Effects of fluctuating thermal regimes on cold survival and life history traits of the spotted wing Drosophila(Drosophila suzukii)

Drosophila suzukii is an invasive pest causing severe damages to a large panel of cultivated crops.To facili tate its biocontrol with stratcgies such as sterile or incompatible insect techniques,D.suzukid must be mass-produced and then stored and transported under low temperature.Prolonged cold exposure induces chill injuries that can be mitigated if the cold period is interrupted with short warming intervals,referred to as fluctuating thermal regimes(FTR).In this study,we tested how to optimally use FTR to extend the shelf life of D.suzukii under cold storage.Several FTR parameters were asessed:temperature(15,20,25℃),duration(0.5,1,2,3 h),and frequency(every 12,24,36,48 h)of warming intervals,in two wild-type lines and in two developmental stages(pupac and adults).Generally,FTR improved cold storage tolerance with respect to constant low temperatures(CLT).Cold mortality was lower when recovery temperature was 20℃ or higher,when duration was 2 h per day or longer,and when warming interruptions occurred frequently(every 12 or 24 h).Applying an optimized FTR protocol to adults greatly reduced cold mortality over long-term storage(up to 130 d).Consequences of FTR on fitness-related traits were also investigated.For adults,poststorage survival was unaffected by FTR,as was the case for female fecundity and male mating capacity.On the other hand,when cold storage occurred at pupal stage,postorage survival and male mating capacity were altered under CLT,but not under FTR.After storage of pupae,female fecundity was lower under FTR compared to CLT,suggesting an energy trade-off between repair of chill damages and C22 production.This study provides detailed information on the application and optimization of an FTR-based protocol for cold storage of D.suzuki that could be useful for the biocontrol of this pest.     

Seasonal variation in life history traits in two Drosophila species

Seasonal environmental heterogeneity is cyclic, persistent and geographically widespread. In species that reproduce multiple times annually, environmental changes across seasonal time may create different selection regimes that may shape the population ecology and life history adaptation in these species. Here, we investigate how two closely related species of Drosophila in a temperate orchard respond to environmental changes across seasonal time. Natural populations of Drosophila melanogaster and Drosophila simulans were sampled at four timepoints from June through November to assess seasonal change in fundamental aspects of population dynamics as well as life history traits. D. melanogaster exhibit pronounced change across seasonal time: early in the season, the population is inferred to be uniformly young and potentially represents the early generation following overwintering survivorship. D. melanogaster isofemale lines derived from the early population and reared in a common garden are characterized by high tolerance to a variety of stressors as well as a fast rate of development in the laboratory environment that declines across seasonal time. In contrast, wild D. simulans populations were inferred to be consistently heterogeneous in age distribution across seasonal collections; only starvation tolerance changed predictably over seasonal time in a parallel manner as in D. melanogaster. These results suggest fundamental differences in population and evolutionary dynamics between these two taxa associated with seasonal heterogeneity in environmental parameters and associated selection pressures.     

Susceptibility of cold hardy grapes to Drosophila suzukii (Diptera: Drosophilidae)

Drosophila suzukii (Diptera: Drosophilidae), known commonly as spotted wing drosophila, is a vinegar fly originating from South‐East Asia and a major pest to many soft‐skinned fruits. Due to the species recent arrival in North America in 2008, many fruit varieties are yet untested for susceptibility to infestation. While previous work has focused on Vitis vinifera, this study aimed to determine grape susceptibility of cold hardy varieties based on hybrids of V. labrusca, V. riparia and V. vinifera. Field sampling was conducted in Southern Wisconsin (USA) vineyards to establish adult and larval abundance and determine whether the number of adults caught in traps correlates with fruit infestation. Host susceptibility was further assessed through no‐choice bioassays of both intact and damaged fruits. The field study found D. suzukii adults present in all varieties, low larval abundance and no correlation between adult abundance and larval presence. Peak adult abundance occurred mid‐season between veraison and harvest, while larval infestation rates were highest near harvest. In laboratory no‐choice tests, significantly more eggs, larvae and adults occurred in damaged than undamaged grapes. In damaged grapes, larvae and adult abundance was comparable between varieties and to the highly susceptible control of undamaged raspberry; however, D. suzukii developed significantly faster in raspberry than grapes. Fruit characteristics (°Brix, titratable acidity, pH) in grapes were uncorrelated with D. suzukii performance. Together, these findings suggest that cold hardy grapes are overall resistant to D. suzukii if intact and highly susceptible if damaged.     

Evaluating Drosophila suzukii immunomarking for mark‐capture research

Drosophila suzukii Matsumura (Diptera: Drosophilidae) utilizes ‘Himalaya’ blackberry, Rubus armeniacus Focke (Rosaceae), as a host and may invade berry and stone fruit crops from field margins containing this invasive weed. Laboratory and semi‐field studies were conducted to determine (1) the persistence of protein marks including 10% chicken egg whites (egg albumin protein), 20% bovine milk (milk casein protein), and 20% soy milk (soy trypsin inhibitor protein) on topically sprayed D. suzukii, (2) protein retention on blackberry leaves, and (3) D. suzukii acquisition of protein after exposure to marked blackberry leaves for up to 14 days after application. All flies and leaves were assayed for the presence of the protein marks using protein‐specific enzyme‐linked immunosorbent assays. Egg albumin, milk casein, and soy trypsin proteins persisted on 94, 49, and 25% of the topically marked D. suzukii, respectively, throughout the 14‐day study period. Egg albumin was retained on 100% of treated leaves for 14 days, regardless of environmental conditions. At least 50% of flies exposed residually to egg albumin‐treated leaves were marked for 3 days, regardless of exposure time and environmental conditions. However, increasing fly exposure time to treated leaves in April and June appeared to improve protein mark acquisition. Acquisition of protein by flies from treated leaves for milk casein was inconsistent, and poor for soy trypsin, despite detectable levels on treated leaves. Egg albumin had the longest and most consistent persistence on flies, leaves, and flies exposed to leaves in laboratory and semi‐field studies, under a variety of environmental conditions and exposure times.     

Nutritional geometry and fitness consequences in Drosophila suzukii,the Spotted‐Wing Drosophila

Since its arrival to North America less than a decade ago, the invasive Spotted‐Wing Drosophila (Drosophila suzukii) has inflicted substantial economic losses on soft fruit agriculture due to its ability to oviposit into ripening fruits. More effective management approaches for this species are needed, but little is known about the factors that influence behavioral choices made by D. suzukii when selecting hosts, or the consequences that their offspring experience when developing in different environments. Using a nutritional geometry methodology, we found that the ratio of proteins‐to‐carbohydrates (P:C) present in media greatly influenced adult D. suzukii behavior and subsequent offspring development. Whereas adult flies showed a strong bias in their oviposition and association behaviors toward carbohydrate‐rich foods, larval survival and eclosion rate were strongly dependent on protein availability. Here, we explore the preference–performance hypothesis (PPH), in which females are predicted to oviposit on medias that provide the greatest offspring benefits, in regard to its relevance in D. suzukii behavior and consequences for management. Our results provide valuable insight into the ecology and evolution of this species that may hopefully lead to more effective management strategies.     

Oviposition and development potential of the spotted‐wing drosophila,Drosophila suzukii (Diptera: Drosophilidae), on uninjured Campbell Early grape

The spotted‐wing drosophila (SWD), Drosophila suzukii (Diptera: Drosophilidae), originally distributed across a few Asian countries including South Korea, has invaded North America and Europe but is absent from Australia. In order to export the South Korean grape cultivar Campbell Early to Australia, its potential to serve as an oviposition and development medium for SWD must first be determined. In this study, we determined the oviposition and development potential of SWD on Campbell Early, after elucidating the SWD life cycle and establishing an artificial diet‐based mass‐culturing system. An investigation of the life cycle under five temperature regimes (16, 19, 22, 25 and 28°C) showed that the durations of the egg, larval and adult stages were shortened when temperature was increased from 16, 19, 22, 25 and 28°C, but pupal duration was shortest at 25°C and extended again at 28°C. A test of oviposition and development potential of SWD on Campbell Early grape clusters showed oviposition of 30.8 ± 6.8 eggs per cluster of injured grapes and 157.7 ± 16.2 eggs on a culture dish of artificial diet. However, in a similar experiment using uninjured grape clusters, only a single egg was deposited on the grape skin, which soon dried. In light of these results, newly harvested grapes left at vineyards during daily harvests are unlikely to serve as an oviposition and development medium for SWD, as long as the grapes remain uninjured.     

Variation of within‐crop microhabitat use by Drosophila suzukii (Diptera: Drosophilidae) in blackberry

Drosophila suzukii Matsumara (Diptera: Drosophilidae) is an invasive vinegar fly that infests ripe and ripening soft skinned fruits. In the south‐eastern United States, blackberry (Rubus spp.) crops are heavily impacted by D. suzukii, and current management tactics rely on the use of broad‐spectrum insecticides targeted to adult populations. An improved understanding of D. suzukii biology and ecology are necessary to create sustainable management options. Knowledge of how D. suzukii interacts with resources will enable targeted management actions in the future. In this present study, we monitored larval infestation throughout the blackberry canopy and found that infestation was highest in the inner portion of the canopy and lower in more exposed locations. We also documented higher humidity within the cane canopy relative to the edge of the field. A difference in abiotic conditions may create within‐crop microhabitats that D. suzukii is able to exploit. Future research will explore how to take advantage of these microhabitats in pest management programs.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Effect of temperature on reproduction,development, and phenotypic plasticity of Drosophila suzukii in Brazil

Temperature is a determining factor for the development and establishment potential of insect pests. The present study describes the impact of temperature (13, 18, 23, 25, 28, 30, and 33 °C) on the life cycle parameters and phenotypic plasticity of South American populations of Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) in the laboratory. Secondary objectives were to determine the lower thermal threshold and thermal constant to estimate the number of annual generations of the insect in small-fruit-producing regions in Brazil. The highest egg-to-adult survival was recorded at 23 and 25 °C. At 30 and 33 °C, no emergence of D. suzukii was observed. The egg-to-adult development time was shortest at 25 and 28 °C (ca. 10 days). The net reproductive rate (R₀), and the intrinsic rate of population increase (r_m) were highest at 23 and 25 °C. In contrast, temperatures of 13 and 28 °C generated largest and smallest body sizes, respectively, and caused reductions of 99 and 93% in R₀. The estimated lower thermal threshold was 7.8 °C for egg-to-adult survival. The estimated thermal constant was 185.8 degree days, and the estimated annual number of generations of D. suzukii ranged from 17.1 in cold regions to 27.2 in warm regions. The results of the present study are important for understanding D. suzukii occurrence in the field, contributing to more informed and precise pest management.     

The effect of border sprays and between‐row soil tillage on Drosophila suzukii in organic blackberry production

Drosophila suzukii (Matsumura) is a major pest of soft‐skinned fruit. Females have an enlarged serrated ovipositor that is used to cut into ripening fruit and lay their eggs. Larvae develop inside infested fruit, rendering fruit unmarketable. Previous research has indicated that D. suzukii can move from adjacent woodlands into cultivated fields. Furthermore, multiple generations can occur in a single season as a result of fallen, infested fruit in the fields. Our hypothesis was that border sprays and soil tillage of field aisles can reduce D. suzukii presence in commercial blackberry fields (Rubus spp.). To test our hypothesis, we conducted split‐plot field trials in organic blackberry fields for 3 and 4 weeks in 2014 and 2015, respectively. Treatments were border sprays (whole plot, pyrethrins + azadirachtin) and tillage (subplot, ~15 cm). We evaluated adult D. suzukii in both years and berry infestation and natural enemies in 2015 only. We found that plots with border treatments had fewer D. suzukii (larvae and adults) than plots without border sprays. Tilling the soil between rows of blackberry bushes did not have a significant effect on adult captures or larval infestation of fruit. Natural enemies were unaffected by the border spray and tillage treatments. Our results confirmed our hypothesis that border sprays can be utilized to reduce populations of D. suzukii in organic blackberry fields, while maintaining populations of natural enemies. However, the effect of soil tillage is unclear and requires further investigation. Additional research should investigate the timing of border sprays and their effect on high infestations of D. suzukii as well as quantify fruit fall and depth of burial to reduce D. suzukii emergence using soil tillage.     

Selection for life‐history traits to maximize population growth in an invasive marine species

Species establishing outside their natural range, negatively impacting local ecosystems, are of increasing global concern. They often display life‐history features characteristic for r‐selected populations with fast growth and high reproduction rates to achieve positive population growth rates (r) in invaded habitats. Here, we demonstrate substantially earlier maturation at a 2 orders of magnitude lower body mass at first reproduction in invasive compared to native populations of the comb jelly Mnemiopsis leidyi. Empirical results are corroborated by a theoretical model for competing life‐history traits that predicts maturation at the smallest possible size to optimize r, while individual lifetime reproductive success (R₀), optimized in native populations, is near constant over a large range of intermediate maturation sizes. We suggest that high variability in reproductive tactics in native populations is an underappreciated determinant of invasiveness, acting as substrate upon which selection can act during the invasion process.     

Trade‐offs between life‐history traits at range‐edge and central locations

The allocation of resources to different life‐history traits should represent the best compromise in fitness investment for organisms in their local environment. When resources are limiting, the investment in a specific trait must carry a cost that is expressed in trade‐offs with other traits. In this study, the relative investment in the fitness‐related traits, growth, reproduction and defence were compared at central and range‐edge locations, using the seaweed Ascophyllum nodosum as a model system. Individual growth rates were similar at both sites, whereas edge populations showed a higher relative investment in reproduction (demonstrated by a higher reproductive allocation and extended reproductive periods) when compared to central populations that invested more in defence. These results show the capability of A. nodosum to differentially allocate resources for different traits under different habitat conditions, suggesting that reproduction and defence have different fitness values under the specific living conditions experienced at edge and central locations. However, ongoing climate change may threaten edge populations by increasing the selective pressure on specific traits, forcing these populations to lower the investment in other traits that are also potentially important for population fitness.     

Life history evolution and cellular mechanisms associated with increased size in high‐altitude Drosophila

Understanding the physiological and genetic basis of growth and body size variation has wide‐ranging implications, from cancer and metabolic disease to the genetics of complex traits. We examined the evolution of body and wing size in high‐altitude Drosophila melanogaster from Ethiopia, flies with larger size than any previously known population. Specifically, we sought to identify life history characteristics and cellular mechanisms that may have facilitated size evolution. We found that the large‐bodied Ethiopian flies laid significantly fewer but larger eggs relative to lowland, smaller‐bodied Zambian flies. The highland flies were found to achieve larger size in a similar developmental period, potentially aided by a reproductive strategy favoring greater provisioning of fewer offspring. At the cellular level, cell proliferation was a strong contributor to wing size evolution, but both thorax and wing size increases involved important changes in cell size. Nuclear size measurements were consistent with elevated somatic ploidy as an important mechanism of body size evolution. We discuss the significance of these results for the genetic basis of evolutionary changes in body and wing size in Ethiopian D. melanogaster.     

Life‐history trade‐offs under different larval diets in Drosophila suzukii (Diptera: Drosophilidae)

Most larval drosophilids eat the microorganisms that develop in rotting fruit, a relatively protein‐rich resource. By contrast, the spotted‐wing Drosophila suzukii Matsumara (Diptera: Drosophilidae) uniquely develops in ripening fruit, a protein‐poor, carbohydrate‐rich resource. This shift in larval nutritional niche has led to D. suzukii being a significant agricultural pest in the U.S.A. and Europe. Although occupying a new niche may benefit a species by reducing competition, adaptation in host use may generate trade‐offs affecting fitness. To test the hypothesis that fitness trade‐offs will change with adaptation to novel larval diets, D. suzukii larval development on either a diet of a fresh, ripe blueberry (a natural host) or standard artificial Drosophila media (protein‐rich) is compared and the effect of diet on development time from egg to adult, adult body size and male wing spot area, and female fecundity is assessed. Larval development time differs, with larvae on the blueberry emerging as adults earlier than those on the artificial medium, although other fitness measures do not vary between the two diets. In addition, the faster development time on a blueberry does not trade off with body size as expected, although early fecundity is delayed in females that develop on blueberries. Thus, adaptation to a novel larval diet environment does not come at a cost to the ability to develop in protein‐rich resources.     

Evaluating the life‐history trade‐off between dispersal capability and reproduction in wing dimorphic insects: a meta‐analysis

A life‐history trade‐off exists between flight capability and reproduction in many wing dimorphic insects: a long‐winged morph is flight‐capable at the expense of reproduction, while a short‐winged morph cannot fly, is less mobile, but has greater reproductive output. Using meta‐analyses, I investigated specific questions regarding this trade‐off. The trade‐off in females was expressed primarily as a later onset of egg production and lower fecundity in long‐winged females relative to short‐winged females. Although considerably less work has been done with males, the trade‐off exists for males among traits primarily related to mate acquisition. The trade‐off can potentially be mitigated in males, as long‐winged individuals possess an advantage in traits that can offset the costs of flight capability such as a shorter development time. The strength and direction of trends differed significantly among insect orders, and there was a relationship between the strength and direction of trends with the relative flight capabilities between the morphs. I discuss how the trade‐off might be both under‐ and overestimated in the literature, especially in light of work that has examined two relevant aspects of wing dimorphic species: (1) the effect of flight‐muscle histolysis on reproductive investment; and (2) the performance of actual flight by flight‐capable individuals.     

Evolution of life‐history traits and mating strategy in males: a case study on two populations of a Drosophila parasitoid

Abiotic and biotic factors affect life‐history traits and lead populations to exhibit different behavioural strategies. Due to the direct link between their behaviour and fitness, parasitoid females have often been used to test the theories explaining these differences. In male parasitoids, however, such investigations are vastly understudied, although their mating strategy directly determines their fitness. In this study, we compared the pattern of life history traits and the mating strategy of males in two populations of the Drosophila parasitoid Asobara tabida, exposed to different biotic and abiotic conditions, with the major difference being that one of them was recently exposed to strong competition with the dominant competitor Leptopilina boulardi after recent climate change, the other population being settled in a location where L. boulardi has not been recorded. The results showed that individuals of both populations have a different reproductive strategy: in one population, females produced a more female‐biased sex ratio, while males accumulated more lipids during their larval development, invested more energy in reproduction and decreased their locomotor activity, suggesting a higher proportion of matings on their emergence patch, all of these factors being possibly linked to the new competition pressure. In both populations, mating without sperm transfer may persist for several days after males become sperm‐depleted, and may be more frequent than mating with sperm transfer over their whole lifespan. This point is discussed from an evolutionary point of view.     

Interplay of robustness and plasticity of life‐history traits drives ecotypic differentiation in thermally distinct habitats

Phenotypic plasticity describes the ability of an individual to alter its phenotype in response to the environment and is potentially adaptive when dealing with environmental variation. However, robustness in the face of a changing environment may often be beneficial for traits that are tightly linked to fitness. We hypothesized that robustness of some traits may depend on specific patterns of plasticity within and among other traits. We used a reaction norm approach to study robustness and phenotypic plasticity of three life‐history traits of the collembolan Orchesella cincta in environments with different thermal regimes. We measured adult mass, age at maturity and growth rate of males and females from heath and forest habitats at two temperatures (12 and 22 °C). We found evidence for ecotype‐specific robustness of female adult mass to temperature, with a higher level of robustness in the heath ecotype. This robustness is facilitated by plastic adjustments of growth rate and age at maturity. Furthermore, female fecundity is strongly influenced by female adult mass, explaining the importance of realizing a high mass across temperatures for females. These findings indicate that different predicted outcomes of life‐history theory can be combined within one species' ontogeny and that models describing life‐history strategies should not assume that traits like growth rate are maximized under all conditions. On a methodological note, we report a systematic inflation of variation when standard deviations and correlation coefficients are calculated from family means as opposed to individual data within a family structure.     

Tracking changes in life‐history traits related to unnecessary virulence in a plant‐parasitic nematode

Evaluating trade‐offs in life‐history traits of plant pathogens is essential to understand the evolution and epidemiology of diseases. In particular, virulence costs when the corresponding host resistance gene is lacking play a major role in the adaptive biology of pathogens and contribute to the maintenance of their genetic diversity. Here, we investigated whether life‐history traits directly linked to the establishment of plant–nematode interactions, that is, ability to locate and move toward the roots of the host plant, and to invade roots and develop into mature females, are affected in Meloidogyne incognita lines virulent against the tomato Mi‐1.2 resistance gene. Virulent and avirulent near‐isogenic lines only differing in their capacity to reproduce or not on resistant tomatoes were compared in single inoculation or pairwise competition experiments. Data highlighted (1) a global lack of trade‐off in traits associated with unnecessary virulence with respect to the nematode ability to successfully infest plant roots and (2) variability in these traits when the genetic background of the nematode is considered irrespective of its (a)virulence status. These data suggest that the variation detected here is independent from the adaptation of M. incognita to host resistance, but rather reflects some genetic polymorphism in this asexual organism.     

Distribution and activity of Drosophila suzukii in cultivated raspberry and surrounding vegetation

Spotted wing drosophila, Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), may utilize wild ‘Himalaya’ blackberry (HB) Rubus armeniacus Focke or other non‐crop plants as refugia and possibly exploit adjacent field margins before colonizing cultivated fruiting crops. Studies were conducted to determine the role of field margins containing HB and their effect on D. suzukii activity, density and distribution in an adjacent commercial red raspberry crop. One‐ha plots adjacent to field margins containing HB or known non‐host (NH) grass crops were established in 2011 and 2012 and replicated three times. Each plot contained two transects with monitoring traps for D. suzukii in the field margin (0 m) and spaced approximately 10 (crop boundary), 40, 70 and 100 m into the adjacent crop (n = 10 traps/plot). Field margin vegetation was treated with a 10% chicken egg white mark solution weekly from pre‐harvest until the end of harvest using a cannon sprayer. Adult D. suzukii were collected from traps weekly and analysed for the presence of the egg white mark using an egg white‐specific enzyme‐linked immunosorbent assay (ELISA). During both years, marked flies and total flies were captured in higher numbers in HB field margins, whereas virtually no flies were captured in field margins containing no known alternative host. Similarly, more flies were captured in the crop near HB than near NH. Spatial Analysis by Distance IndicEs (SADIE) and mean D. suzukii trap captures additionally displayed significantly higher fly densities in the raspberry field near HB than near NH. These results suggest that HB may contribute to elevated D. suzukii populations and pest pressure in comparison with field margins containing no known alternate host vegetation for D. suzukii. Having closely adjacent non‐crop alternate host landscapes may result in increased D. suzukii pest pressure.     

Entanglement is a costly life‐history stage in large whales

Individuals store energy to balance deficits in natural cycles; however, unnatural events can also lead to unbalanced energy budgets. Entanglement in fishing gear is one example of an unnatural but relatively common circumstance that imposes energetic demands of a similar order of magnitude and duration of life‐history events such as migration and pregnancy in large whales. We present two complementary bioenergetic approaches to estimate the energy associated with entanglement in North Atlantic right whales, and compare these estimates to the natural energetic life history of individual whales. Differences in measured blubber thicknesses and estimated blubber volumes between normal and entangled, emaciated whales indicate between 7.4 × 10¹⁰ J and 1.2 × 10¹¹ J of energy are consumed during the course to death of a lethal entanglement. Increased thrust power requirements to overcome drag forces suggest that when entangled, whales require 3.95 × 10⁹ to 4.08 × 10¹⁰ J more energy to swim. Individuals who died from their entanglements performed significantly more work (energy expenditure × time) than those that survived; entanglement duration is therefore critical in determining whales’ survival. Significant sublethal energetic impacts also occur, especially in reproductive females. Drag from fishing gear contributes up to 8% of the 4‐year female reproductive energy budget, delaying time of energetic equilibrium (to restore energy lost by a particular entanglement) for reproduction by months to years. In certain populations, chronic entanglement in fishing gear can be viewed as a costly unnatural life‐history stage, rather than a rare or short‐term incident.