Host selection and sex allocation in Leptomastidea abnormis, a parasitoid of the citrus mealybug Planococcus citri

Leptomastidea abnormis (Girault) and Leptomastix dactylopii (Howard) [Hymenoptera: Encyrtidae] are the two parasitoid species frequently used for biological control of the citrus mealybug Planococcus citri (Risso) [Homoptera: Pseudococcidae]. As part of a programme to determine the best control practice, host size selection and sex allocation were studied in L. abnormis, and compared with those of L. dactylopii. L. abnormis showed a preference for the second size class, but also attacked third and fourth size class and adult mealybugs, while in previous studies L. dactylopii was found to prefer the third and fourth size class and adults, and never attacked second size class mealybugs. Female L. abnormis reared from parasitized mealybugs had a higher mean body mass than males, as is predicted by Charnov's theory on sex allocation. However, sex ratios of emerging L. abnormis did not differ between mealybug size classes. These results imply possibile coexistence of the two parasitoids. As field evidence suggests a more complex interaction between the two species, studying direct competition would be a useful next step.     

Predicting development of the mealybug parasitoids Anagyrus pseudococci, Leptomastix dactylopii and Leptomastidae abnormis under glasshouse conditions

Developmental rates of three encyrtid mealybug parasitoids were examined under constant, cycling and glasshouse temperatures. Under constant conditions the rate of development increased with rising temperature up to 35°C, but a constant 40°C was lethal. Under cycling conditions, 12 h periods at 40°C could be tolerated. Developmental rates under cycling conditions could be calculated from constant temperature data, by incrementing development on an h/h basis. A new technique was developed to determine the lower thresholds for development, using cycling regimes and to calculate the rates of development over short periods at high temperature. A previously untested polynomial regression technique was used to predict mean developmental periods under glasshouse conditions. These predictions were considerably more accurate than day-degree (D^o) predictions when substantial periods at extreme temperatures were involved, but not significantly different from day-degree predictions under hot-house conditions. In addition a method was developed to calculate a 95% range for emergence, which gave consistently good estimates of first and last eclosion observed under fluctuating conditions.

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Résumé Les vitesses de développement de 3 parasitoïdes de cochenilles ont été étudiées dans les conditions de température de serre, avec thermopériode ou en température constante. En conditions constantes, la vitesse augmente jusqu'à 35°C, mais est létale à 40°C. Avec thermopériode, une thermophase de 40°C peut être tolérée pendant 12 heures. Les vitesses de développement sous thermopériodes pourraient être calculées à partir des données obtenues en conditions constantes, modifiées en fonction des durées respectives des cryophases et thermophases. Une méthode originale est présentée pour obtenir les seuils inférieurs de développement, en utilisant des thermopériodes, et pour calculer les vitesses de développement à températures élevées.Une méthode de régression polynomiale, non vérifiée préalablement, a été utilisée pour prévoir les durées moyennes des développements sous serre. Ces estimations sont beaucoup plus sûres que les prédictions basées sur le concept de degré-jour (D^o), lorsque les températures extrêmes durent longtemps, mais en serre chaude elles ne présentent aucune différence significative. De plus, une méthode d'évaluation des 95% d'émergences, a donné de bonnes estimations des premières et dermières émergences en conditions périodiques.

     

Kairomonal response of the parasitoid Anagyrus spec. nov. near pseudococci to the sex pheromone of the vine mealybug

The occurrence of a kairomonal response of the parasitoid Anagyrus spec. nov. near pseudococci (Hymenoptera: Encyrtidae) to (+)‐(1R,3R)‐cis‐2,2‐dimethyl‐3‐isopropenyl‐cyclobutanemethanol acetate (PcA, namely, planococcyl acetate) and (S)‐(+)‐lavandulyl senecioate (LS), the respective female sex pheromones of its hosts, the citrus mealybug, Planococcus citri (Risso) and the vine mealybug, Planococcus ficus (Signoret) (Homoptera: Pseudococcidae) was investigated. Attraction to the pheromones was tested by employing pheromone traps in field trials and by static air olfactometer bioassays in the laboratory. Female wasps showed a significant response to LS, in both field and olfactometer experiments. No significant response was registered to the sex pheromone of P. citri. Despite the similarity between the structures of LS and its analogue (S)‐(+)‐lavandulyl isovalerate (LI), no significant response to the latter compound was observed. It seems that differences between the structures of the carboxylate moiety of the respective molecules (LS and LI) markedly affect the kairomonal attractiveness to the parasitoid. The kairomonal response of Anagyrus spec. nov. near pseudococci was neither influenced by the host habitat nor by the host species on which it developed. This suggested innate behaviour of Anagyrus spec. nov. near pseudococci, possibly derived from evolutionary relationships between the parasitoid and P. ficus. The practical implications of the results are discussed.     

Developmental time,longevity, reproductive capacity and sex ratio of the mealybug parasitoid Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera: Encyrtidae)

Key biological parameters of Anagyrus sp. nr. pseudococci reared on the citrus mealybug Planococcus citri (Risso) were determined. The mean longevity of female wasps was 21.1 days, the median lethal time (LT₅₀) was 22 days, the mean number of offspring per female was 30.2 wasps and the sex ratio of the progeny was slightly male biased (1.1 males per female). Wasps fed with water showed a significantly lower longevity compared to that recorded for wasps fed with syrup for bee nutrition mixed with water. The longevity of female wasps was significantly higher compared to males when the syrup-water solution was provided as food. Longevity was significantly higher for female wasps fed with syrup and water in comparison with those exposed to host mealybugs. The mean development time of female and male wasps was 14.7±0.1 and 14.5±0.1 days, respectively.     

Predation of the apefly,Spalgis epius (Lepidoptera: Lycaenidae) on citrus mealybug,Planococcus citri (Hemiptera: Pseudococcidae)

The citrus mealybug, Planococcus citri (Risso) (Hemiptera: Pseudococcidae) is a serious pest of economically important crops worldwide. The apefly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae) is a potential predator of various species of mealybugs. Earlier investigation on its daily preying capacity and preference for prey stages on P. citri is incomplete. Hence, a study was conducted to find out the daily prey consumption ability and preference for prey stages by different larval instars of S. epius reared on P. citri in the laboratory. Through the 8-day developmental period with four larval instars, the daily prey consumption of S. epius increased from the first to the seventh day and decreased on the eighth day prior to the prepupal stage. Generally, there was a significant difference in the prey consumption on different days. When the prey stages were offered separately, the first to fourth instar larva of S. epius consumed, respectively, a mean of 199.6, 722.6, 1908.8, and 4625.6 eggs or 21.5, 77.0, 168.5, and 670.5 nymphs or 3.2, 7.2, 16.0, and 35.1 adults of P. citri. When an S. epius larva was fed on P. citri eggs, nymphs and adults separately, it consumed a mean of 7456.7 eggs, 937.6 nymphs, or 62.3 adults during its entire development. When the prey stages were offered all together, a single S. epius larva consumed 2618.4 eggs, 170.4 nymphs, and 39.7 adults of P. citri throughout its entire development. The study revealed that S. epius is a voracious predator of P. citri and thus could be utilized as a major biological control agent.     

Host selection behaviour and specificity of the solitary parasitoid of mealybugs Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera,Encyrtidae)

The host selection behaviour of Anagyrus sp. nr. pseudococci was compared in no-choice tests among five mealybug species of different geographical and phylogenetic origin, including the Mediterranean native host, Planococcus ficus and four exotic mealybug species, one of the same genus, Pl. citri, two Pseudococcus species, Ps. calceolariae and Ps. viburni and a more distant one, Phenacoccus peruvianus. All five studied mealybug species were recognised by the parasitoid as potential hosts and parasitised, but the behavioural pattern of host recognition, host handling and the level of host acceptance of Anagyrus sp. nr. pseudococci significantly varied among the five studied species, indicating a clear preference for the two Planococcus species, Pl. ficus in particular. The results suggest that A. sp. nr. pseudococci has a broader host range and a more generalist behaviour in comparison with other Anagyrus species. Practical implications of the findings are discussed.     

Evaluation of the synthetic sex pheromone of the obscure mealybug,Pseudococcus viburni,as an attractant to conspecific males,and to females of the parasitoid Acerophagus maculipennis

The obscure mealybug, Pseudococcus viburni (Signoret) (Hemiptera: Pseudococcidae), is a cosmopolitan pest. In New Zealand, recently introduced management tools include the host‐specific parasitoid Acerophagus maculipennis (Mercet) (Hymenoptera: Encyrtidae) established in 2001, and pheromone‐baited monitoring traps available since 2005. Red delta traps baited with rubber septum lures impregnated with 4.0 μg of the mealybug synthetic sex pheromone, placed in apple orchards in Hawke's Bay and Nelson, trapped both male P. viburni and female A. maculipennis. Two generations of both species per year were discernible, but numbers were low in spring and parasitoids were not trapped during winter (June to September). Male P. viburni catches reached a plateau at a pheromone dose of ca. 1.0 μg per lure but numbers of A. maculipennis per trap increased up to 100 μg per lure, the maximum dose tested. A mathematical model showed that the lures had a half‐life of about 7.4 days and were most attractive to P. viburni with a dose of 0.19 μg, and that the trap effectiveness decreased rapidly once the release rate dropped below the optimum. The model also predicted that the initial pheromone dose should be increased from 0.19 to 5.41 μg per lure as the desired period of deployment increased from 0 to 9 weeks. A dose of 4.0 μg had an initial relative effectiveness of about 55%, reached peak effectiveness after about 5 weeks, and fell to 55% relative effectiveness again after about 8.3 weeks. We conclude that an initial pheromone load of 4.0 μg is appropriate for practical monitoring of P. viburni during the New Zealand summer. Future applications of the sex pheromone for managing the pest and parasitoid are discussed.     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.     

Influence of egg load and oviposition time interval on the host discrimination and offspring survival of Anagyrus pseudococci (Hymenoptera: encyrtidae), a solitary endoparasitoid of citrus mealybug, ++Planococcus citri (Hemiptera: pseudococcidae)

Oviposition and host discrimination behaviour of unmated Anagyrus pseudococci (Girault), an endoparasitoid of the citrus mealybug Planococcus citri (Risso), were investigated in the laboratory. Female parasitoids were able to discriminate between parasitized hosts and healthy ones. The mean number of ovipositions was significantly higher in unparasitized than in parasitized hosts. Conspecific-superparasitism occurred more often than self-superparasitism. Changes in consecutive ovipositions over three hours by A. pseudococci suggested that egg load influenced the discrimination behaviour of the parasitoids, with females which had low egg loads mostly avoiding oviposition in already parasitized hosts at time intervals ranging from 0 h to 96 h, and distributing their eggs in the high quality (unparasitized) hosts. The parasitized hosts were rejected more commonly through antennal perception of external markers than during ovipositor probing which could have encountered internal markers but this relationship changed with increasing time after oviposition. The parasitoid's oviposition rate in unparasitized and conspecific-parasitized hosts varied at the different oviposition time intervals when the females had fewer eggs in the ovaries. Percentage emergence of parasitized offspring was not significantly influenced by whether they developed in single or superparasitized mealybugs. The significance of host discrimination by A. pseudococci is discussed.     

不同离体寄主及其成熟度对亚洲柑桔木虱存活的影响

【目的】亚洲柑桔木虱Diaphorina citri Kuwayama的虫口密度及活动与柑桔黄龙病的田间传播、流行有十分密切的关系,寻找合适的室内饲养条件能够便于观察和研究其生物学特性和传病机制。【方法】本文以九里香Murraya exotica(L.)Mant.、酸桔Citrus sunki Hort.ex Tanaka和马水桔Citrus reticulata Blanco.cv.Mashuiju 3种寄主植物不同成熟度离体梢为研究材料,(1)比较亚洲柑桔木虱卵分别在3种寄主植株和离体嫩梢上的孵化率;(2)比较3种寄主不同成熟度离体梢对各龄若虫存活率和蜕皮的影响;(3)比较3种寄主不同成熟度离体梢对成虫存活率的影响。【结果】(1)亚洲柑桔木虱在植株嫩梢上卵的孵化率高于离体嫩梢,九里香表现最明显;(2)低龄若虫在叶片未完全展开的离体嫩梢上存活率最高,而高龄若虫在完全展开的嫩梢上最高;(3)用离体成熟梢饲养柑桔木虱成虫存活率高于离体嫩梢,酸桔和马水桔离体梢饲养的存活率较九里香高。【结论】用寄主植物离体梢饲养的亚洲柑桔木虱卵孵化率和若虫、成虫存活率都较高而且较稳定,该方法可用于这种木虱的室内繁殖中。     

Effect of host availability on reproduction and survival of the parasitoid wasp Trichogramma minutum

Abstract.

• 1 We tested the hypothesis that females of the egg parasitoid, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), could adjust their fecundity schedule according to host availability and that there was a negative correlation between reproduction and survival in these wasps.
• 2 Newly-emerged females were provided with an unlimited or limited number of hosts in the first trial and with either unlimited, limited or zero hosts in the second trial.
• 3 When hosts were unlimited, wasps had the highest rate of reproduction in the first day, which decreased dramatically thereafter. When hosts were limited, wasps from the two trials differed in their response. In Trial I, females with limited hosts had lower first-day fecundity than, and the same subsequent-day fecundity as, those with unlimited hosts. However, in Trial II, females with limited host had a lower first-day but a higher subsequent-day fecundity than those with unlimited hosts. This indicates variation in Trichogramma's ability to shift its fecundity schedule in response to host availability.
• 4 There was a positive (rather than a negative) correlation between reproduction and survival. Wasps that oviposited (in host-unlimited treatment) had greater longevity than those that could not (in host-unavailable treatment).
• 5 The sex ratio of the progeny produced by wasps in both host-unlimited and limited treatments shifted gradually from a female to a male bias as the wasps aged.
• 6 We consider the ability of parasitoids to adjust their fecundity schedule as an adaptation to changing host resources and discuss our findings with regard to theories of life history evolution.

     

Temperature-dependent growth rates for the cassava mealybug, Phenacoccus herreni, and two of its encyrtid parasitoids, Epidinocarsis diversicornis and Acerophagus coccois in Colombia

Developmental rates for Phenacoccus herreni Cox & Williams were determined at 18°, 20°, 22°, 25°, 30° and 35°C for the egg, all juvenile male and all juvenile female stadia. Longevity was determined for adult females and adult males. Developmental rates for the P. herreni parasitoid Epidinocarsis diversicornis (Howard) were determined at 18°, 20°, 25° and 30°C for the oviposition-to-mummy-formation period and the mummy-formation-to-adult-eclosion period. Developmental rates were determined for the P. herreni parasitoid Acerophagus coccois Smith for the same two life stages at 20°, 25° and 30°C. Least-squares-derived polynomial equations or logistic equations were fitted to each data set (except for A. coccois) so that rates could be interpolated for temperatures between observed points for use in an analysis of the impact of these parasitoids on population dynamics of P. herreni. Results of this analysis are presented separately.

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Résumé Les vitesses de développement des oeufs et de tous les stades larvaires mâles et femelles de P. herreni Cox & Williams ont été déterminées à 18°, 20°, 22°, 25°, 30° et 35°C. La longévité des adultes mâles et femelles a été déterminée. Les vitesses de développement de la ponte à la formation du cocon et de celleci à l'émergence de E. diversicornis Howard, encyrtide parasite de P. herreni ont été déterminées à 18°, 20°, 25° et 30°C. Il en a été de même pour un second encyrtide parasite A. coccois Smith à 20°, 25° et 30°C. Les équations polynomiales des derniers carrés dérivés et les équations logistiques ont été ajustées pour chaque lot de données (à l'exception de A. coccois) de façon à ce qu'elles aient pu être interpolées pour analyser l'impact de ces parasites sur la dynamique de population de P. herreni. Les résultats de cette analyse sont présentés à part.

     

Brood parasitism by cowbirds: risks and effects on reproductive success and survival in indigo buntings

We observed brood parasitism by brown-beaded cowbirds (Molothrusater) on indigo buntings (Passerina cyanea) and estimated dieimpact of parasitism on the success of the individual buntingsin their current nests and in their future survival and reproduction.Rates of parasitism over 8 years were 26.6% in 1040 nests and19.8% in 693 nests in two areas in southern Michigan. Risk ofparasitism was high early in the season; half the bunting nestswere begun after the end of the cowbird season. Risk was independentof female age, plant containing the nest, or habitat The immediatecost of parasitism was 1.19 and 1.26 fewer buntings fledgedper nest. Bunting success was lower in parasitized nests withcowbird eggs (nests were more likely to be deserted or predated),lower when the cowbird nestling failed (nests were more likelyto be predated), and lower when the cowbird fledged (fewer buntingsfledged) compared to nonparasitized nests. Costs were due toremoval of a bunting egg when die cowbird laid its own egg andto competition for parental care of the cowbird and buntingnestlings. Buntings that fledged from nests where a cowbirdalso fledged were only 18% as likely to survive and return totheir natal area in the next year as buntings from nests wherea cowbird did not fledge. Long-term effects of cowbird parasitismon adult breeding later in the season, survival to the nextseason, and reproductive success in the next season were negligiblewhen compared between birds that reared a cowbird and birdsthat reared only a bunting brood, or between birds that wereparasitized and birds that escaped parasitism. The results indicatelittle long-term cost of brood parasitism on individual fitnessof adult buntings beyond the impact on the current nest andthe survival of buntings that fledge from it; nearly all costis to the parasitized brood.     

The effect of Jacobin Cuckoo Clamator jacobinus parasitism on the body mass and survival of young in a new host species

The southern African subspecies of Jacobin Cuckoo Clamator jacobinus serratus is a brood parasite of a range of host species. While Jacobin Cuckoos do not evict host young, previous research has found that host young rarely survive the nestling period. Here we provide the first records of Jacobin Cuckoo parasitism of a new host species, the Southern Pied Babbler Turdoides bicolor. We investigate rates of brood parasitism and the survival of host young. The Southern Pied Babbler is one of the largest recorded hosts for Jacobin Cuckoos and, unusually, we find that host young tend to survive the nestling period and maintain similar body mass to host young in unparasitized broods. However, host young were less likely to survive to independence than young raised in unparasitized nests, suggesting a post‐fledging reproductive cost to hosts.     

The effects of body size on mating and reproduction in Brachinus lateralis (Coleoptera: Carabidae)

Abstract. 1. The causes and reproductive consequences of body size variation of Brachinus lateralis Dejean, a parasitic carabid beetle, were investigated.
2. Body size variation occurs within and between sites. Host size has a major influence on body size of the adult.
3. Fecundity is positively correlated with body size. Egg size is not correlated with body size.
4. Mating males tend to be larger than non-mating males. There is a positive correlation of body sizes in mating pairs.
5. Limited opportunity for host choice may maintain size variation despite the advantages of large size.
6. The non-random patterns of mating for a species without obvious intrasexual aggression suggest that subtle means of male-male competition or female choice may be important.     

Host feeding and ovipositor insertion as sources of mortality in the mealybug Phenacoccus herreni caused by two encyrtids, Epidinocarsis diversicornis and Acerophagus coccois

Females of Epidinocarsis diversicornis (Howard) killed cassava mealybugs, Phenacoccus herreni Cox & Williams, through host feeding and piercing (i.e. ovipositor insertion and its consequences). Parasitoids fed on 9% of 592 hosts in which ovipositor insertion was achieved. For hosts which were attacked but not fed on, mortality in the first 72 h was significantly greater than that of controls and was highest in the first nymphal instar. No host feeding by Acerophagus coccois Smith was observed in 494 attacks. Mortality due to piercing was significantly greater than control values, but did not differ between host life stages.

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Résumé La prise de nourriture et les piqûres (c'est-à-dire l'insertion de la tarière et ses conséquences) de Epidinocarsis diversicornis tuent Phenacoccus herreni. Les adultes s'alimentent sur 9% des 592 cochenilles dans lesquelles la tarière a pénétré. La mortalité au cours des 72 premières heures est beaucoup plus élevée chez les hôtes attaqués mais qui n'avaient pas subi une prise de nourriture, que chez les témoins; elle était particulièrement élevée chez les cochenilles du premier stade larvaire. Pour 494 attaques par Acerophagus coccois, aucune prise de nourriture n'a été observée; la mortalité était plus élevée que chez les temoins, sans qu'il y ait eu de différences suivant les stades.

     

Beyond fecundity and longevity: trade-offs between reproduction and survival mediated by behavioural responses of the seed beetle, Callosobruchus maculatus

Abstract.  Many studies of life-history traits have failed to find trade-offs where they are predicted by theory. A hypothesis that explains the lack of trade-offs between fecundity and longevity in the seed beetle, Callosobruchus maculatus , is proposed. By manipulating host availability time and host size, trade-offs mediated by behavioural responses of the female to adapt to environmental change are tested. Females show no decrease in lifetime fecundity when host availability time is limited to only 4 h on each day. However, longevity significantly increases when the female is provided with small beans after host deprivation. Because neither acquisition, nor utilization by females of these four manipulation treatments significantly differs, studies are carried out to demonstrate whether the energy shifted from increased longevity without decreasing fecundity. Providing abundant small or large beans each day directly after host deprivation, significantly increases the number of daily eggs laid by the female for several days, whereas the female decreases the uniformity of her egg dispersion only when small beans are provided. Therefore, the female shows a response to a change in the environment by adjusting egg-laying rate and/or egg-dispersion pattern. This may change the traits of reproduction and survival. Because energy allocations can be shifted between components of reproduction (e.g. host-selection behaviour and fecundity) or between reproduction and survival, fecundity and longevity may be inappropriate indices for trade-off analyses in this study. A framework for exploring the costs of reproduction mediated by physiological and behavioural changes in C. maculatus is proposed and discussed.     

Interacting effects of age,density, and weather on survival and current reproduction for a large mammal

Individual‐based study of natural populations allows for accurate and precise estimation of fitness components and the extent to which they might vary with ecological conditions. By tracking the fates of all 701 horses known to have lived on Sable Island, Canada, from 2009 to 2013 (where there is no predation, human interference, or interspecific competition for food), we present a detailed analysis of structured population dynamics with focus on interacting effects of intraspecific competition and weather on reproduction and survival. Annual survival of adult females (0.866 ± 0.107 [ ± SE]) was lower than that of 3‐year‐olds (0.955 ± 0.051), although annual fecundity (producing a foal in a year that was observed during our census) was higher in adults (0.616 ± 0.023) compared to 3‐year‐olds (0.402 ± 0.054). Milder winters and lower densities during gestation increased fecundity. Density negatively impacted survival for all age and sex categories; however, highest adult female survival was observed during high‐density years coupled with a harsh winter, the result expected if pregnancy loss during winter or loss of foals in spring improved survival. Three‐year‐old females, which reproduced at lower rates, experienced higher survival than adults. Our results contrast with a previous study of feral horses that suggested recently feral ungulates might be artificially selected to reproduce even when costs to survival are high. In part, this may be because of the comparably long history of feralization (250 years; at least 25 generations) for Sable Island horses.