisms underpinning bird migration and provides a strong basis for predicting how migratory species will respond to future change.     

Why migrate during the day: a comparative analysis of North American birds

Migration can take place primarily during the day or at night, or during both depending on the species. Why the timing of migration varies among species has been the object of much research but the underlying ecological processes are still unclear. Proximally, migration timing may be influenced by the timing of other activities or may be more prevalent in species that migrate over long distances. Adaptive scenarios emphasize the reduction in flight costs at night especially in smaller species and the advantages of travelling in groups during the day to locate staging sites more efficiently. I used phylogenetic independent contrasts to examine these hypotheses in all North American nesting birds. I uncovered 24 evolutionary transitions in migration timing, most of which involved a switch from nocturnal to diurnal migration. Few of these transitions involved a concomitant change in the timing of foraging habits or migration distance. However, species in diurnal clades were larger, travelled in larger flocks and were generally more sociable than their nocturnal counterparts. The results support the hypotheses that a reduction in flight costs and the ability to pool information from companions are associated with migration timing in North American bird species.     

‘It is hard being the different one all the time’: gringos and racialized identity in lifestyle migration to Ecuador

This paper explores the identities that North American lifestyle migrants in Ecuador adopt as they adjust to life in a new racialized social environment. It is based on qualitative interviews with migrants from North America, as well as ethnographic field notes. North Americans describe their growing community in racialized terms and adopt a series of practices that demonstrate anxiety about their position in the racialized social order of Ecuador. The paper discusses the strategies that North American migrants engage in to diminish the importance of their racialized identities in Ecuador. I identify two main practices that complicate North American incorporation in Ecuador: self-policing practices that aim to optimize Ecuadorians' perceptions of them; and desires for integration and ethnic mobility, which seek to erase their ‘Otherness’.     

Conflicting constraints on male mating success shape reward size in pollen-rewarding plants

Premise

Pollen-rewarding plants face two conflicting constraints: They must prevent consumptive emasculation while remaining attractive to pollen-collecting visitors. Small pollen packages (the quantity of pollen available in a single visit) may discourage visitors from grooming (reducing consumptive loss) but may also decrease a plant's attractiveness to pollen-collecting visitors. What package size best balances these two constraints?

Methods

We modeled the joint effects of pollinators' grooming behaviors and package size preferences on the optimal package size (i.e., the size that maximizes pollen donation). We then used this model to examine Darwin's conjecture that selection should favor increased pollen production in pollen-rewarding plants.

Results

When package size preferences are weak, minimizing package size reduces grooming losses and should be favored (as in previous theoretical studies). Stronger preferences select for larger packages despite the associated increase to grooming loss because loss associated with nonremoval of smaller packages is even greater. Total pollen donation increases with production (as Darwin suggested). However, if floral visitation declines or packages size preference increases with overall pollen availability, the fraction of pollen donated may decline as per-plant pollen production increases. Hence, increasing production may result in diminishing returns.

Conclusions

Pollen-rewarding plants can balance conflicting constraints on pollen donation by producing intermediate-sized pollen packages. Strictly pollen-rewarding plants may have responded to past selection to produce more pollen in total, but diminishing returns may limit the strength of that selection.

     

An interaction between feeding rate and season affects fertility of sows

Trials were conducted in two piggeries in the summer-autumn and winter-spring periods to determine the effects of rate of feeding and type of housing (group or individual stalls) on farrowing rate. Sows were fed a low (1.6–2.0 kg per sow per day), moderate (2.5 kg per sow per day) or high (> 3.6 kg per sow per day) rate for the first 4 weeks postmating and from then until farrowing fed at a moderate (2.5 or 3.2 kg per sow per day) rate. The low level of feeding in the summer-autumn was associated with increased numbers of delayed returns to oestrus and low farrowing rate (< 75%) typical of the infertility that occurs at this time of the year. This same rate of feeding in winter-spring was associated with few delayed returns to oestrus and a high farrowing rate (> 85%). Feeding sows at the high rate during the summer-autumn significantly improved the farrowing rate reducing the adverse effect of season. Litter size was not affected by feeding rate during early pregnancy. The seasonal effect on farrowing rate was not evident in individually stalled sows fed a moderate level during pregnancy. This study demonstrated effects on farrowing rate of interactions between rate of feeding and season and type of accommodation and season. Current practice restricting feed intake postmating, aimed at reducing embryonic mortality and increasing litter numbers, appeared to contribute substantially to the decrease in farrowing rate commonly observed in summer-autumn. The optimal level of feeding to minimise the effect of season on fertility has yet to be defined. Accommodation of sows in individual stalls, rather than in groups, removed the seasonal effect on fertility.