Coupled social and ecological change drove the historical extinction of the California grizzly bear (Ursus arctos californicus).

Large carnivores (order Carnivora) are among the world's most threatened mammals due to a confluence of ecological and social forces that have unfolded over centuries. Combining specimens from natural history collections with documents from archival records, we reconstructed the factors surrounding the extinction of the California grizzly bear (Ursus arctos californicus), a once-abundant brown bear subspecies last seen in 1924. Historical documents portrayed California grizzlies as massive hypercarnivores that endangered public safety. Yet, morphological measurements on skulls and teeth generate smaller body size estimates in alignment with extant North American grizzly populations (approx. 200 kg). Stable isotope analysis (δ13C, δ15N) of pelts and bones (n = 57) revealed that grizzlies derived less than 10% of their nutrition from terrestrial animal sources and were therefore largely herbivorous for millennia prior to the first European arrival in this region in 1542. Later colonial land uses, beginning in 1769 with the Mission era, led grizzlies to moderately increase animal protein consumption (up to 26% of diet), but grizzlies still consumed far less livestock than otherwise claimed by contemporary accounts. We show how human activities can provoke short-term behavioural shifts, such as heightened levels of carnivory, that in turn can lead to exaggerated predation narratives and incentivize persecution, triggering rapid loss of an otherwise widespread and ecologically flexible animal.

Small mammal responses to fire severity mediated by vegetation characteristics and species traits.

The frequency of large, high-severity "mega-fires" has increased in recent decades, with numerous consequences for forest ecosystems. In particular, small mammal communities are vulnerable to post-fire shifts in resource availability and play critical roles in forest ecosystems. Inconsistencies in previous observations of small mammal community responses to fire severity underscore the importance of examining mechanisms regulating the effects of fire severity on post-fire recovery of small mammal communities. We compared small mammal abundance, diversity, and community structure among habitats that burned at different severities, and used vegetation characteristics and small mammal functional traits to predict community responses to fire severity three years after one mega-fire in the Sierra Nevada, California. Using a model-based fourth-corner analysis, we examined how interactions between vegetation variables and small mammal traits associated with their resource use were associated with post-fire small mammal community structure among fire severity categories. Small mammal abundance was similar across fire severity categories, but diversity decreased and community structure shifted as fire severity increased. Differences in small mammal communities were large only between unburned and high-severity sites. Three highly correlated fire-dependent vegetation variables affected by fire and the volume of soft coarse woody debris were associated with small mammal community structures. Furthermore, we found that interactions between vegetation variables and three small mammal traits (feeding guild, primary foraging mode, and primary nesting habit) predicted community structure across fire severity categories. We concluded that resource use was important in regulating small mammal recovery after the fire because vegetation provided required resources to small mammals as determined by their functional traits. Given the mechanistic nature of our analyses, these results may be applicable to other fire-prone forest systems, although it will be important to conduct studies across large biogeographic regions and over long post-fire time periods to assess generality.

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers.

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.

Analogies for a No-Analog World: Tackling Uncertainties in Reintroduction Planning.

Species reintroductions involve considerable uncertainty, especially in highly altered landscapes. Historical, geographic, and taxonomic analogies can help reduce this uncertainty by enabling conservationists to better assess habitat suitability in proposed reintroduction sites. We illustrate this approach using the example of the California grizzly, an iconic species proposed for reintroduction.

Aquatic macroinvertebrates stabilize gravel bed sediment: A test using silk net-spinning caddisflies in semi-natural river channels.

Organisms can have large effects on the physical properties of the habitats where they live. For example, measurements in laboratory stream microcosms have shown that the presence of silk net-spinning insect larvae (Trichoptera: Hydropsychidae) can increase the shear force required to initiate movement of riverbed sediments. Few studies, however, have moved beyond laboratory settings to quantify the engineering impacts of aquatic insects under more complex field conditions. To bridge the gap between small-scale laboratory experiments and natural stream ecosystems, we conducted experiments in large (50 m2) outdoor river channels where net-spinning aquatic insects were manipulated in sediment patches that were 5 to 25 times larger than in previous studies. We tested whether larvae of two caddisfly species (Arctopsyche californica and Ceratopsyche oslari) influenced the stability of gravel during simulated floods when alone in monoculture and together in polyculture. On average, populations of caddisflies increased the critical shear stress required to initiate sediment movement by 20% compared to treatments without caddisflies. Per capita effects of caddisflies on sediment stability were similar between previous laboratory studies and this field experiment, and Arctopsyche had a larger per capita effect than Ceratopsyche, perhaps because of its larger size and stronger silk. Contrary to prior laboratory flume results, the effects of the two species on critical shear stress when together were similar to the additive expectation of both species when alone, but effects of the two species together were higher than the additive expectation when we accounted for density. Comparisons of total population and per capita effects suggest that caddisfly density, identity, and coexisting species likely have effects on the magnitude of caddisfly impacts on critical shear stress. Our findings imply that consideration of both the abundances and traits of ecosystem engineers is needed to describe and model their effects on sediment mobility.

Complex impacts of an invasive omnivore and native consumers on stream communities in California and Hawaii.

The effects of invasive species on native communities often depend on the characteristics of the recipient community and on the food habits of the invasive species, becoming complicated when the invader is omnivorous. In field enclosure experiments, we assessed the direct and interactive effects of an invasive omnivorous crayfish (Procambarus clarkii) and either native herbivorous snails (Physella gyrina) or shrimp (Atyoida bisulcata) on stream communities in California and Hawaii, respectively. Based on literature data and the characteristics of each study site, we predicted that crayfish would affect primarily algal-based trophic linkages in an open California stream but detritus-based trophic linkages in a shaded Hawaiian stream, with trophic cascades mediated through crayfish effects on primary consumers being observed in both systems. As predicted, crayfish in California directly reduced periphyton, filamentous algae, sediment, and snail levels, but generated a cascade by decreasing snail densities and increasing periphyton biomass. Contrary to prediction, crayfish did not reduce total invertebrate biomass. As predicted, crayfish in Hawaii reduced leaf litter, filamentous algae, and benthic invertebrate biomass. Contrary to our predictions, however, a trophic cascade was not observed because shrimp did not affect periphyton levels, crayfish did not reduce shrimp abundance, and crayfish had greater negative impacts on filamentous algae than did shrimp. Our findings highlight that the same invasive species can generate different effects on disparate systems, probably as mediated through the availability of different food types, flexibility in the invasive species' food habits, and complex pathways of trophic interaction.

Scaling population responses to spatial environmental variability in advection-dominated systems.

We model the spatial dynamics of an open population of organisms that disperse solely through advection in order to understand responses to multiscale environmental variability. We show that the distance over which a population responds to a localized perturbation, called the response length, can be characterized as an organisms average lifetime dispersal distance, unless there is strong density-dependence in demographic or dispersal rates. Continuous spatial fluctuations in demographic rates at scales smaller than the response length will be largely averaged in the population distribution, whereas those in per capita emigration rates will be strongly tracked. We illustrate these results using a parameterized example to show how responses to environmental variability may differ in streams with different average current velocities. Our model suggests an approach to linking local dynamics dominated by dispersal processes to larger-scale dynamics dominated by births and deaths.

Effects of trout on the diel periodicity of drifting in baetid mayflies.

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.

Effects of an invertebrate grazer on the spatial arrangement of a benthic microhabitat.

We demonstrated the effect of an aquatic herbivore on the spatial arrangement of benthic algal biomass within artificial stream channels. Transects of ceramic tiles were exposed to a gradient of snail (Physella) densities in a 30 d experiment. We observed positive effects of snails on the mean abundance of "overstory" algae (the filamentous chlorophyte Cladophora and associated epiphytes), an important benthic microhabitat in streams. Snails altered several aspects of the spatial arrangement of overstory algae. Snails reduced the strength of downstream gradients in overstory biomass, as well as residual variability around these gradients. Geostatistical analysis revealed that snails also reduced the strength of spatial dependence, and so reduced spatial heterogeneity of the overstory, at small scales (<40 cm). As a result, organisms inhabiting the overstory might experience a more fragmented habitat landscape at high snail densities. In addition, snails increased the scale of spatial dependence in understory algal biomass (algae remaining on tiles after overstory was removed) from 10 cm to 40 cm. Consumer effects on the spatial arrangement of a microhabitat argue for the inclusion of feedbacks between the biota and the environment in spatially-explicit models.

The effects of trout on water striders in stream pools.

Observations made in Rattlesnake Creek, Santa Barbara County, California, U.S.A. indicate that rainbow trout (Salmo gairdneri) affect the microdistribution of adult water striders (Gerris remigis). Gerrids occupied all areas of stream pools lacking trout but stayed in margins of pools containing trout. To test the hypothesis that the presence of trout controlled the distribution of water striders within pools, trout were removed from some pools and transferred to pools lacking trout. Undistrubed pools with and without trout served as controls for the trout transferrals. The results supported the hypothesis; within one week, gerrids moved to the edge of pools where trout were added and to the middle of pools where trout were removed. Time budgets and gut analyses suggest that local patterns of gerrid distribution were maintained by harrassment of gerrids by trout. At times trout had significant effects on components of gerrid fitness.

The effect of physical disturbance on the relative abundances of two filter-feeding insects in a small stream.

We examined the importance of disturbance in determining the relative abundances of two lotic filter-feeders, Simulium virgatum and Hydropsyche oslari, in a small, coastal stream in southern California, USA.In most years, winter spates effectively scour substrata in fast-flowing areas, thereby drastically reducing stream insect populations. Newly-opened space in these areas is quickly colonized by simuliids. The abundance of simuliids, however, gradually declines as hydropsychid abundance increases in early summer. To determine if these changes in insect abundance represent seasonal changes or successional changes following disturbance, we performed a field experiment where hard substrates were disturbed at 2 wk, 4 wk, or 8 wk intervals, or were left undisturbed. We found that the numbers of simuliids increased and the numbers of hydropsychids decreased as the frequency of disturbance increased. Although seasonal recruitment patterns and longitudinal position in the strem had important effects on the colonization rates of these insects, time since last disturbance was a prime determinant of the relative abundances of Simulium and Hydropsyche. These results and additional observations suggest that Simulium virgatum is an opportunistic species that quickly colonizes new space, but that it is displaced by the slower-colonizing but competitively superior Hydropsyche oslari. Disturbance promotes the coexistence of these two species by preventing the attainment of a climax state where Hydropsyche monopolizes available space.