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Demography of an apex predator at the edge of its range: impacts of changing sea ice on polar bears in Hudson Bay

Changes in the abundance and distribution of wildlife populations are common consequences of historic and contemporary climate change. Some Arctic marine mammals, such as the polar bear ( Ursus maritimus ), may be particularly vulnerable to such changes due to the loss of Arctic sea ice. We evaluated the impacts of environmental variation on demographic rates for the Western Hudson Bay (WH), polar bear subpopulation from 1984 to 2011 using live-recapture and dead-recovery data in a Bayesian implementation of multistate capture–recapture models. We found that survival of female polar bears was related to the annual timing of sea ice break-up and formation. Using estimated vital rates (e.g., survival and reproduction) in matrix projection models, we calculated the growth rate of the WH subpopulation and projected population responses under different environmental scenarios while accounting for parametric uncertainty, temporal variation, and demographic stochasticity. Our analysis suggested a long-term decline in the number of bears from 1185 (95% Bayesian credible interval [BCI] = 993–1411) in 1987 to 806 (95% BCI = 653–984) in 2011. In the last 10 yr of the study, the number of bears appeared stable due to temporary stability in sea ice conditions (mean population growth rate for the period 2001–2010 = 1.02, 95% BCI = 0.98–1.06). Looking forward, we estimated long-term growth rates for the WH subpopulation of ~1.02 (95% BCI = 1.00–1.05) and 0.97 (95% BCI = 0.92–1.01) under hypothetical high and low sea ice conditions, respectively. Our findings support previous evidence for a demographic linkage between sea ice conditions and polar bear population dynamics. Furthermore, we present a robust framework for sensitivity analysis with respect to continued climate change (e.g., to inform scenario planning) and for evaluating the combined effects of climate change and management actions on the status of wildlife populations.

Hudson Bay

Coupling contaminants with demography: Effects of lead and selenium in Pacific common eiders

We coupled intensive population monitoring with collection of blood samples from 383 nesting Pacific common eiders ( Somateria mollisima v-nigrum ) at two locations in Alaska (USA) from 2002 to 2004. We investigated annual, geographic, and within-season variation in blood concentrations of lead and selenium; compared exposure patterns with sympatrically nesting spectacled eiders ( Somateria fischeri ); and examined relationships with clutch size, egg viability, probability of hatching, and apparent survival of adult females. Lead concentrations were elevated in 3.6% of females, and all individuals exhibited elevated selenium, most (81%) at concentrations associated with death in captive waterfowl. Blood lead and selenium concentrations varied both within and among site-years and were lower than those of spectacled eiders. During incubation, blood lead concentrations in females increased significantly (possibly via re-release of stored lead from bone), whereas selenium concentrations decreased (likely because of natural excretion). Probability of a nest containing at least one nonviable egg was positively related to blood selenium in hens, but adverse effects in other life-history variables were not supported. Although reproduction appeared to be sensitive to selenium toxicity, our data suggest that high rates of nonviability are unlikely in this population and that selenium-related reductions to clutch size would be inconsequential at the scale of overall population dynamics. We conclude that Pacific common eiders and other wild marine birds likely have higher selenium tolerances than freshwater species and that interspecific differences in exposure levels may reflect differences in reproductive strategies.

Environmental Toxicology and Chemistry

Reproduction and demography of the Florida Everglade (Snail) Kite

An 18-year study of reproduction and survival of the Florida Everglade (Snail) Kite (Rostrhamus sociabilis plumbeus) has revealed the following: extremely poor nesting success (only 13.6% of nests found at the nest-building stage successful); extremely long breeding seasons (some reproductive activity in almost all months in good years); frequent multiple brooding and frequent multiple brooding and frequent renesting after failure; low egg hatchability (81%); high failure rates due to nest collapse, desertion, and predation; extremely high survival of juveniles and adults under good water conditions; and high vulnerability to drought due to near total dependency on a single species of drought-sensitive snail for food. Despite low nesting success, the species has increased rapidly under good conditions, mainly because of multiple nesting attempts within long breeding seasons and high survival rates of free-flying birds. Nesting success varied significantly between regions and nest substrates, but not as a function of seasons or solitary vs. colonial nesting. While nesting success was reduced in low water years, this effect was at least partly due to heavy use of poor nest substrates under such conditions. Clutch size and numbers of young per successful nest varied with regions, but not as a function of seasons or water levels. The effects of coloniality on clutch size and numbers of young were inconsistent. Significant effects of nest-substrate types on clutch size and numbers of young were apparently artifacts of substrate differences between regions.

Condor

Demography and movements of the omao (Myadestes obscurus)

Density, age-specific survival, timing of breeding and molting, and movements of the Omao or Hawaiian Thrush (Myadestes obscurus) were studied at four sites on the island of Hawaii. Mean monthly density (birds/ha) was 3.23 +- 0.57, 1.07 +- 0.33, 3.23 +- 0. 16, and 3.74 +- 0.36 at Kau Forest, Hamakua. Keauhou Ranch, and Kilauea Forest study areas, respectively. Annual survival of juvenile Omao (0.40 +- 0.09) was lower than that of adults (0.66 +- 0.08). Emigration and mortality was greatest during November through May. Breeding and molting occurred throughout the year, with peak breeding in May through July. Omao showed strong site fidelity and were highly sedentary. Mean home range size (n = 39) was 2.20 t 0.26 ha and did not differ between sexes or study sites.

Condor

Demography of an introduced red-billed Leiothrix population in Hawaii

Relative abundance, timing of breeding and molting, annual survival, and philopatry of an introduced population of Red-billed Leiothrix ( Leiothrix lutea ) were studied at four sites on the island of Hawaii. Numbers of leiothrix on our study areas showed a regular cyclical pattern, with highest numbers during the breeding season. Peak breeding occurred May-August, followed by peak molting of flight and body feathers in August-October. Flocking behavior increased post-breeding, and many leiothrix left the study areas during the fall and winter months. Populations on two intensive study areas were stable, with high annual survival (x ± SE = 0.581 ± 0.115 for hatching-year birds and 0.786 ± 0.047 for adults). Leiothrix seem to be relatively unaffected by avian diseases that have decimated some Hawaiian bird populations, and yet reasons for their large historical population fluctuations remain unexplained.

Hawaii

Demography, environmental uncertainty, and the evolution of mate desertion in the snail kite

The Snail Kite (Rostrhamus sociabilis), an endangered hawk, has a unique mating system in Florida (Beissinger and Snyder 1987): when food is abundant, males or females desert their mates at nearly equal frequency (ambisexual mate desertion) in the midst of a nesting cycle. I examined the demographic and environmental constraints selecting for a clutch size that permits one parent to desert, yet optimizes the number of offspring produced by each parent. Demographic studies, conducted from 1979—1983, indicated that kites have (1) a very high nest failure rate (° = 68%) due most often to unstable nest sites and predation, (2) a variable nesting season (5—10 mo/yr), (3) an early age of first reproduction for a bird this size (10 mo), (4) a high degree of iteroparity (double and potentially triple clutching within a season), and (5) unstable populations. Both nesting success and population size were directly related to Everglades water levels and resultant snail densities. Kites responded to large annual changes in food abundance, not by adjusting clutch size but by deserting their mates and presumably attempting to renest. Kite demographic traits appear to be adaptations to or results of an uncertain environment. Based on 67 yr of Everglades water levels, environmental predictability, measured by spectral analysis and Colwell's (1974) index, was low and influenced by water management regimes: (1) water levels were lowered, (2) annual variation in levels increased and annual cycles became stronger, (3) the period length of long—term drought—flood cycles shifted from 10 or more yr toward 5 yr, and (4) levels became a less predictive cue for favorable nesting conditions. A potential evolutionary pathway from biparental care to mate desertion in Snail Kites is proposed. It is unlikely that mate desertion evolved solely from a context of conflict between the sexes, because kite nesting success is so low that it is probably maladaptive for a parent to desert and jeopardize the survival of any of its first brood. Instead, mate desertion behavior probably evolved in response to a smaller average clutch size; this would allow females to be highly iteroparous and avoid the costs of overinvestment, and should be strongly favored in a highly uncertain environment. Analysis of clutches in museum collections suggests that an apparent decline in clutch size may have occurred in Florida during the past century. The potential for ambisexual mate desertion to occur in other vertebrates is during the past century. The potential for ambisexual mate desertion to occur in other vertebrates is examined.

Ecology

Demography of birds in a neotropical forest: Effects of allometry, taxonomy, and ecology

Comparative demographic studies of terrestrial vertebrates have included few samples of species from tropical forests. We analyzed 9 yr of mark—recapture data and estimated demographic parameters for 25 species of birds inhabiting lowland forests in central Panama. These species were all songbirds (Order Passeriformes) ranging in mass from 7 to 57 g. Using Jolly—Seber stochastic models for open populations, we estimated annual survival rate, population size, and recruitment between sampling periods for each species. We then explored relationships between these parameters and attributes such as body size, phylogenetic affiliation, foraging guild, and social behavior. Larger birds had comparatively long life—spans and low recruitment, but body size was not associated with population size. After adjusting for effects of body size, we found no association between phylogenetic affiliation and any demographic trait. Ecological attributes, especially foraging guild, were more clearly associated with interspecific variation in all demographic traits. Ant—followers had comparatively long life—spans, but species that participate in flocks did not live longer than solitary species. The allometric associations we observed were consistent with those demonstrated in other studies of vertebrates; thus, these relationships appear to be robust. Our findings that ecological factors were more influential than phylogenetic affiliation contrasts with comparative studies of temperate—zone birds and suggests that the relative importance of environmental vs. historical factors varies geographically.

Ecology

Stem demography and post-fire recruitment of a resprouting serotinous conifer

The contribution of resprouts and seedling recruitment to post-fire regeneration of the South African fynbos conifer Widdringtonia nodiflora was compared eight months after wildfires in 1990. Stems on all trees were killed by fire but resprouting success was > 90 % at all but one site. A demographic study of burned skeletons revealed that prior to these fires, nearly all plants were multi-stemmed (4–9 stems/plant) and multi-aged, indicating continuous sprout production between fires. All stems were killed by these 1990 fires and at most sites > 90 % of the stems were burned to ground level. All diameter stems were susceptible to such incineration as, at most sites, there was no difference in average diameter of stems burned to ground level and those left standing. Individual genets usually had all ramets incinerated to ground level or all ramets charred, but intact, suggesting certain micro-sites burned hotter, whereas other sites were somewhat protected. Although not true of the 1990 fires, there was evidence that occasionally Widdring-tonia stems may survive fire. At one site, four of the 16 plants sampled had a burned stem twice as old as the oldest burned stem on the other 12 plants at the site, suggesting some stems had survived the previous fire (ca. 1970) and this conclusion was supported by fire-scars on these four stems that dated to ca. 1970. Based on the highly significant correlation between stem diameter and cone density left standing after the 1990 fires, we calculated that for most sites > 80 % of the initial cone crop was incinerated by fire. This is important because we observed a strong relationship between size of the canopy cone crop surviving fire and post-fire seedling recruitment. Under these conditions we hypothesize that sprouting confers a selective advantage to genets when fires cause heavy losses of seed. The infrequent occurrence of sprouting in the Cupressaceae suggests the hypothesis that resprouting is an apomorphic or derived trait in Widdringtonia. Data from this study suggests resprouting provides a selective advantage under severe fynbos fires, which are not only 'stand-replacing fires,’but also are intense enough to incinerate cone-bearing stems.

Journal of Vegetation Science

Distribution and demography of coastal Cactus Wrens (Campylorhynchus brunneicapillus) in southern San Diego County, California — 2021 Data summary

We surveyed for coastal Cactus Wren ( Campylorhynchus brunneicapillus ) in 378 established plots in southern San Diego County in 2021, encompassing 3 genetic clusters (Otay, Lake Jennings, and Sweetwater/Encanto). Two surveys were completed at each plot between March 1 and July 31. Cactus Wrens were detected in 130 plots (34 percent of plots), remaining virtually the same as the percentage of plots occupied in 2020 (35 percent). There were 113 Cactus Wren territories detected across all survey plots in 2021, an increase from 109 in 2020. At least 86 percent of Cactus Wren territories were occupied by pairs, and 50 fledglings were observed in 2021. We observed 48 color-banded Cactus Wrens in 2021, 44 of which we could identify to individual. Adults of known age ranged from 2 to at least 7 years old. Adult Cactus Wrens moved, on average, 0.1 kilometers (maximum 0.5 kilometers) from their 2020 territories to their 2021 territories. No known-identity Cactus Wrens moved between genetic clusters from 2020 to 2021. Vegetation at Cactus Wren plots typically was dominated by coastal sage scrub shrubs, such as California sagebrush ( Artemisia californica ), California buckwheat ( Eriogonum fasciculatum ), lemonadeberry ( Rhus integrifolia ), and San Diego sunflower ( Bahiopsis laciniata ). Twenty-nine percent of plots contained blue elderberry ( Sambucus nigra ssp. caerulea ), and Cactus Wrens occupied proportionally more plots with elderberry than plots without elderberry. Very little dead or unhealthy cactus was observed within all survey plots, and the plots that were occupied by Cactus Wrens were likely to contain more healthy cactus than plots that were not occupied by Cactus Wrens. Thirteen percent of plots had 5 percent or less of the cactus crowded or overtopped by vines and shrubs. Although in 2020, Cactus Wrens occupied proportionally more plots with 5 percent or less of cactus crowded or overtopped by vines and shrubs, this pattern was not found in 2021. Non-native annual cover was 5 percent or less at 29 percent of plots and, unlike in 2020, Cactus Wrens appeared to occupy proportionally more plots with less non-native annual cover than plots with more than 5-percent annual cover.

California

Distribution and demography of coastal Cactus Wrens ( Campylorhynchus brunneicapillus ) in San Diego County, California—2022 data summary

We surveyed for coastal Cactus Wren ( Campylorhynchus brunneicapillus ) in 507 established plots in San Diego County in 2022, encompassing 4 genetic clusters (Otay, Lake Jennings, Sweetwater/Encanto, and San Pasqual). Two surveys were completed at each plot between March 1 and July 31. Cactus Wrens were detected in 214 plots (42 percent of plots). Cactus Wrens were detected in 31 percent of plots that have been consistently surveyed since 2020, indicating less plot occupancy than in 2021 (34 percent) and 2020 (35 percent). There were 202 Cactus Wren territories detected across all survey plots in 2022. In plots that have been consistently surveyed since 2020, we documented 94 territories, which is a decrease from 113 territories in 2021 and 109 in 2020. The number of territories declined from 2021 to 2022 in the Lake Jennings and Otay genetic clusters but remained virtually the same in the Sweetwater/Encanto genetic cluster. At least 78 percent of Cactus Wren territories were occupied by pairs, and 134 fledglings were observed in 2022. We observed 24 banded Cactus Wrens in 2022, 20 of which we could identify individually by band combination. Adults of known age ranged from 3 to at least 7 years old. All individually identifiable adult Cactus Wrens remained on the same territory in 2022 that they occupied in 2021, and we detected no movement of banded Cactus Wrens between genetic clusters. Vegetation at Cactus Wren survey plots was dominated by coastal sage scrub shrubs, such as California sagebrush ( Artemisia californica ), California buckwheat ( Eriogonum fasciculatum ), lemonadeberry ( Rhus integrifolia ), jojoba ( Simmondsia chinensis ), and San Diego sunflower ( Bahiopsis laciniata ). No definitive signs of fungal pathogens were observed on cactus within and around survey plots. Blue elderberry ( Sambucus nigra ssp. caerulea ) was detected at 39 percent of plots, and Cactus Wrens occupied proportionally more plots with elderberry than plots without elderberry. Very little dead or unhealthy cactus was observed within all survey plots, and the plots that were occupied by Cactus Wrens were likely to contain fewer dead cactus than plots that were not occupied by Cactus Wrens. Almost 80 percent of plots had more than 5 percent of cactus crowded or overtopped by vines and shrubs. Similar to 2020, Cactus Wrens occupied proportionally more plots with 5 percent or less of cactus crowded or overtopped by vines and shrubs, although this pattern was not observed in 2021. Non-native annual cover was less prevalent in survey plots in 2022 than in 2021, and Cactus Wrens did not preferentially occupy survey plots with 5 percent or less non-native annual cover than plots with more non-native cover.

California

Distribution and demography of coastal cactus wrens (Campylorhynchus brunneicapillus) in southern San Diego County, California—2020 data summary

Executive Summary Surveys for coastal Cactus Wren ( Campylorhynchus brunneicapillus ) were done in 378 established plots in southern San Diego County in 2020, encompassing three genetic clusters (Otay, Lake Jennings, and Sweetwater/Encanto genetic clusters). Two surveys were completed at each plot between March 1 and July 31. Cactus Wrens were detected in 131 plots (35 percent of plots). This is a slight increase over the proportion of occupied plots in 2019. One hundred and nine Cactus Wren territories were detected across all survey plots in 2020, an increase from 83 in 2019. At least 85 percent of Cactus Wren territories were occupied by pairs, and 62 fledglings were observed in 2020. There were 89 color-banded Cactus Wrens observed in 2020, 84 of which we could identify to individual. Adults of known age ranged from 1 to at least 6 years old. Adult Cactus Wrens moved on average 0.2 kilometers (km; maximum 3.8 km) from their 2019 territories to their 2020 territories. Cactus Wrens that fledged in 2019 moved on average 1.2 km (maximum 9.9 km) to their 2020 territories. No known-identity Cactus Wrens moved between genetic clusters from 2019 to 2020. Vegetation at Cactus Wren plots typically was dominated by coastal sage scrub shrubs such as California sagebrush ( Artemisia californica ), lemonadeberry ( Rhus integrifolia ), California buckwheat ( Eriogonum fasciculatum ), and broom baccharis ( Baccharis sarothroides ). Very little dead or unhealthy cactus was observed within Cactus Wren survey plots. Thirty-eight percent of plots had at least 25 percent of the cactus crowded or overtopped by vines and shrubs. Non-native annual cover was greater than 25 percent at 35 percent of plots.

California

Effects of experimental removal of barred owls on population demography of northern spotted owls in Washington and Oregon—2015 progress report

Evidence indicates that competition with newly established barred owls ( Strix varia ) is causing rapid declines in populations of northern spotted owls ( Strix occidentalis caurina ), and that the longterm persistence of spotted owls may be in question without additional management intervention. A pilot study in California showed that lethal removal of barred owls in combination with habitat conservation may be able to slow or even reverse population declines of spotted owls at local scales, but it remains unknown whether similar results can be obtained in larger areas with different forest conditions and where barred owls are more abundant. In 2015, we implemented a before-after-controlimpact (BACI) experimental design on two study areas in Oregon and Washington with at least 20 years of pre-treatment demographic data on spotted owls to determine if removal of barred owls can improve population trends of spatially associated spotted owls. Here we provide an overview of our research accomplishments and preliminary results in Oregon and Washington in 2015.

Oregon, Washington

Effects of experimental removal of barred owls on population demography of northern spotted owls in Washington and Oregon—2016 progress report

Evidence indicates that competition with invasive barred owls ( Strix varia ) is causing rapid declines in populations of northern spotted owls ( S. occidentalis caurina ), and that the long-term persistence of spotted owls may be in question without additional management intervention. A pilot study in California showed that removal of barred owls in combination with habitat conservation may be able to slow or even reverse population declines of spotted owls at local scales, but it remains unknown whether similar results can be obtained in areas with different forest conditions and a greater density of barred owls. In 2015, we implemented a before-after-control-impact (BACI) experimental design on three study areas in Oregon and Washington with at least 20 years of pre-treatment demographic data on spotted owls to determine if removal of barred owls can improve localized population trends of spotted owls. Here, we report on research accomplishments and preliminary results from the first 21 months (March 2015–December 2016) of the planned 5-year experiment.

Oregon, Washington

Distribution and demography of San Francisco gartersnakes (Thamnophis sirtalis tetrataenia) at Mindego Ranch, Russian Ridge Open Space Preserve, San Mateo County, California

San Francisco gartersnakes ( Thamnophis sirtalis tetrataenia ) are a subspecies of common gartersnakes endemic to the San Francisco Peninsula of northern California. Because of habitat loss and collection for the pet trade, San Francisco gartersnakes were listed as endangered under the precursor to the Federal Endangered Species Act. A population of San Francisco gartersnakes resides at Mindego Ranch, San Mateo County, which is part of the Russian Ridge Open Space Preserve owned and managed by the Midpeninsula Regional Open Space District (MROSD). Because the site contained non-native fishes and American bullfrogs ( Lithobates catesbeianus ), MROSD implemented management to eliminate or reduce the abundance of these non-native species in 2014. We monitored the population using capture-mark-recapture techniques to document changes in the population during and following management actions. Although drought confounded some aspects of inference about the effects of management, prey and San Francisco gartersnake populations generally increased following draining of Aquatic Feature 3. Continued management of the site to keep invasive aquatic predators from recolonizing or increasing in abundance, as well as vegetation management that promotes heterogeneous grassland/shrubland near wetlands, likely would benefit this population of San Francisco gartersnakes.

California

Effects of experimental removal of Barred Owls on population demography of Northern Spotted Owls in Washington and Oregon—2017 progress report

Populations of Northern Spotted Owls ( Strix occidentalis caurina ; hereinafter referred to as Spotted Owl) are declining throughout this subspecies’ geographic range. Evidence indicates that competition with invading populations of Barred Owls ( S. varia ) has contributed significantly to those declines. A pilot study in California showed that localized removal of Barred Owls coupled with conservation of suitable forest conditions can slow or even reverse population declines of Spotted Owls. It remains unknown, however, whether similar results can be obtained in areas with different forest conditions, greater densities of Barred Owls, and fewer remaining Spotted Owls. During 2015–17, we initiated a before-after-control-impact (BACI) experiment at three study areas in Oregon and Washington to determine if removal of Barred Owls can improve population trends of Spotted Owls. Each study area had at least 20 years of pre-treatment demographic data on Spotted Owls, and represented different forest conditions occupied by the two owl species in the Pacific Northwest. This report describes research accomplishments and preliminary results from the first 2.5 years (March 2015–August 2017) of the planned 5-year experiment.

Oregon, Washington

Effects of experimental removal of Barred Owls on population Demography of Northern Spotted Owls in Washington and Oregon—2018 Progress Report

Populations of Northern Spotted Owls ( Strix occidentalis caurina ; herineafter referred to as Spotted Owl) have declined throughout the subspecies’ geographic range. Evidence indicates that competition with invading Barred Owls ( S. varia ) has contributed significantly to those declines. A pilot study in California showed that removal of Barred Owls coupled with conservation of suitable habitat conditions can slow or even reverse population declines of Spotted Owls. It is unknown, however, whether similar results can be obtained in areas with different forest conditions, greater densities of Barred Owls, and fewer remaining Spotted Owls. We initiated a before-after-control-impact (BACI) experiment at three study areas in Oregon and Washington to determine if removal of Barred Owls can improve population trends of Spotted Owls. This report describes research accomplishments and initial results from the first 3.5 years of the study (March 2015–August 2018).

Oregon, Washington

Effects of barred owl (Strix varia) removal on population demography of northern spotted owls (Strix occidentalis caurina) in Washington and Oregon—2019 annual report

Strix occidentalis caurina (northern spotted owl; hereinafter referred to as spotted owl) have rapidly declined throughout the subspecies’ geographic range. Competition with invading Strix varia (barred owl) has been identified as an immediate cause of those declines. A pilot study in California showed that removal of barred owls coupled with conservation of suitable habitat conditions can slow or even reverse population declines of spotted owls. It is unknown, however, whether similar results can be obtained in areas with different forest conditions, greater densities of barred owls, and fewer remaining spotted owls. We used a before-after-control-impact experimental design on three study areas with long-term demographic information on spotted owls to determine if removal of barred owls can improve population trends of spotted owls. This report summarizes research accomplishments and initial results from the first 4.5 years (from March 2015 to August 2019) of implementing barred owl removal experiments in Washington and Oregon.

Oregon, Washington

Distribution and demography of Coastal Cactus Wrens in Southern California, 2015–19

Surveys and monitoring for the coastal Cactus Wren ( Campylorhynchus brunneicapillus ) were completed in San Diego County between March 2015 and July 2019. A total of 383 plots were surveyed across 3 genetic clusters (Otay, Lake Jennings, and Sweetwater/Encanto). From 2015 to 2019, 317 plots were surveyed 8 times (twice per year in 2015, 2017–19). Additional plots were added in later years as wrens were discovered in new locations. We found differences in the proportion of plots occupied in the genetic clusters, with a lower proportion of plots occupied in the Otay cluster than in the Lake Jennings and Sweetwater/Encanto clusters in all years. Plot occupancy increased each year in the Otay and Sweetwater/Encanto clusters but not in the Lake Jennings cluster. The number of Cactus Wren territories increased from 2015 through 2018, and then decreased in 2019 in all three genetic clusters. We monitored nesting activities for two populations of Cactus Wrens in southern San Diego County. The Otay population consisted of two sites within the Otay genetic cluster, and the San Diego population consisted of two sites within the Sweetwater/Encanto and Lake Jennings genetic clusters. Nest monitoring occurred at 10–13 territories per year in the Otay population and 14–18 territories in the San Diego population from 2015 through 2019. All territories were occupied by pairs except two territories in 2015, five in 2016, and two in 2019. Between 46 and 74 Cactus Wren nests were monitored each year, which totaled 295 monitored nests from 2015 to 2019. To evaluate the direct influence of precipitation on breeding success, bio-year precipitation (“precipitation”) was calculated from July 1 of the prior year through June 30 of the breeding season year. Overall apparent nest success was positively influenced by precipitation with the lowest apparent nest success of 50 percent in 2015 and the highest apparent nest success of 72 percent in 2017, corresponding to the second lowest and the highest precipitation years, respectively. Apparent nest success also was higher in the Otay population than in the San Diego population. The number of brood nests initiated per pair and the number of renesting attempts per pair also were higher in years with more precipitation. Other metrics of Cactus Wren nesting success and productivity were positively influenced by the amount of precipitation, including clutch size and egg hatching success. The percent of hatchlings that fledged was greater in the Otay population than in the San Diego population but was not influenced by precipitation. The number of fledglings per pair was higher in years with more precipitation and was greater in the Otay population than in the San Diego population. Predation was the predominant cause of nest failure in both populations. Analysis of Cactus Wren daily nest survival rate indicated that there was a population, and possibly a precipitation effect on nest survival, with the daily survival rate for the Otay population significantly higher than for the San Diego population and weak increase in the daily survival rate with more precipitation. A total of 629 Cactus Wrens were banded during the course of the study, 360 in the San Diego population and 269 in the Otay population. Between 2015 and 2019, we resighted 301 color-banded adult birds that ranged between 1 and 8 years old. One additional color-banded bird was resighted in San Pasqual Valley (as part of a separate study); this bird originated in the San Diego population and was excluded from our analyses. Annual survival was higher for adult Cactus Wrens (ranging from 60 to 70 percent) than for first-year wrens (ranging from 20 to 28 percent) and varied by year. Annual survival was also weakly but positively correlated with precipitation. Annual survival was higher for first year and adult Cactus Wrens following years with increased precipitation. We found no evidence that survival differed by population. Banding also allowed us to examine whether there were differences in movement of adult and first-year Cactus Wrens by year or by population. We found that average dispersal distance for first-year Cactus Wrens was 1.9 kilometers in the Otay population and 1.6 kilometers in the San Diego population and did not differ by population or year. Dispersal between populations was not common. We detected five instances of movement of first-year wrens between the San Diego and Otay populations. All movements into and out of the San Diego population were from or into territories in the Sweetwater area. We detected no movement between the Lake Jennings site and either of the Sweetwater or Otay sites; however, we did detect one wren that dispersed from Lake Jennings to the San Pasqual Valley population in 2019, which was a distance of 26.4 kilometers. Adult Cactus Wrens were site-faithful, with 87 percent of adults remaining on the same territory between breeding seasons. Precipitation may be a weak driver of movement for adult Cactus Wrens, with adults more likely to remain on the same territory following years of increased precipitation. There was no difference in adult movement between populations. Arthropods were collected in pitfall traps and by vacuum in 23 Cactus Wren territories during 3 sampling periods in 2016 (early nesting, peak nesting, and late nesting). Arthropods of 19 orders and at least 128 families were collected. Analysis of 43 Cactus Wren fecal samples identified 10 arthropod orders that were present in more than 10 percent of fecal samples. The most abundant arthropod order collected was Hymenoptera; however, Cactus Wrens consumed arthropods in the order Hymenoptera significantly less than their availability, suggesting that this order was avoided. No other orders were significantly selected or avoided; however, selection indices of arthropod families identified that two families of arthropods (Isopoda Porcellionidae [woodlice] and Hymenoptera Formicidae [ants]) were avoided. After excluding the taxa that were avoided or not represented in fecal samples, 95 percent of Cactus Wren prey items were collected in pitfall traps and 5 percent were collected by vacuum. The most abundant prey orders captured were Diptera, Coleoptera, Hemiptera, Hymenoptera, and Aranea. Analysis of the abundance of Cactus Wren prey items by vegetation type and sampling period indicated that vegetation type by itself was not a significant predictor of arthropod abundance but interacted with sampling period. Seasonal availability of arthropods was highest in the peak nesting period, followed by early and late nesting periods for California sagebrush ( Artemisia californica ), lemonadeberry ( Rhus integrifolia ), non-native grass, and bare ground, whereas availability increased from early to late nesting periods for blue elderberry ( Sambucus mexicana spp. caerulea ), cactus ( Opuntia spp. and Cylindropuntia spp.), California buckwheat ( Eriogonum fasciculatum ), native bunch grasses, and black mustard ( Brassica nigra ). During the early nesting period, arthropods were most abundant in native bunch grasses and least abundant in lemonadeberry. During the peak nesting period, arthropods were most abundant in native bunch grasses and in areas of bare ground and were least abundant in cactus and blue elderberry. During late nesting, arthropods were most abundant in blue elderberry and non-native grass and least abundant in lemonadeberry and mustard. Each year from 2015 to 2019, vegetation data were collected at the same 23 territories where arthropods were sampled: 9 territories in the Otay population and 14 territories in the San Diego population. Cactus, California buckwheat, and non-native grasses were detected within at least 60 percent of sampling points in the Otay population. Cactus, California sagebrush, California buckwheat, non-native grass, and black mustard each were detected within an average of 40 percent of sampling points in the San Diego population. No native bunch grass or lemonadeberry were recorded at the Lake Jennings site within the San Diego population. The cover of shrub species was relatively stable throughout the 5 years. Cover of herbaceous species and bare ground had greater annual variation than shrub species. We found that vegetation cover varied widely among territories, with territory accounting for 69 percent of the variation in vegetation cover. Redundancy analysis allowed us to identify the vegetation types that accounted for the most variation. We used the top scores from the redundancy analysis to identify six vegetation types to be used in generalized linear mixed models analyzing the relationships between vegetation type, precipitation, and Cactus Wren breeding productivity. Three vegetation variables influenced the number of fledglings produced per pair. California sagebrush had a positive effect on the number of fledglings per pair whereas non-native grass and black mustard had a negative effect. Breeding productivity, survival, and movements of adult and first-year Cactus Wrens indicated that the Otay population behaved similarly to, if not out-performed, the San Diego population during the span of our project, suggesting that the driving forces behind low numbers of Cactus Wrens in the Otay population before 2015 were no longer in effect. The Cactus Wren populations in Otay and San Diego reached a peak in 2018, which followed a year of high productivity and survivorship, both of which were correlated with high precipitation. This peak in population size was consistent with reproductive timing and productivity in other bird populations in semi-arid ecosystems that were linked to precipitation and arthropod abundance. We did not find a strong link among arthropod abundance, vegetation composition, and Cactus Wren breeding productivity, likely in part because arthropod abundance varied by vegetation type and sampling period, suggesting that different vegetation types provided important sources of prey at different periods of the breeding season. Arthropod abundance also may not represent arthropod availability when vegetation structure discourages the ground foraging behavior of species such as Cactus Wrens. Cover of non-native grass negatively influenced breeding productivity, although arthropods were abundant in non-native grass. Other factors that could have influenced differential breeding productivity between the Otay and San Diego populations were habitat restoration, control of annual herbaceous vegetation, human disturbance, lingering effects of wildfire, and nest predation. Overall, precipitation appeared to be a driver of Cactus Wren breeding productivity and possibly survival, potentially obscuring proximate effects of arthropod or vegetation composition.

California