Poaching is a widespread and well-appreciated problem for the conservation of many threatened species. Because poaching is illegal, there is strong incentive for poachers to conceal their activities, and consequently, little data on the effects of poaching on population dynamics are available. Quantifying poaching mortality should be a required knowledge when developing conservation plans for endangered species but is hampered by methodological challenges. We show that rigorous estimates of the effects of poaching relative to other sources of mortality can be obtained with a hierarchical state–space model combined with multiple sources of data. Using the Scandinavian wolf (Canis lupus) population as an illustrative example, we show that poaching accounted for approximately half of total mortality and more than two-thirds of total poaching remained undetected by conventional methods, a source of mortality we term as ‘cryptic poaching’. Our simulations suggest that without poaching during the past decade, the population would have been almost four times as large in 2009. Such a severe impact of poaching on population recovery may be widespread among large carnivores. We believe that conservation strategies for large carnivores considering only observed data may not be adequate and should be revised by including and quantifying cryptic poaching.

Predatory behavior of wolves (Canis lupus) was studied in 2 wolf territories in Scandinavia. We used hourly data from Global Positioning System (GPS)‐collared adult wolves in combination with Geographic Information System (GIS) for detailed analyses of movement patterns. We tested the hypothesis that wolves spend 1–2 days close to larger prey such as moose (Alces alces) and reasoned that 1–2 locations per day would be enough to find all larger prey killed by the wolves. In total, the study period comprised 287 days and yielded 6,140 hourly GPS positions, with an average of 21.4±2.4 (SD) daily positions. Depending on the radius used to define clusters, 4,045‐5,023 (65.9–81.8%) positions were included in 622–741 GPS‐clusters. We investigated all positions within clusters in the field, and 244 (22%) single positions. In total, we found 68 moose and 4 roe deer (Capreolus capreolus) and classified them as wolf‐killed within the study period. Another 10–15 moose may have been killed but not found. The GIS analyses indicated the proportion of wolf‐killed ungulates included in GPS clusters to be strongly dependent on both number of positions per day and the radius used for defining a set of spatially aggregated GPS positions as a cluster. A higher proportion (78%) of killed prey in clusters based on nighttime (2000‐0700) than those based on daytime (0800–1900) positions (41%). Simulation of aerial search during daylight hours for killed moose resulted in a serious underestimation (>60%) as compared to the number of wolf‐killed moose found during the study. The average kill rate, corrected for 14% nondetected moose, in the territories was 3.6‐4.0 days per killed moose. We concluded that the feeding behavior of wolves in Scandinavia was either different from wolves preying on moose and living at the same latitude in North America, or that estimates of wolf kill rates on moose may have been seriously underestimated in previous North American studies.

Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult‐sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r= —0.37) and time to next reproduction (r= —0.36). Time to breeder replacement correlated strongly with time to next reproduction (r=0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad‐sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.

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