Western pond turtles in the Mojave Desert? A review of their past, present, and possible future

The western pond turtle (WPT) was formerly considered a single species (Actinemys or Emys marmorata) that ranged from southern British Columbia, Canada to Baja California, México. More recently it was divided into a northern and a southern species. WPTs are found primarily in streams that drain into the Pacific Ocean, although scattered populations exist in endorheic drainages of the Great Basin and Mojave deserts. Populations in the Mojave Desert were long thought to be restricted to the Mojave River, but recently another population was documented in Piute Ponds, a terminal wetland complex associated with Amargosa Creek on Edwards Air Force Base. WPT fossils in the Mojave Desert are known from the Miocene to the Pleistocene. Recently, Pleistocene fossils have been found as far into the desert as Salt Springs, just south of Death Valley. The oldest fossil records suggest that WPTs were present in wetlands and drainages of the geological feature known as the Mojave block prior to the uplift of the Sierra Nevada Range about 8 Ma and prior to the ~ 3 Ma uplift of the Transverse Ranges. Archaeological records document use of turtles by Native Americans for food and cultural purposes 1,000 or more years ago at the Cronese Lakes on the lower Mojave River and Oro Grande on the upper river. The first modern publication documenting their presence in the Mojave River was 1861. Museum specimens were collected as early as 1937. These fossil and early literature records support the indigenous status of WPTs to the Mojave River. However, mtDNA-based genetic evidence shows that Mojave River turtles share an identical haplotype with turtles on the California coast. Limited nuclear data show some minor differences. Overdraft of water from the Mojave River for municipal and agricultural uses, urban development, and saltcedar expansion are threats to the continued survival of WPTs in the Mojave River.

Historical museum collections and contemporary population studies implicate roads and introduced predatory bullfrogs in the decline of western pond turtles

The western pond turtle (WPT), recently separated into two paripatrically distributed species (Emys pallida and Emys marmorata), is experiencing significant reductions in its range and population size. In addition to habitat loss, two potential causes of decline are female-biased road mortality and high juvenile mortality from non-native predatory bullfrogs (Rana catesbeiana). However, quantitative analyses of these threats have never been conducted for either species of WPT. We used a combination of historical museum samples and published and unpublished field studies shared with us through personal communications with WPT field researchers (B. Shaffer, P. Scott, R. Fisher, C. Brown, R. Dagit, L. Patterson, T. Engstrom, 2019, personal communications) to quantify the effect of roads and bullfrogs on WPT populations along the west coast of the United States. Both species of WPT shift toward increasingly male biased museum collections over the last century, a trend consistent with increasing, female-biased road mortality. Recent WPT population studies revealed that road density and proximity were significantly associated with increasingly male-biased sex ratios, further suggesting female-biased road mortality. The mean body size of museum collections of E. marmorata, but not E. pallida, has increased over the last 100 years, consistent with reduced recruitment and aging populations that could be driven by invasive predators. Contemporary WPT population sites that co-occur with bullfrogs had significantly greater average body sizes than population sites without bullfrogs, suggesting strong bullfrog predation on small WPT hatchlings and juveniles. Overall, our findings indicate that both species of WPT face demographic challenges which would have been difficult to document without the use of both historical data from natural history collections and contemporary demographic field data. Although correlational, our analyses suggest that female-biased road mortality and predation on small turtles by non-native bullfrogs are occurring, and that conservation strategies reducing both may be important for WPT recovery.

Experimental removal of introduced slider turtles offers new insight into competition with a native, threatened turtle

The red-eared slider turtle (Trachemys scripta elegans; RES) is often considered one of the world’s most invasive species. Results from laboratory and mesocosm experiments suggest that introduced RES outcompete native turtles for key ecological resources, but such experiments can overestimate the strength of competition. We report on the first field experiment with a wild turtle community, involving introduced RES and a declining native species of conservation concern, the western pond turtle (Emys marmorata; WPT). Using a before/after experimental design, we show that after removing most of an introduced RES population, the remaining RES dramatically shifted their spatial basking distribution in a manner consistent with strong intraspecific competition. WPT also altered their spatial basking distribution after the RES removal, but in ways inconsistent with strong interspecific competition. However, we documented reduced levels of WPT basking post-removal, which may reflect a behavioral shift attributable to the lower density of the turtle community. WPT body condition also increased after we removed RES, consistent with either indirect or direct competition between WPT and RES and providing the first evidence that RES can compete with a native turtle in the wild. We conclude that the negative impacts on WPT basking by RES in natural contexts are more limited than suggested by experiments with captive turtles, although wild WPT do appear to compete for food with introduced RES. Our results highlight the importance of manipulative field experiments when studying biological invasions, and the potential value of RES removal as a management strategy for WPT.

Physiological consequences of rising water salinity for a declining freshwater turtle

Sea-level rise, drought and water diversion can all lead to rapid salinization of freshwater habitats, especially in coastal areas. Increased water salinities can in turn alter the geographic distribution and ecology of freshwater species including turtles. The physiological consequences of salinization for freshwater turtles, however, are poorly known. Here, we compared the osmoregulatory response of two geographically separate populations of the freshwater Western Pond Turtle (Actinemys marmorata)—a species declining across its range in western North America—to three constant salinities: 0.4 ppt, 10 ppt and 15 ppt over 2 weeks. We found that turtles from a coastal estuarine marsh population regulated their plasma osmolality at lower levels than their conspecifics from an inland freshwater creek population 45 km away. Plasma osmolalities were consistently lower in estuarine marsh turtles than the freshwater creek turtles over the entire 2-week exposure to 10 ppt and 15 ppt water. Furthermore, estuarine marsh turtles maintained plasma osmolalities within 1 SD of their mean field osmolalities over the 2-week exposure, whereas freshwater creek turtles exceeded their field values within the first few days after exposure to elevated salinities. However, individuals from both populations exhibited body mass loss in 15 ppt water, with significantly greater loss in estuarine turtles. We speculate that the greater ability to osmoregulate by the estuarine marsh turtles may be explained by their reduced feeding and drinking in elevated salinities that was not exhibited by the freshwater creek population. However, due to mass loss in both populations, physiological and behavioural responses exhibited by estuarine marsh turtles may only be effective adaptations for short-term exposures to elevated salinities, such as those from tides and when traversing saline habitats, and are unlikely to be effective for long-term exposure to elevated salinity as is expected under sea-level rise.

Archaeology and Biogeography of the Western Pond Turtle (Actinemys marmorata) in the Puget Sound Region

The modern distribution of the western pond turtle (Actinemys marmorata) is discontinuous, with a historic but extirpated population in the Puget Sound region that was isolated from populations south along the Columbia River. To better understand this distribution, a review of the archaeological literature for the Puget Sound region was conducted to determine the prehistoric biogeography of the species in the Puget Sound area. Western pond turtles are nearly absent from the regional archaeological record, represented at best by four tentative specimens. This may be explained by extremely low population levels throughout the Holocene at the northernmost extent of its range.

Effects of Drought on Western Pond Turtle Survival and Movement Patterns

Drought has the ability to affect the persistence of small animal populations, especially those tied to aquatic habitats. We studied the response of western pond turtles Actinemys marmorata to California's worst drought on record. From 2009 through 2015 we used telemetry to track movements and assess survival of 19 western pond turtles in a stock pond at the San Joaquin Experimental Range in the western foothills of the Sierra Nevada, in Madera County, California. In 2013 the pond dried in late summer and winter rains were insufficient for pond formation. The pond remained dry through the end of the study in March 2015. In years with below average precipitation the pond often dried completely in late summer; however, the lack of a pond forming in winter and spring had not been previously documented. We observed no mortalities of radiotagged western pond turtles in years with normal precipitation. All observed mortalities occurred in drought years and in years when the pond completely dried up in the summer or never formed. Results from known-fate survival models revealed that survival decreased with increasing drought. Model results also indicated that male survival was slightly higher than female survival (19.1% vs. 11.5%), although the 95% confidence intervals overlapped. We observed high variability in western pond turtle movement distances from the pond in the final 2 y of the study. Two individuals that survived to the end of the study showed unique movement patterns. One young male moved frequently, accumulating a large total distance, moved into new areas, and eventually found his way into a livestock water trough. The other, a young female, moved 2.6 km from the pond (a minimum total distance traveled of 3.3 km based on telemetry locations) and emigrated to a pond on a neighboring ranch. Turtles that died exhibited no distinctive behaviors. After the pond dried western pond turtles remained terrestrial for long periods, with one surviving individual remaining out of water for 617 consecutive days, which is an unprecedented finding for this species to our knowledge. Our findings suggest that increased frequency and severity of droughts can affect the resiliency of small, isolated western pond turtle populations, especially those in ephemeral aquatic environments. These small populations are essential to the long-term survival of the species because of the current fragmented distribution of the species.