Effects of density, climate, and supplementary forage on body mass and pregnancy rates of female red deer in Spain

The influence of short- and long-term (cohort) effects of climate and density on the life-histories of ungulates in temperate regions may vary with latitude, habitat, and management practices, but the life-histories of ungulates in the Mediterranean region are less well known. This study examined the short- and long-term effects of rainfall and absolute density on hinds in two of the southernmost populations of red deer (Cervus elaphus hispanicus) in Europe. One population received supplementary forage. Unlike more northerly latitudes, where red deer hinds lose body mass in winter as a result of adverse weather, in the Spanish populations, hinds did not lose body mass. Hinds in the population that received supplementary forage were heavier and more likely to become pregnant than were the hinds in the unsupplemented population. The likelihood of pregnancy occurring was strongly influenced by hind body mass; the proportion of yearlings that became pregnant was consequently lower in the unsupplemented population than in the population that received supplementary forage. Cohort effects on hind body mass (negative for density and positive for rainfall at birth) and on the probability of pregnancy (negative for density at birth) were apparent only in the unsupplemented population, which implies that supplemental feeding may partially compensate for negative density-dependent factors during early growth, and that supplemented deer hinds may experience reduced selection pressures. These results reflect the particular seasonal variation in the abundance and quality of food in Mediterranean habitats. The delayed effects of climate and density at birth on adult hind body mass and the prevalence of pregnancy probably affects population dynamics and constitutes a mechanism by which cohort effects affect the population dynamics in Iberian red deer. The management of Iberian red deer populations should take into account cohort effects and supplemental feeding practices, which can buffer density- and climate-dependent effects and reduce natural selection pressures.     

Seasonal change in the above-ground phytomass of an Agrostis-with-Festuca grassland in North-east Scotland

Summary

The above-ground standing crop of an Agrostis-with-Festuca grassland near Cabrach, Banff, was measured nine times in a 16-month period by harvesting sample plots. Standing crop peaked in late summer when the weight of live graminoid phytomass reached its seasonal maximum.     

Characteristics of Sasa nipponica grassland as a summer forage resource for sika deer on Mt Ohdaigahara, central Japan

Recently, the sika deer, Cervus nippon Temminck, population has increased on Mt Ohdaigahara, central Japan. The dwarf bamboo, Sasa nipponica Makino et Shibata, is a primary forage plant for sika deer in this area. To demonstrate the characteristics of S. nipponica grassland, especially as summer forage for sika deer, the habitat use intensity of sika deer was estimated by fecal densities, and biomass, growth rate, removal by deer and crude protein content were examined. Sika deer utilized the S.nipponica grassland on Mt Ohdaigahara during summer when the biomass, growth rate and crude protein content of S. nipponica were high. The recent increase in the deer population seems to be partly due to S.nipponica grassland being a favorable summer habitat.     

Lifetime reproductive success and density-dependent, multi-variable resource selection

Individuals are predicted to maximize lifetime reproductive success (LRS) through selective use of resources; however, a wide range of ecological and social processes may prevent individuals from always using the highest-quality resources available. Resource selection functions (RSFs) estimate the relative amount of time an individual spends using a resource as a function of the proportional availability of that resource. We quantified the association between LRS and coefficients of individual-based RSFs describing lifetime resource selection for 267 female red deer (Cervus elaphus) of the Isle of Rum, Scotland, from 1970 to 2001. LRS was significantly related to first- and second-order effects of selection for Agrostis/Festuca grassland and proximity to the sea coast (quality of forage within Agrostis/Festuca grassland was highest nearest the coast (ratio of short:long grassland)). The benefits of selecting for quality in Agrostis/Festuca grassland, however, traded-off with increases in LRS gained by avoiding conspecific density. LRS was inversely associated with local density, which was highest along the coast, and reproductive benefits of selecting Agrostis/Festuca grassland diminished with increasing density. We discuss the relevance of these results to our understanding of the spatial distribution of red deer abundance, and potential applications of our approach to evolutionary and applied ecology.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.     

Social rank,feeding and winter weight loss in red deer: any evidence of interference competition?

During winter at northern latitudes, large herbivores often exploit patches of concentrated, relatively high quality forage, which may lead to interference competition. The factors affecting success in contests and subsequent dominance rank, such as age and body weight, remain key issues in ungulate behavioural ecology. Maternal effects on offspring body weight are well known, but few studies have investigated if mothers social rank influence offspring rank. Moreover, no study has related dominance rank in ungulates to weight loss during winter. Outcomes of social interactions (n=7,609), feeding time and spatial position in red deer (Cervus elaphus) hinds and calves, and weight loss of calves, were registered from 1981 to 1996 at six winter-feeding sites within the county of Sør-Trøndelag in Norway. The level of aggressiveness was higher among calves than among adult hinds, and the factors determining the outcome of contests also differed. The initiator won the majority of interactions (more than 90% in both hinds and calves). Social rank was related to both age and body weight in adult hinds, and related to body weight and mother rank in calves. The relationship between feeding time and rank was non-linear. Feeding time was correlated with rank only among high ranked hinds, while there was no such relationship among low ranked hinds or calves. There was no correlation between winter weight loss and social rank in calves. Our study therefore underlines that, although frequent aggression is observed at artificial feeding sites of northern herbivores, this is not necessarily sufficient to give rise to interference competition.     

Dry season diets of sympatric ungulates in lowland Nepal: competition and facilitation in alluvial tall grasslands

Based on microhistological analyses of faecal material, we compared the early dry season diets of greater one-horned rhinoceros Rhinoceros unicornis, swamp deer Cervus duvauceli and hog deer Axis porcinus, which inhabit the same alluvial grassland habitat complex in lowland Nepal. Their diets were quite similar, both at the forage category level and within subcategories of graminoids and woody plants. Early successional tall grasses, especially Saccharum spontaneum, were the dominant food of all three species, underlining the key role of the threatened alluvial floodplains in large mammal conservation in South Asia. The two deer species ate significantly more graminoids (>66.5%) than did rhino (45.5%), and although they did not differ in proportions of graminoids, swamp deer ate significantly more late successional tall grasses (Narenga porphyrocoma and Themeda spp.) and short grasses (mainly Imperata cylindrica) than hog deer. The two deer consumed almost equal proportions of woody browse (ca. 10%), significantly less than that of rhino (33.0%). The prediction of the Jarman–Bell hypothesis, that large-bodied herbivores are less selective and subsist on lower quality graminoids than smaller ruminants, was not supported by the data. Based on this and previous studies in the same area we propose a conceptual model where the larger megaherbivores (rhino and elephant Elephas maximus) facilitate the smaller swamp deer and hog deer during the monsoonal growing season, while the smaller and more selective deer species outcompete the larger during the dry season when food is more limited. Owing to the all-year sprouting ability of S. spontaneum, facilitation may occur also in the dry season with low numbers of megaherbivores, thus accentuating competitive exclusion at higher deer densities.     

Habitat Interference by Axis Deer on White-Tailed Deer

Abstract: Many studies of interactions between exotic and native ungulates have not had temporal and spatial controls nor have they considered the types of competitive interactions that would allow coexistence. For exotic axis deer (Axis axis) and native white-tailed deer (Odocoileus virginianus) to coexist one species should be superior at interference competition and the other species should be superior at exploitative competition. We generated and tested predictions, based on body size and diet breadth, about habitat selection by white-tailed deer in the presence and absence of axis deer, dominance relationships, and time at sites provisioned with high quality forage. We conducted our study in treatment (axis and white-tailed deer) and control (white-tailed deer only) areas when both species were present and after axis deer were removed. We conducted vehicle surveys to determine habitat use of both species. At provisioned feeding sites we recorded aggressive behaviors and amount of time species spent at feeding sites alone and together. In the treatment area white-tailed deer selection for wooded habitat increased 2.1 times after axis deer were removed, whereas habitat selection by white-tailed deer was constant in the control area over the same time. At feeding sites axis deer were dominant to white-tailed deer; both species spent a significantly greater amount of time alone than at feeders together, but amount of time that individuals of each species spent at feeders did not differ. Axis deer were superior at interference competition, but white-tailed deer were not superior at exploitative competition; thus, species coexistence is unlikely. Whether white-tailed deer are negatively impacted by axis deer at spatial scales larger than our experiment probably depends on abundance of axis deer at larger spatial scales. Experiments of species interactions with temporal and spatial controls that consider types of competitive interactions increase a manager's understanding of when and how native ungulates may be negatively impacted by exotic ungulates.     

Free‐Ranging Red Deer Hinds Show Greater Attentiveness to Roars with Formant Frequencies Typical of Young Males

Acoustic cues present in the reproductive calls of many animal species potentially encode important information about the caller. Here, we test the response of a free‐ranging population of peri‐oestrus red deer hinds to variation in a specific acoustic cue to body size in the male roar, the formant frequencies. Our results revealed: (1) that hinds showed greater overall attention (judged by longer looking responses and lower response latencies) to roars simulating males of sub‐adult body size than to those simulating a large adult male and (2) that hinds without dependent offspring had greater looking responses to male roars and lower response latencies than hinds with dependent offspring to roars simulating sub‐adult males. These findings indicate that free‐ranging red deer hinds may use formants as acoustic cues to gauge the body size and maturity of males in their natural environment, possibly to facilitate earlier detection and avoidance of young stags that are known to harass them.     

Reproductive performance of farmed red deer (Cervus elaphus) in New Zealand. I. Descriptive data.

A longitudinal observational study of 15 red deer (Cervus elaphus) farms was carried out in New Zealand for 2 years from Mar 1992. Management of hind mobs and their composition during mating, calving, and weaning were recorded. About 2700 hinds were individually monitored for live weight, body condition score and reproductive success. All hinds were pregnancy tested in June by ultrasound and classified as having conceived before May 1, after May 1, or as being not pregnant. Calving dates and dam-offspring pairs were recorded on four farms. The lactational status of hinds and live weight of calves were recorded at weaning. Mean pre-mating live weights of yearling hinds were 81.3-82.8 kg and of adult hinds 98.0-98.7 kg, respectively, for each year of study. The percentage of yearling and adult hinds conceiving before May ranged from 8.3 to 95.0% and from 77.6 to 98.4%, respectively, between farms. Overall, the proportion of yearling and adult hinds not pregnant was 15.3% and 3.2%, respectively. Pregnancy rates at scanning of adult hinds within mating mobs were generally over 90%, and 41.7% of mating mobs had 100% pregnancy rates. In contrast, the pregnancy rates of yearling hind mobs were more variable, with five mobs below 50%, and 34.1% of mating mobs achieving 100% pregnancy rate. Estimated in utero loss rates from pregnancy diagnosis to calving were 0.66 and 0.79% for yearling and adult hinds, respectively. From four farms, median calving dates of yearling and adult hinds were Dec 13 and Nov 30, respectively. Overall, calf survival rates to weaning of yearling and adult offspring were 84.1 and 91.6%, respectively, while the reproductive efficiencies (number of calves weaned per hind mated) were 70 and 83.6%, respectively. Farm mean weaner live weights standardised on Apr 1 ranged from 42-59 kg and 39-51 kg for stags and hinds, respectively. These data are currently the best estimates of reproductive parameters of New Zealand red deer herds, and highlight a wide variability in reproductive indices between farms.     

Kinship Discrimination and Effects on Social Rank and Aggressiveness Levels in Iberian Red Deer Hinds

Kin recognition is a widespread phenomenon that allows individuals to benefit by enhancing their inclusive fitness, and one of its most common forms is reducing aggressiveness towards relatives. We carried out an experiment with Iberian red deer hinds (Cervus elaphus hispanicus) in order to examine kin biases in dominance behaviour and its consequences on social rank. Three enclosed groups (n = 36, 23 and 21, respectively) were monitored during two lactation seasons and social rank hierarchies were assessed by analysing aggressive interactions matrices with Matman 1.1 software. Aggressive interactions between related hinds was significantly smaller than expected (χ² = 5.02, df = 1, p = 0.025), not only between mother and daughter but also in second and third kinship degrees. Although rates of aggressiveness were similar to data published relating free‐ranging C. e. scoticus, aggressive interactions with relatives were significantly smaller (χ² = 39.0, df = 1, p < 0.001). This reduction of aggressiveness between related hinds was not the result of these hinds having a lower social rank: social rank was only related to age and weight, but not to kinship degree, calf sex or calving date. The decrease of aggressiveness towards first‐, second‐ and third‐degree relatives shows a complex kin recognition system in deer. Possible nepotistic roles in lactation include preventing milk thefts by non‐kin and disturbing feeding of unrelated hinds.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

Movement pattern of red deer during drive hunts in Sweden

We investigated the movement pattern of GPS-collared red deer (Cervus elaphus) during drive hunts with loose dogs. In 46 flights of hinds (N?=?9), the median flight distance was 2.5 km (range 0.4–15.0). In 28 % of the flights, the hind left its home range. Average time before returning to home range was 23 h (range 2–88). Hinds in a less forested site left their home ranges more often, fled longer distances, moved at higher speed, and returned later than hinds in a homogenous forest. Speed of movement increased with number of hunts in the less forested site. In eight flights of stags (N?=?4), the median flight distance was 5.1 km (range 2.2–13.3). The start of hunting season seemed to trigger stags’ departure to their wintering areas. Knowledge of reactions to disturbance can aid game management to choose hunting methods and dogs that cause less disturbance but may also be used to deter deer from areas where they damage crops and forest plantations.     

Migratory strategies, fawn recruitment, and winter habitat use by urban and rural mule deer (Odocoileus hemionus)

We examined the migratory strategies of mule deer (Odocoileus hemionus) using adjacent urban and rural winter ranges in northern Utah in relation to deer demography and patterns of habitat use. Urban deer were more likely to be migratory than rural deer, even though migratory animals from the two herds intermixed on a common, high-elevation summer range. Urban deer exhibited lower fawn recruitment than rural deer; but within each herd, demographic characteristics of migratory and nonmigratory animals suggested that game theory explained the ratios of deer adopting each behavior. Estimates of animal numbers and available habitat did not reveal clearly whether deer densities differed on the two winter ranges. However, patterns of habitat use by urban deer were so clustered around areas of concealment vegetation that animals probably experienced higher local densities than rural animals. In addition, the clustered patterns of habitat use by urban deer resulted in incomplete use of available forage. This may have contributed to the relatively poor fawn recruitment by urban deer, a phenomenon that appeared to be perpetuated by their strong fidelity to winter range.     

Advertising individual identity by mother and adolescent contact calls in Siberian wapiti Cervus elaphus sibiricus

Individualistic contact calls facilitate mother‐offspring reunion after separation. However, in many mammals, both the acoustic structure and individuality of contact calls differ between mother and young. In contrast, in Siberian wapiti Cervus elaphus sibiricus, contact calls are similar in the acoustics between mother and young, whereas effects of this similarity on vocal individuality were not investigated. In this study, we analyzed acoustic differences between closed‐mouth (nasal) and open‐mouth (oral) contact calls and examined individuality of the most usual oral calls of 19 Siberian wapiti (9 hinds and 10 5–6‐month adolescents) emitted in response to mother‐offspring separation. In the oral calls, the values of frequency and power variables were higher than in the nasal calls. Calls of hinds and adolescents did not differ by the maximum fundamental frequency and duration, whereas the peak frequency was higher in the young. Discriminant function analysis (DFA) based on 11 acoustic variables of oral calls accurately classified to individual 92.5% of hind calls and 96.9% of adolescent calls (chi‐square test for differences between hinds and adolescents, p = 0.19). Variables mainly contributing to vocal identity (duration, start, and maximum fundamental frequency) were the same in calls of mothers and adolescents. We conclude that similarities in the acoustics calls of mothers and adolescents mean that they do not differ in their potential for encoding individual identity, suggesting a mutual process of mother‐offspring vocal recognition in Siberian wapiti.     

Why do migrants move downhill? The effects of increasing predation and density on red deer altitudinal migration in temperate Carpathian forests

Resource selection by ungulates is driven by trade-offs between foraging and predation avoidance or by intraspecific competition. Ungulates use migratory flexibility to optimize access to spatially and temporally variable resources across seasons, sometimes even adaptively switching between migrant and resident strategy as conditions change. After an 80% increase in red deer (Cervus elaphus) population and simultaneous recovery of wolves (Canis lupus) in the Kremnica Mountains, Slovakia, a significant portion of the deer population started to migrate downhill (<?700 m) to marginal habitats during winter. Building on available spatial data on forage availability, predation risk, and deer abundance, we tested for differences in habitat selection of migrant and resident male red deer to assess possible reasons for this change. On high-altitude (700–1100 m) summer ranges, deer were not forced to trade-off forage to avoid predation within their home ranges. However, during winter, residents remaining on high-altitude ranges selected for areas with highly abundant forage only under low predation risk or at high deer abundance. Downhill migration exposed migrants to 15% lower forage availability but simultaneously reduced wolf predation risk by 39% relative to residents. Consequently, the limited access to forage resources at low-altitude ranges have reduced antler growth, especially in young males. Our study represents one of few that address the role of predation risk in driving seasonal migrations in temperate systems where snow is not likely to be the major driver of migration to low-altitude winter ranges.     

Effects of lactation on feeding behaviour and habitat use in wild Red deer hinds

In many mammals, the energetic costs of lactation double a female's daily energy requirements on her activity budgets, food choice and ranging behaviour–but the effects of lactation have rarely been documented. In this paper, the ecological consequences of lactation in free-ranging Red deer ( Cervus elaphus ) hinds on the lsle of Rhum is examined by comparing the feeding behaviour, habitat use and ranging behaviour of hinds that were supporting calves (milk hinds) with hinds that did not have calves (yeld hinds). In summer, milk hinds graze for approximately two hours longer per day then yeld hinds. This difference has disappeared by the latter half of the winter, when the grazing budgets of milk and yeld hinds are similar. Both in summer and winter milk hinds spend more time grazing on the most strongly selected plant communities (short, herb-rich greens) where standing crops are low but food quality is comparatively high. Inter-year variation in habitat use by milk hinds is also closely related to variation in rainfall—while this is not the case among yeld hinds. Both in summer and winter, yeld hinds range more widely than milk hinds.     

Influence of level of nutrition during late pregnancy on reproductive productivity of red deer (2) Adult hinds gestating wapitixred deer crossbred calves

The present study aimed to relate feed intake of red deer hinds in the later stages of gestating wapitixred deer crossbred foetuses on dam body condition, gestation length, birth weight and calf growth. Multiparous hinds (N=18) conceiving at known dates to either wapiti (n=12) or red deer (n=6) sires were housed in individual pens from days 150-220 of pregnancy, during which time they were offered either ad libitum access to pelletised rations (n=6 crossbred-bearing hinds [HH] and n=6 red deer-bearing hinds [RH]) or a restricted offer (n=6 crossbred-bearing hinds [HL]) set at 70% of the average ad libitum intake of HH hind in the previous week. Hinds were returned to pasture at day 220 and calving was closely monitored. Liveweights, body condition score (BCS), and lactation score (LS) of hinds were recorded weekly from day 130 of pregnancy until calves were weaned at 12 weeks of age. Calves were tagged and weighed at birth, and subsequently weighed at 7 and 12 weeks of age. HH and RH hinds exhibited similar patterns and levels of MEI/kg0.75, which peaked at 7.8 MJME/kg0.75 at day 220. HL hinds peaked at approximately 5 MJME/kg0.75 and showed significantly lower rates of liveweight gain during pregnancy. Interestingly, both crossbred-bearing groups initiated mammary development in advance of the RH hinds. While there were significant effects of foetal genotype on mean gestation length (239 days versus 234 days for crossbred versus red deer) and mean birth weight (14.5 kg versus 10 kg), the nutritional contrast for gestation length of crossbred-bearing hinds (i.e. HH versus HL) was not significant but approached significance for birth weight (14.5 kg versus 11.9 kg; P=0.06). Regression analysis revealed weak relationships between changes in hind liveweight and gestation length (P>0.05) but a significant relationship with birth weight (P<0.05). However, change in hind BCS was significantly related to both gestation length and birth weight. Crossbred calves reared by HH hinds were 30% heavier at 7 and 12 weeks of age than the red deer calves. However, those reared by HL hinds were significantly lighter than their genotype contemporaries and only marginally heavier than the red deer calves. These results generally contrast with the previous studies on red deer hinds gestating red deer foetuses [Asher, G.W., Mulley, R.C., O'Neill, K.T., Scott, I.C., Jopson, N.B., Littlejohn, R. 2004. Influence of level of nutrition during late pregnancy on reproductive productivity of red deer, (1) Adult and primiparous hinds gestating red deer calves. Anim. Reprod. Sci., in press] and indicate that the genetically determined higher growth requirements of crossbred foetuses may override any mechanism of compensatory control of gestation length at the expense of calf birth weight. Furthermore, there were marked carryover effects of late gestational feeding on crossbred calf growth and their dam's BCS that highlight the high nutritional demands of lactation.     

Lactation under induced five-month delay reproduction in Iberian red deer (Cervus elaphus hispanicus)

In Iberian deer, calf birth date affects both milk production and composition, as well as the weight change of hind during lactation and calf growth. This study examines a five-month induced delay in calving (autumn group) in a group of 19 Iberian hinds compared to a group of 13 standard calving hinds (spring group). Fertility in the autumn group was half that of the spring one. From the former, a further two calves out of six died before or soon after birth. Successfully lactating hinds showed a greater concentration in protein, fat and lactose (P < 0.001), but did not differ from spring group in total milk yield, calf gain or weight change of hinds during lactation. However, after correcting for the greater calf or hind body weight at calving of the autumn group, this group showed lower values for these variables than the spring group. Results show that at least in Iberian deer, counter-season reproduction is possible, and that the effects on lactation variables are similar to those of a natural delay in calving.     

Skewed distributions of lifetime reproductive success: beyond mean and variance

Lifetime reproductive performance is quantified here by the LRS (lifetime reproductive success), the random number of offspring an individual produces over its lifetime. Many field studies find that distributions of LRS among individuals are non‐normal, zero‐inflated and highly skewed. These results beg the question, what is the distribution of LRS predicted by demographic models when the only source of randomness is demographic stochasticity? Here we present the first exact analysis of the probability distribution of LRS for species described by age + stage models; our analysis starts with estimated vital rates. We illustrate with three examples: the Hadza, human hunter‐foragers (age‐only), the evergreen tree Tsuga canadensis (stage‐only) and Roe deer, Capreolus capreolus (age + stage). For each we obtain the exact distribution of LRS, but also calculate and discuss the first three moments. Our results point to important questions about how such LRS distributions affect natural selection, and life history evolution.