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

Abstract

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.