Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park

Grizzly bears (Ursus arctos) and American black bears (U. americanus) are sympatric in much of Yellowstone National Park. Three primary bear foods, cutthroat trout (Oncorhynchus clarki), whitebark pine (Pinus albicaulis) nuts, and elk (Cervus elaphus), have declined in recent years. Because park managers and the public are concerned about the impact created by reductions in these foods, we quantified bear diets to determine how bears living near Yellowstone Lake are adjusting. We estimated diets using: 1) stable isotope and mercury analyses of hair samples collected from captured bears and from hair collection sites established along cutthroat trout spawning streams and 2) visits to recent locations occupied by bears wearing Global Positioning System collars to identify signs of feeding behavior and to collect scats for macroscopic identification of residues. Approximately 45 ± 22% ( ± SD) of the assimilated nitrogen consumed by male grizzly bears, 38 ± 20% by female grizzly bears, and 23 ± 7% by male and female black bears came from animal matter. These assimilated dietary proportions for female grizzly bears were the same as 10 years earlier in the Lake area and 30 years earlier in the Greater Yellowstone Ecosystem. However, the proportion of meat in the assimilated diet of male grizzly bears decreased over both time frames. The estimated biomass of cutthroat trout consumed by grizzly bears and black bears declined 70% and 95%, respectively, in the decade between 1997–2000 and 2007–2009. Grizzly bears killed an elk calf every 4.3 ± 2.7 days and black bears every 8.0 ± 4.0 days during June. Elk accounted for 84% of all ungulates consumed by both bear species. Whitebark pine nuts continue to be a primary food source for both grizzly bears and black bears when abundant, but are replaced by false-truffles (Rhizopogon spp.) in the diets of female grizzly bears and black bears when nut crops are minimal. Thus, both grizzly bears and black bears continue to adjust to changing resources, with larger grizzly bears continuing to occupy a more carnivorous niche than the smaller, more herbivorous black bear. © 2012 The Wildlife Society.     

Environmental stress and developmental stability in dentition of the Yellowstone grizzly bears

Asymmetry in bilateral traits is often used to assess an individual'squality and stress resistance, but stress-induced variationin developmental stability is largely undocumented for free-livingpopulations. Over many years, grizzly bears (Ursus arctos horribilis)extensively foraged around garbage dumps in Yellowstone NationalPark. Abrupt closure of these dumps 26 years ago was a severelystressful event and was followed by excessive mortality anda many-fold increase in grizzly home-range size. I examine howthis stress affected developmental stability by comparing dentitionof bears born before and after the dump closure. I predictedthat (1) asymmetry in dentition should be greater in bears bornafter dump closure compared to before closure, and asymmetryin sexually selected canines should change more than nonsexuallyselected premolars following dump closure and (2) the relationshipbetween tooth asymmetry and tooth size should change in thepopulations following the stressful events as compared withpopulations before stressful events. I found that developmentalstability of canines, which are under directional sexual selectionin males, was more responsive to stress compared to that ofmale premolars or female dentition (both under stabilizing selection),and, because of the increased cost of canine production, feweranimals were able to achieve both large size and symmetricaldevelopment of these teeth, and thus the slope of the relationshipbetween fluctuating asymmetry and canine size increased. I concludethat stress appears to act as an honesty-reinforcement mechanismin sexual selection for symmetrical dentition.     

Multiple estimates of effective population size for monitoring a long‐lived vertebrate: an application to Yellowstone grizzly bears

Effective population size (N_e) is a key parameter for monitoring the genetic health of threatened populations because it reflects a population's evolutionary potential and risk of extinction due to genetic stochasticity. However, its application to wildlife monitoring has been limited because it is difficult to measure in natural populations. The isolated and well‐studied population of grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem provides a rare opportunity to examine the usefulness of different N_e estimators for monitoring. We genotyped 729 Yellowstone grizzly bears using 20 microsatellites and applied three single‐sample estimators to examine contemporary trends in generation interval (GI), effective number of breeders (N_b) and N_e during 1982–2007. We also used multisample methods to estimate variance (N_eV) and inbreeding N_e (N_eI). Single‐sample estimates revealed positive trajectories, with over a fourfold increase in N_e (≈100 to 450) and near doubling of the GI (≈8 to 14) from the 1980s to 2000s. N_eV (240–319) and N_eI (256) were comparable with the harmonic mean single‐sample N_e (213) over the time period. Reanalysing historical data, we found N_eV increased from ≈80 in the 1910s–1960s to ≈280 in the contemporary population. The estimated ratio of effective to total census size (N_e/N_c) was stable and high (0.42–0.66) compared to previous brown bear studies. These results support independent demographic evidence for Yellowstone grizzly bear population growth since the 1980s. They further demonstrate how genetic monitoring of N_e can complement demographic‐based monitoring of N_c and vital rates, providing a valuable tool for wildlife managers.     

Influence of whitebark pine decline on fall habitat use and movements of grizzly bears in the Greater Yellowstone Ecosystem

When abundant, seeds of the high‐elevation whitebark pine (WBP; Pinus albicaulis) are an important fall food for grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem. Rates of bear mortality and bear/human conflicts have been inversely associated with WBP productivity. Recently, mountain pine beetles (Dendroctonus ponderosae) have killed many cone‐producing WBP trees. We used fall (15 August–30 September) Global Positioning System locations from 89 bear years to investigate temporal changes in habitat use and movements during 2000–2011. We calculated Manly–Chesson (MC) indices for selectivity of WBP habitat and secure habitat (≥500 m from roads and human developments), determined dates of WBP use, and documented net daily movement distances and activity radii. To evaluate temporal trends, we used regression, model selection, and candidate model sets consisting of annual WBP production, sex, and year. One‐third of sampled grizzly bears had fall ranges with little or no mapped WBP habitat. Most other bears (72%) had a MC index above 0.5, indicating selection for WBP habitats. From 2000 to 2011, mean MC index decreased and median date of WBP use shifted about 1 week later. We detected no trends in movement indices over time. Outside of national parks, there was no correlation between the MC indices for WBP habitat and secure habitat, and most bears (78%) selected for secure habitat. Nonetheless, mean MC index for secure habitat decreased over the study period during years of good WBP productivity. The wide diet breadth and foraging plasticity of grizzly bears likely allowed them to adjust to declining WBP. Bears reduced use of WBP stands without increasing movement rates, suggesting they obtained alternative fall foods within their local surroundings. However, the reduction in mortality risk historically associated with use of secure, high‐elevation WBP habitat may be diminishing for bears residing in multiple‐use areas.     

Contrafreeloading in grizzly bears: implications for captive foraging enrichment

Although traditional feeding regimens for captive animals were focused on meeting physiological needs to assure good health, more recently emphasis has also been placed on non‐nutritive aspects of feeding. The provision of foraging materials to diversify feeding behavior is a common practice in zoos but selective consumption of foraging enrichment items over more balanced “chow” diets could lead to nutrient imbalance. One alternative is to provide balanced diets in a contrafreeloading paradigm. Contrafreeloading occurs when animals choose resources that require effort to exploit when identical resources are freely available. To investigate contrafreeloading and its potential as a theoretical foundation for foraging enrichment, we conducted two experiments with captive grizzly bears (Ursus arctos horribilis). In Experiment 1, bears were presented with five foraging choices simultaneously: apples, apples in ice, salmon, salmon in ice, and plain ice under two levels of food restriction. Two measures of contrafreeloading were considered: weight of earned food consumed and time spent working for earned food. More free than earned food was eaten, with only two bears consuming food extracted from ice, but all bears spent more time manipulating ice containing salmon or apples than plain ice regardless of level of food restriction. In Experiment 2, food‐restricted bears were presented with three foraging choices simultaneously: apples, apples inside a box, and an empty box. Although they ate more free than earned food, five bears consumed food from boxes and all spent more time manipulating boxes containing apples than empty boxes. Our findings support the provision of contrafreeloading opportunities as a foraging enrichment strategy for captive wildlife. Zoo Biol 29:484–502, 2010. © 2009 Wiley‐Liss, Inc.     

Prevalence of Trichinella spp. in black bears, grizzly bears, and wolves in the Dehcho Region, Northwest Territories, Canada, including the first report of T. nativa in a grizzly bear from Canada

Samples of muscle from 120 black bears (Ursus americanus), 11 grizzly bears (Ursus arctos), and 27 wolves (Canis lupus) collected in the Dehcho Region of the Northwest Territories from 2001 to 2010 were examined for the presence of Trichinella spp. larvae using a pepsin-HCl digestion assay. Trichinella spp. larvae were found in eight of 11 (73%) grizzly bears, 14 of 27 (52%) wolves, and seven of 120 (5.8%) black bears. The average age of positive grizzly bears, black bears, and wolves was 13.5, 9.9, and approximately 4 yr, respectively. Larvae from 11 wolves, six black bears, and seven grizzly bears were genotyped. Six wolves were infected with T. nativa and five with Trichinella T6, four black bears were infected with T. nativa and two with Trichinella T6, and all seven grizzly bears were infected with Trichinella T6 and one of them had a coinfection with T. nativa. This is the first report of T. nativa in a grizzly bear from Canada. Bears have been linked to trichinellosis outbreaks in humans in Canada, and black bears are a subsistence food source for residents of the Dehcho region. In order to assess food safety risk it is important to monitor the prevalence of Trichinella spp. in both species of bear and their cohabiting mammalian food sources.     

Fast carnivores and slow herbivores: differential foraging strategies among grizzly bears in the Canadian Arctic

Categorizing animal populations by diet can mask important intrapopulation variation, which is crucial to understanding a species’ trophic niche width. To test hypotheses related to intrapopulation variation in foraging or the presence of diet specialization, we conducted stable isotope analysis (δ¹³C, δ¹⁵N) on hair and claw samples from 51 grizzly bears (Ursus arctos) collected from 2003 to 2006 in the Mackenzie Delta region of the Canadian Arctic. We examined within-population differences in the foraging patterns of males and females and the relationship between trophic position (derived from δ¹⁵N measurements) and individual movement. The range of δ¹⁵N values in hair and claw (2.0–11.0‰) suggested a wide niche width and cluster analyses indicated the presence of three foraging groups within the population, ranging from near-complete herbivory to near-complete carnivory. We found no linear relationship between home range size and trophic position when the data were continuous or when grouped by foraging behavior. However, the movement rate of females increased linearly with trophic position. We used multisource dual-isotope mixing models to determine the relative contributions of seven prey sources within each foraging group for both males and females. The mean bear dietary endpoint across all foraging groups for each sex fell toward the center of the mixing polygon, which suggested relatively well-mixed diets. The primary dietary difference across foraging groups was the proportional contribution of herbaceous foods, which decreased for both males and females from 42–76 to 0–27% and 62–81 to 0–44%, respectively. Grizzlies of the Mackenzie Delta live in extremely harsh conditions and identifying within-population diet specialization has improved our understanding of varying habitat requirements within the population.     

Population fragmentation and inter-ecosystem movements of grizzly bears in western Canada and the northern United States

Population fragmentation compromises population viability, reduces a species ability to respond to climate change, and ultimately may reduce biodiversity. We studied the current state and potential causes of fragmentation in grizzly bears over approximately 1,000,000 km² of western Canada, the northern United States (US), and southeast Alaska. We compiled much of our data from projects undertaken with a variety of research objectives including population estimation and trend, landscape fragmentation, habitat selection, vital rates, and response to human development. Our primary analytical techniques stemmed from genetic analysis of 3,134 bears, supplemented with radiotelemetry data from 792 bears. We used 15 locus microsatellite data coupled with measures of genetic distance, isolation-by-distance (IBD) analysis, analysis of covariance (ANCOVA), linear multiple regression, multi-factorial correspondence analysis (to identify population divisions or fractures with no a priori assumption of group membership), and population-assignment methods to detect individual migrants between immediately adjacent areas. These data corroborated observations of inter-area movements from our telemetry database. In northern areas, we found a spatial genetic pattern of IBD, although there was evidence of natural fragmentation from the rugged heavily glaciated coast mountains of British Columbia (BC) and the Yukon. These results contrasted with the spatial pattern of fragmentation in more southern parts of their distribution. Near the Canada–US border area, we found extensive fragmentation that corresponded to settled mountain valleys and major highways. Genetic distances across developed valleys were elevated relative to those across undeveloped valleys in central and northern BC. In disturbed areas, most inter-area movements detected were made by male bears, with few female migrants identified. North–south movements within mountain ranges (Mts) and across BC Highway 3 were more common than east–west movements across settled mountain valleys separating Mts. Our results suggest that relatively distinct subpopulations exist in this region, including the Cabinet, Selkirk South, and the decades-isolated Yellowstone populations. Current movement rates do not appear sufficient to consider the subpopulations we identify along the Canada–US border as 1 inter-breeding unit. Although we detected enough male movement to mediate gene flow, the current low rate of female movement detected among areas is insufficient to provide a demographic rescue effect between areas in the immediate future (0–15 yr). In Alberta, we found fragmentation corresponded to major east–west highways (Highways 3, 11, 16, and 43) and most inter-area movements were made by males. Gene flow and movement rates between Alberta and BC were highest across the Continental Divide south of Highway 1 and north of Highway 16. In the central region between Highways 1 and 11, we found evidence of natural fragmentation associated with the extensive glaciers and icefields along the Continental Divide. The discontinuities that we identified would form appropriate boundaries for management units. We related sex-specific movement rates between adjacent areas to several metrics of human use (highway traffic, settlement, and human-caused mortality) to understand the causes of fragmentation. This analysis used data from 1,508 bears sampled over a 161,500-km² area in southeastern BC, western Alberta, northern Idaho, and northern Montana during 1979–2007. This area was bisected by numerous human transportation and settlement corridors of varying intensity and complexity. We used multiple linear regression and ANCOVA to document the responses of female and male bears to disturbance. Males and females both demonstrated reduced movement rates with increasing settlement and traffic. However, females reduced their movement rates dramatically when settlement increased to >20% of the fracture zone. At this same threshold, male movement declined more gradually, in response to increased traffic and further settlement. In highly settled areas (>50%), both sexes had a similar reduction in movements in response to traffic, settlement, and mortality. We documented several small bear populations with male-only immigration, highlighting the importance of investigating sex-specific movements. Without female connectivity, small populations are not viable over the long term. The persistence of this regional female fragmented metapopulation likely will require strategic connectivity management. We therefore recommend enhancing female connectivity among fractured areas by securing linkage-zone habitat appropriate for female dispersal, and ensuring current large source subpopulations remain intact. The fragmentation we documented may also affect other species with similar ecological characteristics: sparse densities, slow reproduction, short male-biased dispersal, and a susceptibility to human-caused mortality and habitat degradation. Therefore, regional inter-jurisdictional efforts to manage broad landscapes for inter-area movement will likely benefit a broad spectrum of species and natural processes, particularly in light of climate change. © 2011 The Wildlife Society.     

Carnivore taphonomy in South America: a review of actualistic studies and their implications in the southern Neotropics

Abstract

Actualistic studies on mammalian carnivore taphonomy in southern South America are reviewed here, including pumas, small cats, foxes, and other, smaller carnivores. Patterns for different carnivore taxa and their variation are elicited. Also temporal and spatial variability is analysed, and comparisons are made to other carnivores and regions. While generally these Neotropical carnivores produce low damage intensity, stronger modifications and their implications are described as well. These patterns and their variation are put in context by taking into account the particular physical and biotic conditions in the southern Neotropics.     

Serologic survey for infectious canine hepatitis virus in grizzly bears (Ursus arctos) from Alaska, 1973 to 1987

Serum antibody prevalence of infectious canine hepatitis virus was 12% (90 of 725) for grizzly bears (Ursus arctos) from Alaska (USA) during the period 1973 to 1987. Prevalence was highest on Kodiak Island at 29% (37 of 127). Prevalence of exposure at individual collection areas did not change significantly over time. There were no significant sex-specific differences in prevalence. Prevalence was directly related to age, but it was 0% for bears less than 2-yr-old. Young bears which are exposed to the virus may develop clinical disease and die as a result of the infection. This disease may be a factor affecting grizzly bear population dynamics.     

Non-random patterns in the Yellowstone ecosystem: inferences from mammalian body size, order and biogeographical affinity

Aim Our aim was to investigate how the environment, species characteristics and historical factors at the subcontinental scale affect patterns of diversity. We used the assembly of the Yellowstone biota over the past 10,000 years as a natural experiment for investigating the processes that generate a modern non‐volant mammal species pool. Location The data represent species from throughout North America with special attention to the non‐volant mammals of Yellowstone National Park, USA. Methods We used digitized range maps to determine biogeographical affinity for all non‐volant mammals in the Rocky Mountains, Deserts and Great Plains biogeographical regions of North America. This biogeographical affinity, along with taxonomic order and body size class, was used to test whether non‐random patterns exist in the assemblage of Yellowstone non‐volant mammals. These characteristics were also used to investigate the strength of non‐random processes, such as habitat or taxon filtering, on particular groups of species or individual species. Results Our results indicated that the Yellowstone fauna is composed of a non‐random subset of mammals from specific body size classes and with particular biogeographical affinities. Analyses by taxonomic order found significantly more Carnivora from the Rocky Mountains region and significantly fewer Rodentia from the Deserts region than expected from random assembly. Analyses using body size classes revealed deviations from expectations, including several significant differences between the frequency distribution of regional body sizes and the distribution of those species found within Yellowstone. Main conclusions Our novel approach explores processes affecting species pool assembly in the Yellowstone region and elsewhere, and particularly identifies unique properties of species that may contribute to non‐random assembly. Focusing on the mechanisms generating diversity, not just current diversity patterns, will assist the design of conservation strategies given future environmental change scenarios.     

Geographic pattern of serum antibody prevalence for Brucella spp. in caribou, grizzly bears, and wolves from Alaska, 1975-1998

Blood samples were collected from 2,635 caribou (Rangifer tarandus), 1,238 grizzly bears (Ursus arctos), and 930 wolves (Canis lupus) from throughout mainland Alaska during 1975-98. Sera were tested for evidence of exposure to Brucella spp. Serum antibody prevalences were highest in the northwestern region of the state. In any specific area, antibody prevalences for caribou and wolves were of a similar magnitude, whereas antibody prevalence for bears in these same areas were two to three times higher.     

Dietary protein content alters energy expenditure and composition of the mass gain in grizzly bears (Ursus arctos horribilis)

Many fruits contain high levels of available energy but very low levels of protein and other nutrients. The discrepancy between available energy and protein creates a physiological paradox for many animals consuming high-fruit diets, as they will be protein deficient if they eat to meet their minimum energy requirement. We fed young grizzly bears both high-energy pelleted and fruit diets containing from 1.6% to 15.4% protein to examine the role of diet-induced thermogenesis and fat synthesis in dealing with high-energy-low-protein diets. Digestible energy intake at mass maintenance increased 2.1 times, and composition of the gain changed from primarily lean mass to entirely fat when the protein content of the diet decreased from 15.4% to 1.6%. Daily fat gain was up to three times higher in bears fed low-protein diets ad lib., compared with bears consuming the higher-protein diet and gaining mass at the same rate. Thus, bears eating fruit can either consume other foods to increase dietary protein content and reduce energy expenditure, intake, and potentially foraging time or overeat high-fruit diets and use diet-induced thermogenesis and fat synthesis to deal with their skewed energy-to-protein ratio. These are not discrete options but a continuum that creates numerous solutions for balancing energy expenditure, intake, foraging time, fat accumulation, and ultimately fitness, depending on food availability, foraging efficiency, bear size, and body condition.     

Genetic relationships of grizzly bears (Ursus arctos) in the Prudhoe Bay region of Alaska: inference from microsatellite DNA, mitochondrial DNA, and field observations

Grizzly bears are abundant in the region of the Prudhoe Bay oil fields in northern Alaska. We used field observations and molecular genetic data to identify parent-offspring and sibling relationships among bears in this region. We determined genotypes at 14 microsatellite DNA loci and the cytochrome b gene of mitochondrial DNA (mtDNA) for 36 bears. We identified 17 possible mother-offspring pairs and 8 possible father-offspring pairs. This includes verification of the relationships of 14 mother-offspring pairs identified from field observations. Three additional mother-offspring pairs and all eight father-offspring pairs were determined from genetic and age data. Relatedness coefficients based on numbers of shared alleles between individuals were as expected: approximately 0.50 for parent-offspring and sibling pairs and approximately 0.75 for a father-offspring pair resulting from a father-daughter mating. The level of genetic variation (mean number of alleles per locus = 6.6, mean heterozygosity = 70%) and allele frequencies in grizzly bears in the Prudhoe Bay region are similar to those in other parts of the species' range.     

Herbivore influence on soil microbial biomass and nitrogen mineralization in a northern grassland ecosystem: Yellowstone National Park

Microorganisms are largely responsible for soil nutrient cycling and energy flow in terrestrial ecosystems. Although soil microorganisms are affected by topography and grazing, little is known about how these two variables may interact to influence microbial processes. Even less is known about how these variables influence microorganisms in systems that contain large populations of free-roaming ungulates. In this study, we compared microbial biomass size and activity, as measured by in situ net N mineralization, inside and outside 35- to 40-year exclosures across a topographic gradient in northern Yellowstone National Park. The objective was to determine the relative effect of topography and large grazers on microbial biomass and nitrogen mineralization. Microbial C and N varied by almost an order of magnitude across sites. Topographic depressions that contained high plant biomass and fine-textured soils supported the greatest microbial biomass. We found that plant biomass accurately predicted microbial biomass across our sites suggesting that carbon inputs from plants constrained microbial biomass. Chronic grazing neither depleted soil C nor reduced microbial biomass. We hypothesize that microbial populations in grazed grasslands are sustained mainly by inputs of labile C from dung deposition and increased root turnover or root exudation beneath grazed plants. Mineral N fluxes were affected more by grazing than topography. Net N mineralization rates were highest in grazed grassland and increased from dry, unproductive to mesic, highly productive communities. Overall, our results indicate that topography mainly influences microbial biomass size, while mineral N fluxes (microbial activity) are affected more by grazing in this grassland ecosystem. Received: 4 June 1997 / Accepted: 16 December 1997     

Future sea ice conditions in Western Hudson Bay and consequences for polar bears in the 21st century

The primary habitat of polar bears is sea ice, but in Western Hudson Bay (WH), the seasonal ice cycle forces polar bears ashore each summer. Survival of bears on land in WH is correlated with breakup and the ice‐free season length, and studies suggest that exceeding thresholds in these variables will lead to large declines in the WH population. To estimate when anthropogenic warming may have progressed sufficiently to threaten the persistence of polar bears in WH, we predict changes in the ice cycle and the sea ice concentration (SIC) in spring (the primary feeding period of polar bears) with a high‐resolution sea ice‐ocean model and warming forced with 21st century IPCC greenhouse gas (GHG) emission scenarios: B1 (low), A1B (medium), and A2 (high). We define critical years for polar bears based on proposed thresholds in breakup and ice‐free season and we assess when ice‐cycle conditions cross these thresholds. In the three scenarios, critical years occur more commonly after 2050. From 2001 to 2050, 2 critical years occur under B1 and A2, and 4 under A1B; from 2051 to 2100, 8 critical years occur under B1, 35 under A1B and 41 under A2. Spring SIC in WH is high (>90%) in all three scenarios between 2001 and 2050, but declines rapidly after 2050 in A1B and A2. From 2090 to 2100, the mean spring SIC is 84 (±7)% in B1, 56 (±26)% in A1B and 20 (±13)% in A2. Our predictions suggest that the habitat of polar bears in WH will deteriorate in the 21st century. Ice predictions in A1B and A2 suggest that the polar bear population may struggle to persist after ca. 2050. Predictions under B1 suggest that reducing GHG emissions could allow polar bears to persist in WH throughout the 21st century.     

Anticipative management for coral reef ecosystem services in the 21st century

Under projections of global climate change and other stressors, significant changes in the ecology, structure and function of coral reefs are predicted. Current management strategies tend to look to the past to set goals, focusing on halting declines and restoring baseline conditions. Here, we explore a complementary approach to decision making that is based on the anticipation of future changes in ecosystem state, function and services. Reviewing the existing literature and utilizing a scenario planning approach, we explore how the structure of coral reef communities might change in the future in response to global climate change and overfishing. We incorporate uncertainties in our predictions by considering heterogeneity in reef types in relation to structural complexity and primary productivity. We examine 14 ecosystem services provided by reefs, and rate their sensitivity to a range of future scenarios and management options. Our predictions suggest that the efficacy of management is highly dependent on biophysical characteristics and reef state. Reserves are currently widely used and are predicted to remain effective for reefs with high structural complexity. However, when complexity is lost, maximizing service provision requires a broader portfolio of management approaches, including the provision of artificial complexity, coral restoration, fish aggregation devices and herbivore management. Increased use of such management tools will require capacity building and technique refinement and we therefore conclude that diversification of our management toolbox should be considered urgently to prepare for the challenges of managing reefs into the 21st century.