Environmental factors and occurrence of horseshoe crabs in the northcentral Gulf of Mexico.

This study provides regional-scale data on drivers of horseshoe crab (Limulus polyphemus) presence along the northcentral Gulf of Mexico coast and has implications for understanding habitat suitability for sparse horseshoe crab populations of conservation concern worldwide. To collect baseline data on the relationship between environmental factors and presence of horseshoe crabs, we surveyed four sites from the Fort Morgan peninsula of Mobile Bay, Alabama (AL) to Horn Island, Mississippi (MS). We documented number, size and sex of live animals, molts, and carcasses as metrics of horseshoe crab presence and demographics for two years. Data were compared to in situ and remotely sensed environmental attributes to assess environmental drivers of occurrence during the time of study. Overall, greater evidence of horseshoe crab presence was found at western sites (Petit Bois and Horn Islands) compared to eastern sites (Dauphin Island, Fort Morgan peninsula), mediated by a combination of distance from areas of high freshwater discharge and interannual variation in weather. Higher sex ratios also were found associated with higher occurrence, west of Mobile Bay. Land cover, particularly Bare Land and Estuarine Emergent Wetland classes that are common to western sites, was most predictive of live animal and to some extent carcass occurrence. Our findings suggest that small-scale variation in habitat quality can affect occurrence of horseshoe crabs in sparse populations where density is not a limiting factor. Data from molts and carcasses were informative to supplement live animal data and may be useful to enhance ecological assessment and support conservation and management in regions with sparse populations.

Ontogenetic resource use and trophic dynamics of endangered juvenile Tachypleus tridentatus among diversified nursery habitats in the northern Beibu Gulf, China.

Horseshoe crabs, the most well-known example of "living fossils", are iconic and ecologically important macroinvertebrates in coastal and estuarine ecosystems. Their blood is a crucial resource for manufacturing Limulus or Tachypleus amebocyte lysate to detect bacterial endotoxins or fungal contamination in drug and medical devices. An enhanced understanding of their ecological roles and trophic interactions in the food webs is fundamental to facilitate resource management for the declining populations in Asia. Foraging information of the Asian species, however, is mainly derived from preliminary, scattered reports from a limited number of study locations. In this study, resource utilization, trophic niche dynamics, and trophic interaction of the juvenile tri-spine horseshoe crab, Tachypleus tridentatus (instars 1-12, approximately 0.5-8 years old) across ontogeny was assessed in diversified nursery habitats along the northern Beibu Gulf, China, using carbon and nitrogen stable isotopes. Our results suggest that: (i) T. tridentatus are ecological generalists given the vast range of carbon isotopic values and trophic niche width estimates exhibited between multiple instar groups; (ii) juvenile T. tridentatus across most habitat types predominantly assimilated energy from a variety of basal production sources in the food web, but primarily depended on sedimentary organic matter and seagrass resource pools; (iii) ontogenetic shifts in juvenile dietary proportions were evident, with decreased reliance on sedimentary organic matter, coupled with increased reliance on benthic macroinvertebrate grazers, detritivores, and omnivores with age; and (iv) nearly all juvenile instars occupied similar trophic positions in the food web with slight shifts in trophic position present with increasing size. Our findings indicate that resource availability and ontogenetic diet shifts strongly influence horseshoe crab trophic dynamics, and age should be accounted when formulating habitat conservation measures based on resource use for Asian horseshoe crabs.

Surf and turf vision: Patterns and predictors of visual acuity in compound eye evolution.

Eyes have the flexibility to evolve to meet the ecological demands of their users. Relative to camera-type eyes, the fundamental limits of optical diffraction in arthropod compound eyes restrict the ability to resolve fine detail (visual acuity) to much lower degrees. We tested the capacity of several ecological factors to predict arthropod visual acuity, while simultaneously controlling for shared phylogenetic history. In this study, we have generated the most comprehensive review of compound eye visual acuity measurements to date, containing 385 species that span six of the major arthropod classes. An arthropod phylogeny, made custom to this database, was used to develop a phylogenetically-corrected generalized least squares (PGLS) linear model to evaluate four ecological factors predicted to underlie compound eye visual acuity: environmental light intensity, foraging strategy (predator vs. non-predator), horizontal structure of the visual scene, and environmental medium (air vs. water). To account for optical constraints on acuity related to animal size, body length was also included, but this did not show a significant effect in any of our models. Rather, the PGLS analysis revealed that the strongest predictors of compound eye acuity are described by a combination of environmental medium, foraging strategy, and environmental light intensity.

Habitat suitability maps for juvenile tri-spine horseshoe crabs in Japanese intertidal zones: A model approach using unmanned aerial vehicles and the Structure from Motion technique.

The tri-spine horseshoe crab, Tachypleus tridentatus, is a threatened species that inhabits coastal areas from South to East Asia. A Conservation management system is urgently required for managing its nursery habitats, i.e., intertidal flats, especially in Japan. Habitat suitability maps are useful in drafting conservation plans; however, they have rarely been prepared for juvenile T. tridentatus. In this study, we examined the possibility of constructing robust habitat suitability models (HSMs) for juveniles based on topographical data acquired using unmanned aerial vehicles and the Structure from Motion (UAV-SfM) technique. The distribution data of the juveniles in the Tsuyazaki and Imazu intertidal flats from 2017 to 2019 were determined. The data were divided into a training dataset for HSM construction and three test datasets for model evaluation. High accuracy digital surface models were built for each region using the UAV-SfM technique. Normalized elevation was assessed by converting the topographical models that consider the tidal range in each region, and the slope was calculated based on these models. Using the training data, HSMs of the juveniles were constructed with normalized elevation and slope as the predictor variables. The HSMs were evaluated using the test data. The results showed that HSMs exhibited acceptable discrimination performance for each region. Habitat suitability maps were built for the juveniles in each region, and the suitable areas were estimated to be approximately 6.1 ha of the total 19.5 ha in Tuyazaki, and 3.7 ha of the total 7.9 ha area in Imazu. In conclusion, our findings support the usefulness of the UAV-SfM technique in constructing HSMs for juvenile T. tridentatus. The monitoring of suitable habitat areas for the juveniles using the UAV-SfM technique is expected to reduce survey costs, as it can be conducted with fewer investigators over vast intertidal zones within a short period of time.

Air Breathing in an Exceptionally Preserved 340-Million-Year-Old Sea Scorpion.

Arachnids are the second most successful terrestrial animal group after insects [1] and were one of the first arthropod clades to successfully invade land [2]. Fossil evidence for this transition is limited, with the majority of arachnid clades first appearing in the terrestrial fossil record. Furthermore, molecular clock dating has suggested a Cambrian-Ordovician terrestrialization event for arachnids [3], some 60 Ma before their first fossils in the Silurian, although these estimates assume that arachnids evolved from a fully aquatic ancestor. Eurypterids, the sister clade to terrestrial arachnids [4-6], are known to have undergone major macroecological shifts in transitioning from marine to freshwater environments during the Devonian [7, 8]. Discoveries of apparently subaerial eurypterid trackways [9, 10] have led to the suggestion that eurypterids were even able to venture on land and possibly breathe air [11]. However, modern horseshoe crabs undertake amphibious excursions onto land to reproduce [12], rendering trace fossil evidence alone inconclusive. Here, we present details of the respiratory organs of Adelophthalmus pyrrhae sp. nov. from the Carboniferous of Montagne Noire, France [13], revealed through micro computed tomography (µ-CT) imaging. Pillar-like trabeculae on the dorsal surface of each gill lamella indicate eurypterids were capable of subaerial breathing, suggesting that book gills are the direct precursors to book lungs while vascular ancillary respiratory structures known as Kiemenplatten represent novel air-breathing structures. The discovery of air-breathing structures in eurypterids indicates that characters permitting terrestrialization accrued in the arachnid stem lineage and suggests the Cambrian-Ordovician ancestor of arachnids would also have been semi-terrestrial.

Roles of the clip domains of two protease zymogens in the coagulation cascade in horseshoe crabs.

The lipopolysaccharide (LPS)-triggered coagulation cascade in horseshoe crabs comprises three protease zymogens: prochelicerase C (proC), prochelicerase B (proB), and the proclotting enzyme (proCE). The presence of LPS results in autocatalytic activation of proC to α-chelicerase C, which, in turn, activates proB to chelicerase B, converting proCE to the clotting enzyme (CE). ProB and proCE contain an N-terminal clip domain, but the roles of these domains in this coagulation cascade remain unknown. Here, using recombinant proteins and kinetics and binding assays, we found that five basic residues in the clip domain of proB are required to maintain its LPS-binding activity and activation by α-chelicerase C. Moreover, an amino acid substitution at a potential hydrophobic cavity in proB's clip domain (V55A-proB) reduced both its LPS-binding activity and activation rate. WT proCE exhibited no LPS-binding activity, and the WT chelicerase B-mediated activation of a proCE variant with a substitution at a potential hydrophobic cavity (V53A-proCE) was ∼4-fold slower than that of WT proCE. The kcat/Km value of the interaction of WT chelicerase B with V53A-proCE was 7-fold lower than that of the WT chelicerase B-WT proCE interaction. The enzymatic activities of V55A-chelicerase B and V53A-CE against specific peptide substrates were indistinguishable from those of the corresponding WT proteases. In conclusion, the clip domain of proB recruits it to a reaction center composed of α-chelicerase C and LPS, where α-chelicerase C is ready to activate proB, leading to chelicerase B-mediated activation of proCE via its clip domain.

Physiological impacts of time in holding ponds, biomedical bleeding, and recovery on the Atlantic horseshoe crab, Limulus polyphemus.

Atlantic horseshoe crabs, Limulus polyphemus (HSC), are commercially harvested along the eastern U.S. coast and bled for hemolymph used in pharmaceutical safety testing. In South Carolina, some HSCs are held in outdoor ponds before transport to facilities where they are bled and then released to the wild. This study examined whether the time HSCs are held before bleeding, bleeding itself, or the duration of the recovery after bleeding affects HSC mortality and physiological condition. Female HSCs were collected from Coffin Point Beach, South Carolina (April 22-24, 2016), held in ponds for 2, 4, 6, or 8 weeks, then bled or held as controls. Body weights, hemocyanin concentrations, and hemocyte densities were measured prior to treatment (bled/control) and at 2, 6, and 12 days of recovery. Hemocyanin concentrations declined significantly in HSCs held in ponds for 8 weeks prior to bleeding and were excluded from further analyses. Compared to some studies, HSC mortalities were low (11%). Impacts of time in holding ponds, bleeding, and recovery from bleeding on physiological measures were assessed using 3-way fixed-effects ANOVA. While duration of recovery had main effects on physiological measures, significant interactions were also present. There was an interaction of treatment and recovery duration, with control crabs having higher hemocyte densities than bled animals at days 2 and 6 of recovery. There were two significant two-way interactions influencing hemocyanin concentration: pond time and recovery, and treatment and recovery. Our study suggests both main and synergistic effects are important when assessing the physiology and mortality of HSCs harvested for biomedical purposes.

Xiphosurid from the Tournaisian (Carboniferous) of Scotland confirms deep origin of Limuloidea.

Horseshoe crabs are archetypal marine chelicerates with a fossil record extending from the Lower Ordovician to today. The major horseshoe crab groups are thought to have arisen in the middle to late Palaeozoic. Here we present the oldest known limuloid from the lower Carboniferous (Tournaisian stage, c. 350 million years ago) of Scotland: Albalimulus bottoni gen. et sp. nov. A comprehensive phylogenetic analysis supports the placement of A. bottoni as a representative of the extant family Limulidae and 100 million years older than any other limulid taxon. The use of geometric morphometric analyses corroborate the erection of the new taxon and illustrates the exploitation of morphospace by xiphosurids. This new taxon highlights the complex evolutionary history of xiphosurids and the importance of documenting these unique Palaeozoic individuals.

Lift and Drag Acting on the Shell of the American Horseshoe Crab (Limulus polyphemus).

The intertidal zone is a turbulent landscape where organisms face numerous mechanical challenges from powerful waves. A model for understanding the solutions to these physical problems, the American horseshoe crab (Limulus polyphemus), is a marine arthropod that mates in the intertidal zone, where it must contend with strong ambient flows to maintain its orientation during locomotion and reproduction. Possible strategies to maintain position include either negative lift generation or the minimization of positive lift in flow. To quantify flow over the shell and the forces generated, we laser-scanned the 3D shape of a horseshoe crab, and the resulting digital reconstruction was used to 3D-print a physical model. We then recorded the movement of tracking particles around the shell model with high-speed video and analyzed the time-lapse series using particle image velocimetry (PIV). The velocity vector fields from PIV were used to validate numerical simulations performed with the immersed boundary (IB) method. IB simulations allowed us to resolve the forces acting on the shell, as well as the local three-dimensional flow velocities and pressures. Both IB simulations and PIV analysis of vorticity and velocity at a flow speed of 13 cm/s show negative lift for negative and zero angles of attack, and positive lift for positive angles of attack in a free-stream environment. In shear flow simulations, we found near-zero lift for all orientations tested. Because horseshoe crabs are likely to be found primarily at near-zero angles of attack, we suggest that this negative lift helps maintain the orientation of the crab during locomotion and mating. This study provides a preliminary foundation for assessing the relationship between documented morphological variation and potential environmental variation for distinct populations of horseshoe crabs along the Atlantic Coast. It also motivates future studies which could consider the stability of the horseshoe crab in unsteady, oscillating flows.

Effects of the Biomedical Bleeding Process on the Behavior of the American Horseshoe Crab, Limulus polyphemus, in Its Natural Habitat.

Horseshoe crabs are harvested by the biomedical industry in order to create Limulus amebocyte lysate to test medical devices and pharmaceutical drugs for endotoxins. Most previous studies on the impacts of the biomedical bleeding process on horseshoe crabs have focused on mortality rates and sublethal impacts in the laboratory. In this study, we investigated the effects of the bleeding process on the behavior of horseshoe crabs after they had been released back into their natural environment. A total of 28 horseshoe crabs (14 control and 14 bled) were fitted with acoustic transmitters and released into the Great Bay Estuary, New Hampshire, during the spring of 2016. The acoustic tags transmitted information about the activity and depth of each animal, and these data were logged by an array of passive acoustic receivers. These data were collected from May to December 2016 and from March to October 2017. Bled animals approached mating beaches less than control animals during the first week after release, with the greatest differences between bled and control females. Bled animals also remained significantly deeper during the spawning season than control animals. However, overall, bled and control animals expressed similar biological rhythms and seasonal migrations. Thus, it appears as if the most obvious impacts of the bleeding process take place during the first one to two weeks after crabs are bled.

2-PHENOXYETHANOL (2-PE) AND TRICAINE METHANESULFONATE (MS-222) IMMERSION ANESTHESIA OF AMERICAN HORSESHOE CRABS (LIMULUS POLYPHEMUS).

Despite extensive literature examining American horseshoe crab physiology, there are comparatively few publications addressing their medical care. Establishing anesthesia protocols for horseshoe crabs is integral to limiting the potential stress and pain associated with invasive procedures and for advancing euthanasia techniques. The objective of this study was to compare the effects of two immersion anesthetics, tricaine methanesulfonate (MS-222) at 1 g/L (buffered with sodium carbonate) and 2-phenoxyethanol (2-PE) at 2 mL/L, on horseshoe crabs. Twenty horseshoe crabs were assigned to one of two anesthetic treatment groups and individually anesthetized in natural seawater. Water quality, cardiac contractility, and hemolymph gas analytes were measured prior to anesthesia and at 30 min Animals were monitored via heart rate, gilling rate, and sedation score every 5 min until recovered. Transcarapacial ultrasonography was used to obtain heart rate, gilling rate, and percent fractional shortening. Light or surgical anesthesia was produced in 10/10 animals in the 2-PE group and 8/10 animals in the MS-222 group. There was no significant difference in sedation scores, induction time (median 15 min), or recovery time (median 20.5 min). Gilling rate and cardiac contractility decreased during anesthesia, whereas heart rate did not. Hemolymph pH and pO2 were not different among treatment groups or time points. Baseline pCO2 was higher than pCO2 at 30 min for both groups but significantly elevated only in the MS-222 group. This is attributed to increased activity during the handling of awake animals. Invasive blood pressure obtained via cardiac catheterization in two animals was markedly decreased during surgical anesthesia. In conclusion, 2-PE and MS-222 provided effective anesthesia with clinically useful induction and recovery times. 2-PE provided a subjectively more reliable and smoother anesthesia compared to MS-222.

Limulus and heart rhythm.

Great interest in the comparative physiology of hearts and their functions in Animalia has emerged with classic papers on Limulus polyphemus and mollusks. The recurrent cardiac activity-heart rate-is the most important physiological parameter and when present the kardia (Greek) is vital to the development of entire organs of the organisms in the animal kingdom. Extensive studies devoted to the regulation of cardiac rhythm in invertebrates have revealed that the basics of heart physiology are comparable to mammals. The hearts of invertebrates also beat spontaneously and are supplied with regulatory nerves: either excitatory or inhibitory or both. The distinct nerves and the source of excitation/inhibition at the level of single neurons are described for many invertebrate genera. The vertebrates and a majority of invertebrates have myogenic hearts, whereas the horseshoe crab L. polyphemus and a few other animals have a neurogenic cardiac rhythm. Nevertheless, the myogenic nature of heartbeat is precursor, because the contraction of native and stem-cell-derived cardiomyocytes does occur in the absence of any neural elements. Even in L. polyphemus, the heart rhythm is myogenic at embryonic stages.

Computational biomechanical analyses demonstrate similar shell-crushing abilities in modern and ancient arthropods.

The biology of the American horseshoe crab, Limulus polyphemus, is well documented-including its dietary habits, particularly the ability to crush shell with gnathobasic walking appendages-but virtually nothing is known about the feeding biomechanics of this iconic arthropod. Limulus polyphemus is also considered the archetypal functional analogue of various extinct groups with serial gnathobasic appendages, including eurypterids, trilobites and other early arthropods, especially Sidneyia inexpectans from the mid-Cambrian (508 Myr) Burgess Shale of Canada. Exceptionally preserved specimens of S. inexpectans show evidence suggestive of durophagous (shell-crushing) tendencies-including thick gnathobasic spine cuticle and shelly gut contents-but the masticatory capabilities of this fossil species have yet to be compared with modern durophagous arthropods. Here, we use advanced computational techniques, specifically a unique application of 3D finite-element analysis (FEA), to model the feeding mechanics of L. polyphemus and S. inexpectans: the first such analyses of a modern horseshoe crab and a fossil arthropod. Results show that mechanical performance of the feeding appendages in both arthropods is remarkably similar, suggesting that S. inexpectans had similar shell-crushing capabilities to L. polyphemus This biomechanical solution to processing shelly food therefore has a history extending over 500 Myr, arising soon after the first shell-bearing animals. Arrival of durophagous predators during the early phase of animal evolution undoubtedly fuelled the Cambrian 'arms race' that involved a rapid increase in diversity, disparity and abundance of biomineralized prey species.

A 3D anatomical atlas of appendage musculature in the chelicerate arthropod Limulus polyphemus.

Limulus polyphemus, an archetypal chelicerate taxon, has interested both biological and paleontological researchers due to its unique suite of anatomical features and as a useful modern analogue for fossil arthropod groups. To assist the study and documentation of this iconic taxon, we present a 3D atlas on the appendage musculature, with specific focus on the muscles of the cephalothoracic appendages. As L. polyphemus appendage musculature has been the focus of extensive study, depicting the muscles in 3D will facilitate a more complete understanding thereof for future researchers. A large museum specimen was CT scanned to illustrate the major exoskeletal features of L. polyphemus. Micro-CT scans of iodine-stained appendages from fresh, non-museum specimens were digitally dissected to interactively depict appendage sections and muscles. This study has revealed the presence of two new muscles: one within the pushing leg, located dorsally relative to all other patella muscles, and the other within the male pedipalp, located in the modified tibiotarsus. This atlas increases accessibility to important internal and external morphological features of L. polyphemus and reduces the need for destructive fresh tissue dissection of specimens. Scanning, digitally dissecting, and documenting taxa in 3D is a pivotal step towards creating permanent digital records of life on Earth.

Simple Eyes, Extraocular Photoreceptors and Opsins in the American Horseshoe Crab.

The eyes and photoreceptors of the American horseshoe crab Limulus polyphemus have been studied since the 1930s, and this work has been critical for understanding basic mechanisms of vision. One of the attractions of Limulus as a preparation for studies of vision is that it has three different types of eyes-a pair of later compound, image-forming eyes and two types of simple eyes, a pair of median ocelli, and three pair of larval eyes. Each eye type is tractable for experimentation. Limulus also has extraocular photoreceptors in its segmental ganglia and tail. The current contribution focuses on photoreceptors in Limulus larval eyes and ocelli and its extraocular photoreceptors with the goal of summarizing what is currently known and not known about their physiology and function and the opsins they express. The Limulus genome encodes a surprisingly large number of opsins (18), and studies of their expression pattern have raised new questions about the role of opsin co-expression, the functions of peropsins expressed outside of eyes, and the physiological relevance of opsins with apparently very low expression levels. Studies of opsin expression in Limulus lead one to wonder whether photoreceptors yet to be discovered might be present throughout its central nervous system.

Movement Patterns and Residency of the Critically Endangered Horseshoe Crab Tachypleus tridentatus in a Semi-Enclosed Bay Determined Using Acoustic Telemetry.

The horseshoe crab Tachypleus tridentatus is critically endangered in Japan due to rapidly decreasing numbers resulting from the loss of tidal flats and sandy beaches, and the deterioration of coastal environments. We monitored the year-round migratory patterns and residency of this species in a coastal embayment at Tsuyazaki, Japan, using acoustic telemetry. Total 20 adult crabs (15 males and 5 females) were tagged with ultrasonic transmitters and tracked during two periods (2006-2008; n = 10 and 2007-2009; n = 10). Adult crabs were more active during periods of higher water temperatures and their activity peaked in July, during the spawning period. Water temperature appeared to be one of the key factors influencing the movement patterns for the species. Moreover, the crabs tended to be more active at night than in the day. The nocturnal activity pattern was clearly evident before and during the reproductive period (May-August). Tracking data also showed that one pair-bond was maintained for a maximum of 17 days after the pair-bonded female had spawned. Overall, 11 males (73% of 15 individuals) remained in the bay area over winter, whereas three females (60% of 5 individuals) overwintered outside of the bay. Telemetry data showed that over 60% (13 of 20) of tagged crabs overwintered within the bay where there are sandy beaches, mudflats, and scattered seagrass beds. This year-round residence by adult T. tridentatus in the bay area identifies it as a critical habitat for the management of this species, regardless of life-stage. Not only is it a comprehensive management strategy that effectively reflects this species' habitat use patterns but also its implementation, such as the establishment of a protected area, would contribute to its conservation.

Colour preference and light sensitivity in trilobite larvae of mangrove horseshoe crab, Carcinoscopius rotundicauda (Latreille, 1802).

The trilobite larvae of C. rotundicauda were tested to determine their colour preference and light sensitivity until their first moulting (25 days post hatching) under laboratory conditions. Maximum congregation size of the trilobite larvae was found in the white zone respectively where (n = 12) followed by yellow (n = 8) and orange (n = 8), which showed the larval preference for lighter zones. Morisita's index calculation showed a clumped/aggregated distribution (yellow, blue, orange and white) and uniform/hyper dispersed distribution (green, red and black) for various colours tested. Trilobite larvae showed least preference for brighter regions while tested in the experiment [black; (n = 4) and red; (n = 5)]. Experiments done to determine the light sensitivity of trilobite larvae showed that the larvae had more preference towards ultraviolet lights. The maximum congregation size of 38.8 and 40.7% of the larvae was encountered under ultraviolet light, when the light sources were kept horizontal and vertical, respectively. Overall, results suggested that the trilobite larvae of C. rotundicauda, preferred light source of shorter wavelengths (UV light) and colours of lighter zone (white, yellow, orange), which might be due to their adaptation to their natural habitat for predator avoidance, prey selection and water quality.

Physiological energetics of the fourth instar of Chinese horseshoe crabs (Tachypleus tridentatus) in response to hypoxic stress and re-oxygenation.

Hypoxia associated with eutrophication is a potential threat to the Chinese horseshoe crab Tachypleus tridentatus which inhabits intertidal sand flats in Asia. This study investigated the effect of dissolved oxygen level (DO) (6, 4 and 2 mg O2 l(-1)) on the physiological energetics in the juvenile T. tridentatus. They were exposed to various oxygen levels for three days and then transferred to normoxia for three days to examine the recovery from low oxygen stress. Feeding rate, respiration rate and scope for growth were reduced at lower DO levels while absorption efficiency and excretion rate were independent of DO levels. Although full recovery of the physiological responses and scope for growth from hypoxis stress was observed when normoxia resumed, their long term survival in suboptimal habitats with frequent occurrence of hypoxia deserves a close monitoring as hypoxia may be even more common in future in a warming world.

Adaptive management: from more talk to real action.

The challenges currently facing resource managers are large-scale and complex, and demand new approaches to balance development and conservation goals. One approach that shows considerable promise for addressing these challenges is adaptive management, which by now is broadly seen as a natural, intuitive, and potentially effective way to address decision-making in the face of uncertainties. Yet the concept of adaptive management continues to evolve, and its record of success remains limited. In this article, we present an operational framework for adaptive decision-making, and describe the challenges and opportunities in applying it to real-world problems. We discuss the key elements required for adaptive decision-making, and their integration into an iterative process that highlights and distinguishes technical and social learning. We illustrate the elements and processes of the framework with some successful on-the-ground examples of natural resource management. Finally, we address some of the difficulties in applying learning-based management, and finish with a discussion of future directions and strategic challenges.