Effect of supplementary feeding of vitamin K1 on difethialone treated Indian gerbil, Tatera indica Hardwicke in laboratory

Two dosages (1 and 2 mg/kg) of vitamin K1 supplementation for 5 and 15 days were given to Indian gerbil T. indica fed on difethialone bait (0.0025%) for one day. The results indicated that the lower dosage could not reverse the anticoagulation process, however the period of mortality was considerably increased from 3-9 days (in control) to 5-14 days (5 days supplementation regime). Subsequently when the vitamin K1 dosage was doubled and given for 15 days, there was 100% reversal of anticoagulation process and all the test gerbils became normal within a month of poisoning with difethialone bait.     

Environmental,parental and adaptive variation in egg size of Tengmalm's owls under fluctuating food conditions

We studied egg size variation of Tengmalm's owls in western Finland during 1981–1990. The owls fed on voles whose population fluctuated in a predictable manner: low (1981, 1984, 1987, 1990), increase (1982, 1985, 1988) and peak (1983, 1986, 1986) phases of the cycle occurred every third year. Eggs were largest in the increase phase of the vole cycle, even though that voles were more abundant and egg-laying started earlier in the peak phase than in the increase phase. This suggests that owls invest mostly in egg size when vole abundance increases along with survival chances of offspring. Territory quality and female age had no effects on egg size, but egg size decreased with laying data in the increase phase of the vole cycle. Egg size was significantly positively related to the male age in the increase phase, but the opposite relationship was significant in the peak phase of the vole cycle. The partners of adult males also decreased their egg volume from the increase to the peak phase, whereas the partners of yearling males produced their largest eggs in the peak phase of the vole cycle. This suggests the importance of experience in prevailing food fluctuations. Possibly male Tengmalm's owls can adjust the intensity of courtship feeding not only in relation to the food abundance on their territories at the time of egg laying, but also to the survival prospects of their offspring. Phenotypic plasticity seems to play a substantial role, as the egg size repeatabilities of individual females and partners of individual males were low. Obviously, under cyclic food conditions, predictability and inter-generational trade-offs are important to life history traits.     

Pygmy owl Glaucidium passerinum and the usage of ultraviolet cues of prey

Birds rely mainly on their vision when foraging. Many diurnal raptors use ultraviolet (UV) vision and ultraviolet‐reflecting vole scent marks to find suitable hunting areas, whereas nocturnal owls seem to lack this ability. We studied if the diurnal pygmy owl Glaucidium passerinum that uses voles and birds as its food can detect vole scent marks using UV‐vision. We conducted a laboratory experiment with eleven owls. Each individual owl had four options to choose from: (1) scent marks with UV light, (2) scent marks without UV light, (3) clean arena with UV light and (4) clean arena without UV light. The owls scanned the scent mark arena more often in the presence of UV light than other arenas. However, owls did not spend more time above the UV arena. We suggest that pygmy owls can detect near UV and use UV to gain information about prey like other diurnal raptors.     

Blood characteristics, tracheal volume and heart mass of burrowing owls (Athene cunicularia) and bobwhite (Colinus virginianus)

1. Measurements of certain hematological and morphological characteristics were made in burrowing owls and bobwhite in search of features that could be associated with the previously described ventilatory adaptations of the burrowing owl to hypoxia and hypercarbia. 2. Values for burrowing owls and bobwhite, respectively, were: P50 = 44.9, 46.0 torr; Hct = 36.6, 38.8 vol%, Hb = 12, 12.3 (g/100 ml); RBC = 2.72 X 10(6), 3.02 X 10(6); log P50/pH = -0.412, -0.485; delta log PCO2/delta pH = -1.39, -1.585. 3. The owls had greater heart weights and smaller tracheal volumes than the bobwhite or the predicted value. 4. No hematological characteristics of the burrowing bird distinguish it from the non-burrowing bird.     

Effects of Radiotransmitter Necklaces on Behaviors of Adult Male Western Burrowing Owls

Abstract: We studied the behavioral effects of necklace-style radiotransmitters on breeding male western burrowing owls (Athene cunicularia hypugaea) in 2 areas of northwestern Texas, USA, in 2004 and 2005. We tested the hypothesis that transmittered owls would spend time interacting with their necklaces and as a result spend less time in vigilance and resting activities than would nontransmittered owls. Nontransmittered owls (n = 6) spent significantly more time being vigilant (P = 0.007) than did transmittered owls (n = 3) in 2004, who spent significant amounts of time interacting with their necklaces. In 2005, behaviors of transmittered owls (n = 8) were significantly different (P < 0.001) from control individuals (n = 4), but behaviors did not vary consistently by treatment period (prenecklace vs. necklace vs. postnecklace periods). Behavioral activity budgets varied considerably among individuals. Although the owls spent a significant amount of time interacting with their necklaces, they appeared to habituate to the presence of the transmitters within a relatively short period (<1week), and necklaces did not affect survivorship or fitness in the short-term.     

Postnatal development of the mixed function oxidase system in nestling barn owls and baby chicks and induction of this system in the mature forms by Aroclor 1254

Postnatal changes in content and activity of the mixed-function oxidase system in nestling barn owls and baby chicks showed the following: 1. Increase in liver weight in both. 2. Significantly higher cytochrome P-450 level in 1-day old chicks but lower in 1-day old owls than at any other age. 3. Ratio of cytochrome b5 to P-450 was lower than one in nestling owls but higher than one in chicks. 4. Aroclor 1254 (PCBs) increased the level and catalytic activity of cytochrome P-450 more in the owl than in the chick. 5. The ratio 455:430 nm characteristic of ethylisocyanide binding was not altered in the barn owl due to PCB treatment but changed significantly in the treated chicken.     

Site occupancy dynamics of northern spotted owls in managed interior Douglas fir forests,California, USA, 1995–2009

Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. For example, public and private forest managers in the Pacific Northwest, USA, conduct surveys to determine presence or absence of spotted owls prior to timber harvest operations. However, although recently developed statistical methods have been applied to presence–absence data collected during research surveys, the effectiveness of operational surveys for detecting spotted owls and evaluating site occupancy dynamics is not known. We used spotted owl survey data collected from 1995 to 2009 on a study area in interior northern California, USA, to evaluate competing occupancy models from Program PRESENCE using Akaike's Information Criterion (AIC). During 1,282 individual surveys, we recorded 480 spotted owl detections (37.4%) and 13 barred owl (1.0%) detections. Average per visit detection probability (85% CL) for single and paired spotted owls was 0.93 (0.90–0.96) for informed daytime, stand-based searches and 0.47 (0.43–0.51) for nighttime, station-based surveys (estimated from the best model); the average per visit detection probability from the null model was 0.67 (0.64–0.70). Average pair-only detection probabilities were 0.86 (0.81–0.90) for informed daytime, stand-based searches and 0.23 (0.18–0.29) for nighttime, station-based surveys; the average per visit detection probability from the null model was 0.63 (0.58–0.68). Site occupancy for any owl declined from 0.81 (0.59–0.93) in 1995 to 0.50 (0.39–0.60) in 2009; pair occupancy declined from 0.75 (0.56–0.87) to 0.46 (0.31–0.61). Our results suggest that a combination of 1 informed stand and 2 station-based operational surveys can support determinations of spotted owl site status (either a single or a pair) at desired levels of confidence. However, our information was collected in an area where barred owls were rarely detected. Surveys conducted in areas that support well-established barred owl populations are likely to be less effective for determining presence or absence of spotted owls and may require more surveys and/or different survey methods to determine site status with confidence. © 2012 The Wildlife Society.     

Roadway mortality of barn owls in Idaho,USA

We examined the temporal, spatial, and demographic factors that influenced roadway mortality of barn owls (Tyto alba) along a 248-km stretch of Interstate 84 in southern Idaho using systematic road surveys. Counts of dead animals from surveys can be underestimated because of sampling biases; therefore, we also conducted experiments to assess the effects of search and removal bias on the estimates of roadway mortality of owls. We conducted surveys every 2 weeks over a 2-year period and detected 812 dead barn owls (unadjusted mortality rate of 1.64 owls/km/yr). After adjusting this estimate for search and removal bias, we documented mortality rates of up to 5.99 owls/km/year. Owl mortality was not random in relation to sex, age class, or location along the highway. Females and juveniles, which represent individuals more likely to disperse long distances, were killed more frequently than males and adults. During the nonbreeding season, owls were killed more often near agricultural lands than in shrub-steppe, but this pattern was not apparent during the breeding season. Owls were also killed more often on portions of the roadway closer to the Snake River canyon, perhaps because of the availability of nest and roost sites. Mortality rates differed markedly between the 2 years of study, which could have been related to variability in weather and its subsequent effect on owl productivity. Our data suggest that barn owls in this region may not persist under this level of mortality without significant immigration or management. Thus, roadway management to reduce or prevent owl use of roadways, reduce rodent populations near major roads, alert motorists to the presence of owls, or otherwise reduce the chances that vehicles and owls collide would improve barn owl survival and population persistence. © 2012 The Wildlife Society.     

Complete mtDNA genome of Otus sunia (Aves,Strigidae) and the relaxation of selective constrains on Strigiformes mtDNA following evolution

Most of owls are nocturnal raptor and usually use their soft and fluffy feathers to flight silently to catch prey while other diurnal raptors prefer fierce attack and swift flight. For energy cost of these different hunting strategies can be greatly different, we speculate that mitochondrial gene of owls may undergo a different evolution pattern following raptors evolution. To test our hypothesis, we sequenced the mtDNA genome of Otus sunia and calculated the ratio of nonsynonymous to synonymous nucleotide substitutions (ω, Ka/Ks, dN/dS) of raptors. The mtDNA genome of O. sunia was 17,609 bp in length, containing 13 PCGs, 2 ribosomal RNAs, 22 transfer RNAs and a control region. Secondly structure of tRNAs and rRNAs were predicted and conserved sequence blocks (CSBs) on control region were identified. The Bayesian inference tree and maximum likelihood tree based on 13 PCGs and 2 rRNAs suggested the owls were related to other raptors. Finally, calculation of ω-values of each owls and other raptors mtDNA PCGs indicated that owls accumulated more nonsynonymous nucleotide substitutions relative to synonymous substitutions compared to other raptors. For mtDNA PCGs associated with energy metabolism, this finding may reveal the degeneration of flight abilities of owls.     

Ventilatory and metabolic responses of burrowing owls, Athene cunicularia, to moderate and extreme hypoxia: analysis of the hypoxic ventilatory threshold vs. hemoglobin oxygen affinity relationship in birds

We measured ventilation, oxygen consumption and blood gases in burrowing owls (Athene cunicularia) breathing moderate and extreme hypoxic gas mixtures to determine their hypoxic ventilatory threshold (HVT) and to assess if they, like other birds and mammals, exhibit a relationship between HVT and hemoglobin O2 affinity (P(50)) of their blood. An earlier report of an attenuated ventilatory responsiveness of this species to hypoxia was enigmatic given the low O2 affinity (high P(50)) of burrowing owl hemoglobin. In the current study, burrowing owls breathing 11% and 9% O2 showed a significantly elevated total ventilation. The arterial partial pressure of oxygen (PaO2) at which ventilation is elevated above normoxic values in burrowing owls was 58 mm Hg. This threshold value conforms well to expectations based on the high P(50) of their hemoglobin and the HVT vs. P(50) relationship for birds developed in this study. Correcting for phylogenetic relatedness in the multi-species analysis had no effect on the HVT vs. P(50) relationship. Also, because burrowing owls in this study did not show a hypometabolic response at any level of hypoxia (even at 9% O2); HVT described in terms of percent change in oxygen convection requirement is identical to that based on ventilation alone.     

Effects of survey methods on burrowing owl behaviors

Monitoring wildlife populations often involves intensive survey efforts to attain reliable estimates of population size. Such efforts can increase disturbance to animals, alter detection, and bias population estimates. Burrowing owls (Athene cunicularia) are declining across western North America, and information on the relative effects of potential survey methods on owl behaviors is needed. We designed a field experiment to compare burrowing owl flight distances, times displaced, and probabilities of being displaced between 4 potential population survey methods (single walking surveyor, single vehicle stop, single vehicle stop with 2 surveyors, and double vehicle stop with 2 surveyors), and an experimental control in the agricultural matrix of Imperial Valley, California. Between 25 April and 1 May 2008, we randomly applied survey methods to 395 adult male owls during daylight hours (0700 hours through 1900 hours). All survey methods increased odds of displacing owls from perches. Survey methods with observers outside the vehicle were 3 times more likely to displace an owl than a single vehicle stop where observers remained inside the vehicle. Owls were displaced farther distances by all survey methods compared to control trials, but distances and time displaced did not differ among survey methods. We recommend that surveys for counting owls during the breeding season in agroecystems like the Imperial Valley where high densities of owls nest primarily along the borders of fields be conducted using single vehicle stops with or without 2 surveyors, depending on conditions for locating owls from roads. © 2011 The Wildlife Society.     

Energy metabolism and body temperature of barn owls fasting in the cold

Energetic adaptation to fasting in the cold has been investigated in a nocturnal raptor, the barn owl (Tyto alba), during winter. Metabolic rate and body temperature (Tb) were monitored in captive birds, (1) after acute exposure to different ambient temperatures (Ta), and (2) during a prolonged fast in the cold (4 degrees C), to take into account the three characteristic phases of body fuel utilization that occur during a long-term but reversible fast. In postabsorptive birds, metabolic rate in the thermoneutral zone was 4. 1+/-0.1 W kg-1 and increased linearly below a lower critical temperature of 23 degrees C. Metabolic rate was 70% above basal at +4 degrees C Ta. Wet thermal conductance was 0.22 W kg-1 degrees C-1. During fasting in the cold, the mass-specific resting metabolic rate decreased by 16% during the first day (phase I) and remained constant thereafter. The amplitude of the daily rhythm in Tb was only moderately increased during phase II, with a slight lowering (0. 6 degrees C) in minimal diurnal Tb, but rose markedly in phase III with a larger drop (1.4 degrees C) in minimal diurnal Tb. Refeeding the birds ended phase III and reversed the observed changes. These results indicate that diurnal hypothermia may be used in long-term fasting barn owls and could be triggered by a threshold of body lipid depletion, according to the shift from lipid to protein fuel metabolism occurring at the phase II/phase III transition. The high cost of regulatory thermogenesis and the limited use of hypothermia during fasting may contribute to the high mortality of barn owls during winter.     

Ventilatory and metabolic responses of burrowing owls, Athene cunicularia, to moderate and extreme hypoxia: analysis of the hypoxic ventilatory threshold vs. hemoglobin oxygen affinity relationship in birds.

We measured ventilation, oxygen consumption and blood gases in burrowing owls (Athene cunicularia) breathing moderate and extreme hypoxic gas mixtures to determine their hypoxic ventilatory threshold (HVT) and to assess if they, like other birds and mammals, exhibit a relationship between HVT and hemoglobin O2 affinity (P(50)) of their blood. An earlier report of an attenuated ventilatory responsiveness of this species to hypoxia was enigmatic given the low O2 affinity (high P(50)) of burrowing owl hemoglobin. In the current study, burrowing owls breathing 11% and 9% O2 showed a significantly elevated total ventilation. The arterial partial pressure of oxygen (PaO2) at which ventilation is elevated above normoxic values in burrowing owls was 58 mm Hg. This threshold value conforms well to expectations based on the high P(50) of their hemoglobin and the HVT vs. P(50) relationship for birds developed in this study. Correcting for phylogenetic relatedness in the multi-species analysis had no effect on the HVT vs. P(50) relationship. Also, because burrowing owls in this study did not show a hypometabolic response at any level of hypoxia (even at 9% O2); HVT described in terms of percent change in oxygen convection requirement is identical to that based on ventilation alone.     

Site Occupancy Dynamics of Northern Spotted Owls in the Eastern Cascades,Washington, USA, 1990–2003

Abstract: Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. Several spotted owl (Strix occidentalis) response variables have been examined in various investigations, but recent advances in statistical modeling permit evaluations of temporal and spatial variability in site occupancy, local-extinction, and colonization probabilities while incorporating imperfect detection probabilities. Following recent work by other researchers on site occupancy dynamics of spotted owls in Oregon, USA, we evaluated temporal variability of detection, occupancy, local-extinction, and colonization probabilities for spotted owls, as well as potential influences of barred owl (Strix varia) presence on these parameters. We used spotted owl survey data collected from 1990 to 2003 on a study area in the eastern Cascades Mountains, Washington, USA, to compare competing occupancy models from Program PRESENCE using Akaike's Information Criterion. Detection probabilities for individual spotted owls ranged from 0.54 to 0.80 if barred owls were not detected during the survey season and from 0.19 to 0.71 if barred owls were detected during the survey season. Pair detection probabilities ranged from 0.27 to 0.67 if barred owls were not detected during an individual survey and from 0.09 to 0.36 if barred owls were detected during an individual survey. During the study, site occupancy probabilities for spotted owl pairs declined by approximately 50%. For all spotted owls, both singles and pairs, site occupancy probabilities declined moderately during the study. Barred owl presence was negatively associated with spotted owl detection probabilities, and it had a positive association with local-extinction probabilities for all spotted owls, both singles and pairs. Given that our study area has supported higher densities of barred owls for longer periods than other study areas, our results may provide insight into how barred owls have influenced spotted owl site occupancy dynamics in adjacent British Columbia, Canada, or will influence spotted owl site occupancy dynamics in Oregon and California, USA, in the future.     

Avian species differences in the intestinal absorption of xenobiotics (PCB, dieldrin, Hg2+)

Intestinal absorption of a polychlorinated biphenyl, dieldrin, and mercury (from HgCl2) was measured in adult Northern bobwhites, Eastern screech owls, American kestrels, black-crowned night-herons and mallards in vivo by an in situ luminal perfusion technique. bobwhites, screech owls and kestrels absorbed much more of each xenobiotic than black-crowned night-herons and mallards. Mallards absorbed less dieldrin and mercury than black-crowned night-herons. Mercury absorption by kestrels was more than twice that in screech owls and eight times that observed in mallards. Pronounced differences in xenobiotic absorption rates between bobwhites, screech owls and kestrels on the one hand, and black-crowned night-herons and mallards on the other, raise the possibility that absorptive ability may be associated with the phylogenetic classification of birds.     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

Barred Owl Space Use and Habitat Selection in the Eastern Cascades,Washington

ABSTRACT Competition with barred owls (Strix varia varia) is an important factor contributing to the continued decline of threatened northern spotted owl (Strix occidentalis caurina) populations in the Pacific Northwest, USA, but basic information on habitat selection and space use patterns of barred owls is lacking for much of the region. We investigated space use and habitat selection by tracking radiotagged barred owls in the Eastern Cascade Range of Washington, USA, from 2004 to 2006. We surveyed for barred owls across the 309-km² study area and confirmed presence of barred owl pairs at 21 sites. We collected movement data on 14 barred owls from 12 sites. Mean annual 95% fixed-kernel home-range size was 194 ha for females (n = 4, SD = 70) and 288 ha for males (n = 5, SD = 114). Home ranges were located more frequently than expected in areas with low topographic position, gentle slopes, large overstory tree-crown diameter, high normalized difference vegetation index (NDVI), overstory tree canopy closure >72%, and a moderate amount of solar insolation. Within home ranges, areas that had large tree-crown diameters, low topographic positions, and gentle slopes were used more frequently than expected. The resource selection function we developed for barred owls in our study area indicated that barred owls used areas with the combination of low values for topographic position and slope and higher values for NDVI, solar insolation, and an interaction term for canopy closure and tree-crown diameter. In comparison to published information on northern spotted owls, barred owls used areas with similar canopy closure and tree size classes, but barred owl home ranges were much smaller and more concentrated on gentler slopes in valley bottoms. This information may contribute to the development of management practices that maintain forest characteristics appropriate for spotted owl habitat and prey in areas where spotted owls are least likely to be excluded by territorial barred owls in the Eastern Cascades of Washington.     

Using Detection Dogs to Conduct Simultaneous Surveys of Northern Spotted (Strix occidentalis caurina) and Barred Owls (Strix varia)

State and federal actions to conserve northern spotted owl (Strix occidentalis caurina) habitat are largely initiated by establishing habitat occupancy. Northern spotted owl occupancy is typically assessed by eliciting their response to simulated conspecific vocalizations. However, proximity of barred owls (Strix varia)-a significant threat to northern spotted owls-can suppress northern spotted owl responsiveness to vocalization surveys and hence their probability of detection. We developed a survey method to simultaneously detect both species that does not require vocalization. Detection dogs (Canis familiaris) located owl pellets accumulated under roost sites, within search areas selected using habitat association maps. We compared success of detection dog surveys to vocalization surveys slightly modified from the U.S. Fish and Wildlife Service's Draft 2010 Survey Protocol. Seventeen 2 km ×2 km polygons were each surveyed multiple times in an area where northern spotted owls were known to nest prior to 1997 and barred owl density was thought to be low. Mitochondrial DNA was used to confirm species from pellets detected by dogs. Spotted owl and barred owl detection probabilities were significantly higher for dog than vocalization surveys. For spotted owls, this difference increased with number of site visits. Cumulative detection probabilities of northern spotted owls were 29% after session 1, 62% after session 2, and 87% after session 3 for dog surveys, compared to 25% after session 1, increasing to 59% by session 6 for vocalization surveys. Mean detection probability for barred owls was 20.1% for dog surveys and 7.3% for vocal surveys. Results suggest that detection dog surveys can complement vocalization surveys by providing a reliable method for establishing occupancy of both northern spotted and barred owl without requiring owl vocalization. This helps meet objectives of Recovery Actions 24 and 25 of the Revised Recovery Plan for the Northern Spotted Owl.     

Interactions Between Inter-Individual and Inter-Task Differences in the Diurnal Variation of Human Performance

Thirty-six undergraduate subjects were divided into ‘larks’ and ‘owls’ by median split using the morningness test devised by Horne and Ostberg. Each was given a battery of two performance tests at six times of day, in a design that counter-balanced for practice effects. Oral temperatures were also recorded. Slight (1 hr) phase differences (in the predicted direction) were observed between ‘larks’ and ‘owls’ in temperature rhythm, and in the performance test that paralleled temperature (a serial search task). Much more striking differences, though, were observed in the cognitive (logical reasoning) performance test which (as predicted) peaked earlier in the day, on average, than the serial search task. Not only did ‘larks’ peak much earlier in the day than ‘owls’, but there were also quite dramatic differences between the two groups in amplitude and shape. Possible mechanisms regarding oscillatory control and performance strategy are discussed.     

Does feeding effort of Tengmalm's owls reflect offspring survival prospects in cyclic food conditions?

In an environment fluctuating in a predicatable manner with wide among-year variation in offspring mortality, fitness is largely influenced by the timing of reproductive investment. In vole-eating nocturnal Tengmalm's owls (Aegolius funereus), within-cycle variation in 1st-year survival of owlets is 3-fold as estimated by the recruitment probability of an offspring. It increases from the peak through the low to the increase phase of the vole cycle. We recorded prey delivery rates of males during a 3-year vole cycle using 4 h continuous recording at night. Males brought significantly more prey items per offspring in a low-vole year than in the increase and peak vole years. In the early night (10 p.m.–12 p.m.), males fed their families equally in the increase and peak vole years, whereas in the late night (0.01–2.00 a.m.) males reduced their feeding rate in the peak year but not in other years. Both prey number and prey mass per offspring were larger in the low and increase vole years than in the peak year, though in the peak phase food is most abundant. We suggest that feeding effort of site-tenacious, long-lived (mean lifespan 3.5 years) male owls culminates in the increase rather than in the peak phase of the vole cycle, because offspring survive better in the former phase.