Sarah J. Kupferberg

BULLFROG (RANA CATESBEIANA) INVASION OF A CALIFORNIA RIVER: THE ROLE OF LARVAL COMPETITION

I studied the invasion of Rana catesbeiana (the bullfrog) into a northern California river system where bullfrogs are not native. Native yellow-legged frogs, Rana boylii, a species of special concern, were almost an order of magnitude less abundant in reaches where bullfrogs were well established. I assessed the potential role of larval com- petition in contributing to this displacement in a series of field manipulations of tadpole density and species composition. The impact of R. catesbeiana on native tadpoles in the natural community agreed with the outcome of more artificial experiments testing pairwise and three-way interactions. In 2-M2 enclosures with ambient densities of tadpoles and natural river biota, bullfrog tadpoles caused a 48% reduction in survivorship of R. boylii, and a 24% decline in mass at metamorphosis. Bullfrog larvae had smaller impacts on Pacific treefrogs, Hyla regilla, causing 16% reduction in metamorph size, and no significant effect on survivorship. Bullfrog tadpoles significantly affected benthic algae, although effects varied across sites. Responses to bullfrogs in field settings were similar qualitatively to results seen in smaller-scale experiments designed to study size-structured competition among disparate age/size classes of species pairs and trios. Competition from large over- wintering bullfrog larvae significantly decreased survivorship and growth of native tadpoles. Competition from recently hatched bullfrog larvae also decreased survivorship of R. boylii and H. regilla. Native species competed weakly, both interspecifically and intraspecifically. The only suggestion of a negative impact of a native species on bullfrogs was a weak effect of H. regilla on recent hatchlings. Competition appeared to be mediated by algal resources, and there was no evidence for behavioral or chemical interference. These results indicate that, through larval interactions, bullfrogs can exert differential effects on native frogs and perturb aquatic community structure.

Effects of Variation in Natural Algal and Detrital Diets on Larval Anuran (Hyla regilla) Life-History Traits

The quality of algae as food for larval Pacific treefrogs, Hyla regilla, varies among algal taxa. Tadpole diets were manipulated in enclosures in the South Fork Eel River, northern California. Enclosed tadpoles were fed ad libitum one of the dominant filamentous green algae (Cladophora, Zygnema, Mougeotia, or Oedogonium), the dominant cyanobacterium in the habitat (Nostoc), flocculent detritus, or a commercial reptile/amphibian food. Tadpoles in a control treatment were not fed but had access to seston which deposited in all enclosures. Diet treatment significantly affected weekly growth rates, as well as time to and weight at metamorphosis. Tadpoles grew most rapidly on filamentous green algae with diatom epiphytes and on the commercial food. Tadpoles fed epiphytized Cladophora metamorphosed 27.1% (11.7 days) sooner and weighed 38.9% (70 mg) more than tadpoles fed Cladophora cleaned of diatoms. In comparison to tadpoles fed Mougeotia, which does not support epiphytes, tadpoles fed epiphytized Cladophora metamorphosed 37.4% (19.7 days) sooner and weighed 31.6% (50 g) more. Tadpoles with the lowest growth rates were those filtering deposited seston or flocculent detritus. Observed changes in life-history traits with diet are assessed in relation to models which predict the optimal timing of metamorphosis.

Effects of Flow Regimes Altered by Dams on Survival, Population Declines, and Range-Wide Losses of California River-Breeding Frogs

Widespread alteration of natural hydrologic patterns by large dams combined with peak demands for power and water delivery during summer months have resulted in frequent aseasonal flow pulses in rivers of western North America. Native species in these ecosystems have evolved with predictable annual flood-drought cycles; thus, their likelihood of persistence may decrease in response to disruption of the seasonal synchrony between stable low-flow conditions and reproduction. We evaluated whether altered flow regimes affected 2 native frogs in California and Oregon (U.S.A.) at 4 spatial and temporal extents. We examined changes in species distribution over approximately 50 years, current population density in 11 regulated and 16 unregulated rivers, temporal trends in abundance among populations occupying rivers with different hydrologic histories, and within-year patterns of survival relative to seasonal hydrology. The foothill yellow-legged frog (Rana boylii), which breeds only in flowing water, is more likely to be absent downstream of large dams than in free-flowing rivers, and breeding populations are on average 5 times smaller in regulated rivers than in unregulated rivers. Time series data (range = 8 - 19 years) from 5 populations of yellow-legged frogs and 2 populations of California red-legged frogs (R. draytonii) across a gradient of natural to highly artificial timing and magnitude of flooding indicate that variability of flows in spring and summer is strongly correlated with high mortality of early life stages and subsequent decreases in densities of adult females. Flow management that better mimics natural flow timing is likely to promote persistence of these species and others with similar phenology.

Development and Evolution of Aquatic Larval Feeding Mechanisms

Publisher Summary This chapter discusses the development and evolution of feeding mechanisms in aquatic larval vertebrates, such as fishes and amphibians. Fish larva is the immature life-history stage that differs morphologically from the juvenile and adult. The time between hatching and metamorphosis may range from a few days in tropical fish species, to a few weeks or months in most temperate fishes, to years in the eel Anguilla. In amphibians, the larval stage begins at hatching, when an embryo escapes from its egg capsule. Larvae are morphologically distinct from adults and are not reproductive. Fish larvae undergo trophic ontogeny as their diet changes markedly with growth. Four types of nutrient acquisitions are recognized in fish larvae: endogenous, absorptive, mixed, and exogenous. For the embryos of oviparous and ovoviviparous fishes, yolk platelets and oil globules serve as the primary source of nutrition. Many marine invertebrate larvae are capable of supplementing their nutrition by the uptake of dissolved organic matter (DOM) via transepidermal transport. Mixed feeding refers to any combination of endogenous, absorptive, and exogenous feeding, and occurs as concurrent endogenous and exogenous feeding during the time between first exogenous feeding and complete resorption of the yolk sac.

Parasitic Copepod (Lernaea cyprinacea) Outbreaks in Foothill Yellow-legged Frogs (Rana boylii) Linked to Unusually Warm Summers and Amphibian Malformations in Northern California

Abstract How climate change may affect parasite–host assemblages and emerging infectious diseases is an important question in amphibian decline research. We present data supporting a link between periods of unusually warm summer water temperatures during 2006 and 2008 in a northern California river, outbreaks of the parasitic copepod Lernaea cyprinacea, and malformations in tadpoles and young of the year Foothill Yellow-legged Frogs (Rana boylii). Relative to baseline data gathered since 1989, both 2006 and 2008 had significantly longer periods when daily mean water temperatures exceeded 20°C compared to years without copepod outbreaks. Infestation varied spatially in the watershed, as prevalence increased concomitantly with temperature along a 5.2 km longitudinal transect. At breeding sites of R. boylii with copepods in 2006, infestation ranged from 2.9% of individuals upstream to 58.3% downstream. In 2008, copepods were absent from the most upstream sites and infested up to 28.6% of individuals sampled at downstream locations. Copepods most frequently embedded near a hind limb or the cloaca. Among individuals with parasites in 2006, 26.5% had morphological abnormalities compared to 1.1% of un-infested individuals. In 2008 when the infestation peak occurred late in development (post Gosner stage 39), abnormalities were not associated with copepod infestation. In both years, recently metamorphosed frogs with copepods were, on average, slightly smaller than those not infested. These occurrences represent a sudden increase in local prevalence atypical for this river ecosystem. Previously we had only once seen copepods on amphibians (on non-native Bullfrogs, Rana catesbeiana), six km further downstream. Pacific Chorus Frogs, Pseudacris regilla, which co-occur with R. boylii in shallow near shore habitats were not used as hosts. The data suggest that increasing summer water temperatures, decreased daily discharge, or a combination of both, promote outbreaks of this non-native parasite on an indigenous host, and could present a threat to the long-term conservation of R. boylii under the flow regime scenarios predicted by climate change models.

Water Velocity Tolerance in Tadpoles of the Foothill Yellow-legged Frog (Rana boylii): Swimming Performance, Growth, and Survival

We explored the effects of large magnitude flow fluctuations in rivers with dams, commonly referred to as pulsed flows, on tadpoles of the lotic-breeding Foothill Yellow-legged Frog, Rana boylii. We quantified the velocity conditions in habitats occupied by tadpoles and then conducted experiments to assess the tolerance to values at the upper limit of, and outside, the natural range. In laboratory flumes and field enclosures we mimicked the velocities observed during pulsed flows. In all experimental venues, the behavioral response of tadpoles was to seek refuge in the channel substrate when velocity increased. In a large laboratory flume, tadpoles moved freely at low water velocities (0–2 cm•s−1) and then sheltered among rocks when velocity increased. In a smaller scale laboratory flume, the median critical velocity was 20.1 cm•s−1. Critical velocity varied inversely with tadpole size, developmental stage, and proportion of time spent swimming. Velocities as low as 10 cm•s−1 caused tadpoles approaching metamorphosis to be displaced. In field mesocosm experiments, tadpoles exposed to repeated sub-critical velocity stress (5–10 cm•s−1) grew significantly less and experienced greater predation than tadpoles reared at ambient velocities. Responses to velocity manipulations were consistent among tadpoles from geographically distinct populations representing the three identified clades within R. boylii. The velocities associated with negative effects in these trials are less than typical velocity increases in near shore habitats when recreational flows for white water boating or peaking releases for hydroelectric power generation occur.

High Time for Conservation: Adding the Environment to the Debate on Marijuana Liberalization

The liberalization of marijuana policies, including the legalization of medical and recreational marijuana, is sweeping the United States and other countries. Marijuana cultivation can have significant negative collateral effects on the environment that are often unknown or overlooked. Focusing on the state of California, where by some estimates 60%–70% of the marijuana consumed in the United States is grown, we argue that (a) the environmental harm caused by marijuana cultivation merits a direct policy response, (b) current approaches to governing the environmental effects are inadequate, and (c) neglecting discussion of the environmental impacts of cultivation when shaping future marijuana use and possession policies represents a missed opportunity to reduce, regulate, and mitigate environmental harm.

Patterns of Freshwater Species Richness, Endemism, and Vulnerability in California.

The ranges and abundances of species that depend on freshwater habitats are declining worldwide. Efforts to counteract those trends are often hampered by a lack of information about species distribution and conservation status and are often strongly biased toward a few well-studied groups. We identified the 3,906 vascular plants, macroinvertebrates, and vertebrates native to California, USA, that depend on fresh water for at least one stage of their life history. We evaluated the conservation status for these taxa using existing government and non-governmental organization assessments (e.g., endangered species act, NatureServe), created a spatial database of locality observations or distribution information from ~400 data sources, and mapped patterns of richness, endemism, and vulnerability. Although nearly half of all taxa with conservation status (n = 1,939) are vulnerable to extinction, only 114 (6%) of those vulnerable taxa have a legal mandate for protection in the form of formal inclusion on a state or federal endangered species list. Endemic taxa are at greater risk than non-endemics, with 90% of the 927 endemic taxa vulnerable to extinction. Records with spatial data were available for a total of 2,276 species (61%). The patterns of species richness differ depending on the taxonomic group analyzed, but are similar across taxonomic level. No particular taxonomic group represents an umbrella for all species, but hotspots of high richness for listed species cover 40% of the hotspots for all other species and 58% of the hotspots for vulnerable freshwater species. By mapping freshwater species hotspots we show locations that represent the top priority for conservation action in the state. This study identifies opportunities to fill gaps in the evaluation of conservation status for freshwater taxa in California, to address the lack of occurrence information for nearly 40% of freshwater taxa and nearly 40% of watersheds in the state, and to implement adequate protections for freshwater taxa where they are currently lacking.