The purpose of this report is to provide the Marine Corps with an annual summary of abundance, breeding activity, demography, and habitat use of endangered Least Bell’s Vireos (Vireo bellii pusillus) at Marine Corps Base Camp Pendleton (MCBCP or “Base”). Surveys for the Least Bell's Vireo were completed at MCBCP, California, between April 5 and July 13, 2021. Core survey areas and a subset of non-core areas in drainages containing riparian habitat suitable for vireos were surveyed three to four times. We detected 551 territorial male vireos and 26 transient vireos in core survey areas. An additional 98 territorial male vireos were detected in non-core survey areas. Transient vireos were detected on 8 of the 10 drainages/sites surveyed (core and non-core areas). Of the vireo territories in core areas, 89 percent were on the four most populated drainages, with the Santa Margarita River containing 70 percent of all territories in areas surveyed on Base. In core areas, 75 percent of male vireos were confirmed as paired; 76 percent of male vireos in non-core areas were confirmed as paired.

The number of documented Least Bell’s Vireo territories in core survey areas on MCBCP decreased 18 percent from 2020 to 2021. The number of territories in all but two core survey area drainages decreased by one or more between 2020 and 2021. The decrease in vireo numbers on MCBCP (18 percent) was consistent with population changes in surrounding areas, including the lower San Luis Rey River (24-percent decrease) and the middle San Luis Rey River (6-percent decrease).

Most core-area vireo territories (59 percent of males) were in willow (Salix spp.) riparian habitat. An additional 7 percent of birds occupied willow habitat co-dominated by Western sycamores (Platanus racemosa) or Fremont cottonwoods (Populus fremontii). Of all the territories surveyed, 25 percent were in riparian scrub dominated by mule fat (Baccharis salicifolia) or sandbar willow (S. exigua). Upland scrub was used by 8 percent of vireos; 1 percent of vireo territories were in non-native vegetation, and less than 1 percent of vireo territories were in alder or drier habitats co-dominated by coast live oak (Quercus agrifolia) and sycamore.

In 2019, MCBCP began operating an artificial seep along the Santa Margarita River; then, in 2021, two additional artificial seeps became operational. The artificial seeps pumped water to the surface starting in March and ending in August each year during daylight hours and were designed to increase the amount of surface water present to enhance Southwestern Willow Flycatcher (Empidonax traillii extimus) breeding habitat. Although this enhancement was designed to benefit flycatchers, few flycatchers have inhabited the seep areas within the past several years; therefore, vireos were selected as a surrogate species to determine effects of the habitat enhancement. This report presents the second year of analyses of vireo and vegetation response to the artificial seeps.

We sampled vegetation in two Seep sites and two Reference sites to determine the effects of a new water diversion dam that was completed in 2019 and two seep pumps that were installed to enhance surface water along the Santa Margarita River in 2019 and 2021. We measured higher total vegetation cover below 2 meters (m) at Seep sites than at Reference sites and lower total vegetation cover above 5 m at Seep sites than at Reference sites. Native herbaceous cover was also higher below 4 m at Seep sites than at Reference sites. Woody cover was lower above 5 m at Seep sites than at Reference sites. Soil moisture did not differ between Seep and Reference sites.

The U.S. Geological Survey has been color banding Least Bell’s Vireos on Marine Corps Base Camp Pendleton since 1995. In 2021, we continued to color band and resight color banded Least Bell’s Vireos to evaluate adult site fidelity, between-year movement, and the effect of surface-water enhancement on vireo site fidelity and between-year movement. We banded 164 Least Bell's Vireos for the first time during the 2021 season. Birds banded included 3 adult vireos and 161 juvenile vireos. All adult vireos were banded with unique color combinations. The juvenile vireos (all nestlings) were banded with a single gold numbered federal band on the right leg.

There were 52 Least Bell's Vireos banded before the 2021 breeding season that were resighted and identified on Base in 2021. Of these vireos, 45 were banded on Base, 6 were originally banded on the San Luis Rey River, and 1 was banded at Marine Corps Air Station, Camp Pendleton. Adult birds of known age ranged from 1 to at least 7 years old.

Base-wide survival of vireos was affected by sex, age, and year. Males had a slightly but significantly higher survival rate than females. Adults had a higher survival rate than first-year vireos. Survival of both adults and first-year birds was high from 2007 to 2008 and from 2012 to 2013 and low from 2020 to 2021. The return rate of adult vireos to Seep or Reference sites ranged from 45 to 57 percent.

Most returning adult vireos showed strong between-year site fidelity. Of the adults present in 2020 and 2021, 84 percent (94 percent of males; no females) returned to within 100 m of their previous territory. The average between-year movement for returning adult vireos was 0.1±0.2 kilometer (km). The average movement of first-year vireos detected in 2021 that fledged from a known nest on MCBCP in 2020 was 1.1±0.7 km.

We monitored Least Bell's Vireo pairs to evaluate the effects of surface-water enhancement on nest success and breeding productivity. Vireos were monitored at two Seep sites and two Reference sites. Early in 2021, a seep was installed in a 2020 Reference site, which changed the status of this monitoring site from Reference to Seep.

Nesting activity was monitored between April 5 and July 22 in 50 territories within the Seep and Reference sites (25 at Seep sites and 25 at Reference sites). All territories, except one, were occupied by pairs and all were fully monitored, meaning all nesting attempts were monitored at these territories. During the monitoring period, 97 nests (42 in Seep sites and 55 in Reference sites) were monitored.

Breeding productivity was similar at the Seep site and Reference sites (3.6 and 3.4 young per pair, respectively), with 84 percent of Seep pairs and 88 percent of Reference pairs successfully fledging at least one young in 2021. Seep sites had a higher proportion of all eggs that hatched and also a higher proportion of nests with eggs that hatched than Reference sites. Seep sites and References sites had similar proportions of hatchlings that fledged and nests with hatchlings that fledged. According to the best model, daily nest survival in 2021 was higher in Seep sites than in Reference sites. Completed nests at the Seep site were more likely to be successful than nests at Reference sites in 2021. At Seep sites, 75 percent of nests fledged young, whereas 53 percent of nests at Reference successfully fledged young. Vireos at Reference sites had to expend more energy in extra nest-building and egg-laying to produce a similar number of young as vireos at Seep sites. Predation was believed to be the primary source of nest failure at both sites. Predation accounted for 100 percent and 83 percent of nest failures at Seep and Reference sites, respectively. Failure of the remaining nests was attributed to infertile eggs and other unknown causes.

There were 11 plant species used as hosts for vireo nests in 2021. Successful vireo nests at Reference sites were further from the edge of host plants (closer to the center) and further from the edge of the nest plant clump than unsuccessful nests. Vireo nests at Seep sites were further from the edge of the host plant and the nest plant clump than vireo nests at Reference sites.