Host Anemones Research Articles

A New Observation about the Symbiotic Relationship of Clownfish with its Host Anemones: Documentation in Captivity

A New Observation about the Symbiotic Relationship of Clownfish with its Host Anemones: Documentation in Captivity

Effects of Personality and Behavioral Syndromes on Competition for Social Hierarchical Status in Anemonefish Amphiprion clarkii.

In this study, the behavioral ethogram of Amphiprion clarkii during the growth phase prior to sexual differentiation was summarized based on behavioral observations in three social environments. These behaviors can be classified into four categories: in addition to normal behaviors, the other three categories of behaviors-threatening, agonistic, and appeasing behaviors-represent different intentions in interactions with other individuals. Subsequently, the personalities of each individual were assessed by testing their reactions to intruders. These individuals mainly exhibited two distinct personality types: bold-aggressive and shy-submissive. In pairing experiments, the interactive behaviors of the anemonefish were observed in pairing combinations of different body sizes and personalities. The impact of personality on the establishment of a stable social hierarchy was confirmed by significant differences in the success rates of different pairing combinations, with the frequency of appeasing behaviors being the main factor influencing the success rate. Our results suggested that in natural waters, when juvenile individuals migrate among host anemones, shy-submissive individuals are more likely to be accepted due to their appeasing behaviors towards larger individuals, thus avoiding the risk of being attacked and bitten, and benefiting the survival of the individual. Conversely, bold-aggressive individuals are more likely to be driven away to another host anemone due to their unwillingness to settle for a lower-ranked status, thereby contributing to population dispersal and increasing opportunities for gene exchange between populations.

Sea anemone-anemonefish symbiosis: Behavior and mucous protein profiling.

Fish species of the genus Amphiprion (Perciformes: Pomacentridae) seek protection from predators among the tentacles of sea anemones as their natural habitat, where they live essentially unharmed from stinging by the host's nematocysts. The skin mucus of these anemonefish has been suggested as a protective mechanism that prevents the discharge of the nematocysts upon contact. Whereas some anemonefish species seem to produce their own protective mucous coating, others may acquire mucus (or biomolecules within) from the sea anemone during an acclimation period. In controlled experiments, we show that Amphiprion ocellaris acclimated successfully to their natural host anemone species Stichodactyla gigantea, and also to Stichodactyla haddoni, and in some cases Heteractis crispa, neither of which are natural host species. No symbiosis was observed for three other anemone species tested, Entacmaea quadricolor, Macrodactyla doreensis, and Heteractis malu. We explored the skin mucous protein profile from naive and experienced A. ocellaris during their acclimation to natural and unnatural host anemones. We confidently report the presence of metabolic and structural proteins in the skin mucus of all samples, likely involved in immunological defense, molecular transport, stress response, and signal transduction. For those anemonefish that established symbiosis, there was a clear increase in ribosomal-type proteins. We additionally provide evidence for the presence of anemone proteins only in the skin mucus of individuals that established symbiosis. Our results support previous speculation of the role of skin mucous-associated proteins in anemonefish-anemone symbiosis. Further exploration of these mucosal proteins could reveal the mechanism of anemonefish acclimation to host anemones.

Anemonefish are better taxonomists than humans

The symbiosis between giant sea anemones, algae of the family Symbiodiniaceae, and anemonefish is an iconic example of a mutualistic trio1,2. Molecular analyses have shown that giant sea anemones hosting anemonefish belong to three clades: Entacmaea, Stichodactyla, and Heteractis3,4,5 (Figure 1A). Associations among 28 species of anemonefish and 10 species of giant sea anemone hosts are complex. Some fish species are highly specialized to only one anemone species (e.g., Amphiprion frenatus with Entacmaea quadricolor), whereas others are more generalist (e.g., Amphiprion clarkii)1,2,6. Reasons for host preferences are obscured, among other things, by the lack of resolution in the giant sea anemone phylogeny. Here, we generated a transcriptomic dataset from 55 sea anemones collected from southern Japan to reconstruct these phylogenetic relationships. We observed that the bubble-tip sea anemone E. quadricolor, currently considered a single species, can be separated into at least four cryptic lineages (A-D). Surprisingly, these lineages can be precisely distinguished by observing their association with anemonefish: A. frenatus only associates with lineage D, whereas A. clarkii lives in the other three lineages.

Counting Nemo: anemonefish Amphiprion ocellaris identify species by number of white bars.

The brilliant colors of coral reef fish have received much research attention. This is well exemplified by anemonefish, which have distinct white bar patterns and inhabit host anemones and defend them as a territory. The 28 described species have between 0 and 3 white bars present, which has been suggested to be important for species recognition. In the present study, we found that Amphiprion ocellaris (a species that displays three white bars) hatched and reared in aquaria, when faced with an intruder fish, attacked their own species more frequently than other species of intruding anemonefish. Additionally, we explicitly tested whether this species could distinguish models with different numbers of bars. For this, 120 individuals of A. ocellaris were presented with four different models (no bars, and 1, 2 and 3 bars) and we compared whether the frequency of aggressive behavior towards the model differed according to the number of bars. The frequency of aggressive behavior toward the 3-bar model was the same as against living A. ocellaris, and was higher than towards any of the other models. We conclude that A. ocellaris use the number of white bars as a cue to identify and attack only competitors that might use the same host. We considered this as an important behavior for efficient host defense.

Insights into the Genomics of Clownfish Adaptive Radiation: The Genomic Substrate of the Diversification.

Clownfishes are an iconic group of coral reef fishes that evolved a mutualistic interaction with sea anemones, which triggered the rapid diversification of the group. Following the emergence of this mutualism, clownfishes diversified into different ecological niches and developed convergent phenotypes associated with their host use. The genetic basis of the initial acquisition of the mutualism with host anemones has been described, but the genomic architecture underlying clownfish diversification once the mutualism was established and the extent to which clownfish phenotypic convergence originated through shared genetic mechanisms are still unknown. Here, we investigated these questions by performing comparative genomic analyses on the available genomic data of five pairs of closely related but ecologically divergent clownfish species. We found that clownfish diversification was characterized by bursts of transposable elements, an overall accelerated coding evolution, incomplete lineage sorting, and ancestral hybridization events. Additionally, we detected a signature of positive selection in 5.4% of the clownfish genes. Among them, five presented functions associated with social behavior and ecology, and they represent candidate genes involved in the evolution of the size-based hierarchical social structure so particular to clownfishes. Finally, we found genes with patterns of either relaxation or intensification of purifying selection and signals of positive selection linked with clownfish ecological divergence, suggesting some level of parallel evolution during the diversification of the group. Altogether, this work provides the first insights into the genomic substrate of clownfish adaptive radiation and integrates the growing collection of studies investigating the genomic mechanisms governing species diversification.

Divergence in metabolomic profile in clownfish and damselfish skin mucus

IntroductionThe clownfish - sea anemone mutualism was suggested to have triggered the adaptive radiation of clownfishes, but the origin of clownfish resistance to stinging tentacles of host anemones remains unclear. The presence of specific compounds in the mucus of clownfishes conferring them the unique ability to prevent nematocyst discharge from their hosts has been the most supported hypothesis. Yet the mystery regarding the types of compounds found in clownfish skin mucus remains unsolved.MethodsWe analyzed the chemical composition of clownfish and damselfish mucus using an untargeted metabolomics (HILIC-HRMS) and lipidomics (RPLC-HRMS) approach.Results and DiscussionThe polar and lipid metabolome signatures were highly specific and allowed to discriminate between the clownfish and damselfish clades. The most discriminative part of the signature was the sphingolipid profile, displaying a broader diversity of ceramides present in significantly higher levels in clownfish mucus. Importantly, the inter-specific variability of metabolic signature was significantly higher in clownfishes, although their diversification is evolutionarily more recent, thus implying the impact of symbiosis on metabolic variability and adaptation. Furthermore, specialists and generalists clownfish species displayed distinctive metabolite signature. Two strict clownfish specialists, which are phylogenetically distant but share the same host species, clustered together based on their molecular signature, suggesting a link with their mutualistic nature. Overall, comparative analyses of metabolic signatures highlight differences in chemical composition of clownfish mucus and provide insight into biochemical pathways potentially implicated in clownfish adaptation to inhabit sea anemones and consequently diversify.

Colour patterns influence symbiosis and competition in the anemonefish-host anemone symbiosis system.

Colour patterns in fish are often used as an important medium for communication. Anemonefish, characterized by specific patterns of white bars, inhabit host anemones and defend the area around an anemone as their territory. The host anemone is used not only by the anemonefish, but also by other fish species that use anemones as temporary shelters. Anemonefish may be able to identify potential competitors by their colour patterns. We first examined the colour patterns of fish using host anemones inhabited by Amphiprion ocellaris as shelter and compared them with the patterns of fish using surrounding scleractinian corals. There were no fish with bars sheltering in host anemones, although many fish with bars were found in surrounding corals. Next, two fish models, one with white bars and the other with white stripes on a black background, were presented to an A. ocellaris colony. The duration of aggressive behaviour towards the bar model was significantly longer than that towards the stripe model. We conclude that differences in aggressive behaviour by the anemonefish possibly select the colour patterns of cohabiting fish. This study indicates that colour patterns may influence not only intraspecific interactions but also interspecific interactions in coral reef ecosystems.

Rank change and growth within social hierarchies of the orange clownfish, Amphiprion percula

Social hierarchies within groups define the distribution of resources and provide benefits that support the collective group or favor dominant members. The progression of individuals through social hierarchies is a valuable characteristic for quantifying population dynamics. On coral reefs, some clownfish maintain size-based hierarchical communities where individuals queue through social ranks. The cost of waiting in a lower-ranked position is outweighed by the reduced risk of eviction and mortality. The orange clownfish, Amphiprion percula, maintains stable social groups with subordinate individuals queuing to be part of the dominant breeding pair. Strong association with their host anemone, complex social interactions, and relatively low predation rates make them ideal model organisms to assess changes in group dynamics through time in their natural environment. Here, we investigate the rank changes and isometric growth rates of A. percula from 247 naturally occurring social groups in Kimbe Island, Papua New Guinea (5° 12′ 13.54″ S, 150° 22′ 32.69″ E). We used DNA profiling to assign and track individuals over eight years between 2011 and 2019. Over half of the individuals survived alongside two or three members of their original social group, with twelve breeding pairs persisting over the study period. Half of the surviving individuals increased in rank and experienced double the growth rate of those that maintained their rank. Examining rank change in a wild fish population provides new insights into the complex social hierarchies of reef fishes and their role in social evolution.

Increased incompatibility of heterologous algal symbionts under thermal stress in the cnidarian-dinoflagellate model Aiptasia

Rising ocean temperatures are increasing the rate and intensity of coral mass bleaching events, leading to the collapse of coral reef ecosystems. To better understand the dynamics of coral-algae symbioses, it is critical to decipher the role each partner plays in the holobiont’s thermotolerance. Here, we investigated the role of the symbiont by comparing transcriptional heat stress responses of anemones from two thermally distinct locations, Florida (CC7) and Hawaii (H2) as well as a heterologous host-symbiont combination composed of CC7 host anemones inoculated with the symbiont Breviolum minutum (SSB01) from H2 anemones (CC7-B01). We find that oxidative stress and apoptosis responses are strongly influenced by symbiont type, as further confirmed by caspase-3 activation assays, but that the overall response to heat stress is dictated by the compatibility of both partners. Expression of genes essential to symbiosis revealed a shift from a nitrogen- to a carbon-limited state only in the heterologous combination CC7-B01, suggesting a bioenergetic disruption of symbiosis during stress. Our results indicate that symbiosis is highly fine-tuned towards particular partner combinations and that heterologous host-symbiont combinations are metabolically less compatible under stress. These results are essential for future strategies aiming at increasing coral resilience using heterologous thermotolerant symbionts.

Vertebrate growth plasticity in response to variation in a mutualistic interaction

Vertebrate growth can be phenotypically plastic in response to predator–prey and competitive interactions. It is unknown however, if it can be plastic in response to mutualistic interactions. Here we investigate plasticity of vertebrate growth in response to variation in mutualistic interactions, using clown anemonefish and their anemone hosts. In the wild, there is a positive correlation between the size of the fish and the size of the anemone, but the cause of this correlation is unknown. Plausible hypotheses are that fish exhibit growth plasticity in response to variation in food or space provided by the host. In the lab, we pair individuals with real anemones of various sizes and show that fish on larger anemones grow faster than fish on smaller anemones. By feeding the fish a constant food ration, we exclude variation in food availability as a cause. By pairing juveniles with artificial anemones of various sizes, we exclude variation in space availability as a single cause. We argue that variation in space availability in conjunction with host cues cause the variability in fish growth. By adjusting their growth, anemonefish likely maximize their reproductive value given their anemone context. More generally, we demonstrate vertebrate growth plasticity in response to variation in mutualistic interactions.

Manganese uptake and partitioning between the tissue of the anemone host Exaiptasia pallida and Symbiodinium spp., including assessment of stress and recovery

Manganese (Mn) is essential for global steel and Mn-iron (Fe) alloy production. The human health effects of elevated Mn concentrations have been well established, but studies on its impact on marine invertebrates are limited. This study is the first to investigate Mn uptake in the sea anemone Exaiptasia pallida after chronic exposure (0.5, 1, 10, and 100 mg/L) for 24-d. Following exposure, E. pallida were transferred to ambient seawater for 6-d to assess Mn depuration. Mn accumulation and partitioning in host tissue and symbionts (Symbiodinium spp.), tentacle retraction, and symbiont cell density were measured during exposure and depuration. Mn concentrations were substantially higher in symbionts than tissue in all treatments after 24-d. No significant difference was observed for symbiont cell density after Mn exposure. Tentacle retractions were significantly higher in all Mn exposed treatments than controls at all time points. Mn depuration was observed for both tissue and symbionts but was more rapid in symbionts. This study reveals that Symbiodinium spp. can play a role in Mn uptake and depuration in anemones, but Mn loading does not affect cell density. These results help understand metal uptake and depuration in complex relationships between Symbiodinium spp. and other host taxa like corals.

Clownfish hosting anemones (Anthozoa, Actiniaria) of the Red Sea: new associations and distributions, historical misidentifications, and morphological variability

BackgroundThe Red Sea contains thousands of kilometers of fringing reef systems inhabited by clownfish and sea anemones, yet there is no consensus regarding the diversity of host anemone species that inhabit this region. We sought to clarify a historical record and recent literature sources that disagree on the diversity of host anemone species in the Red Sea, which contains one endemic anemonefish, Amphiprion bicinctus Rüppell 1830.ResultsWe conducted 73 surveys spanning ~ 1600 km of coastline from the northern Saudi Arabian Red Sea to the Gulf of Aden and encountered seven species of host anemones, six of which hosted A. bicinctus. We revise the list of symbionts for A. bicinctus to include Stichodactyla haddoni (Saville-Kent, 1893) and Stichodactyla mertensii Brandt, 1835 which were both observed in multiple regions. We describe Red Sea phenotypic variability in Heteractis crispa (Hemprich & Ehrenberg in Ehrenberg, 1834) and Heteractis aurora (Quoy & Gaimard, 1833), which may indicate that these species hybridize in this region. We did not encounter Stichodactyla gigantea (Forsskål, 1775), although the Red Sea is the type locality for this species. Further, a thorough review of peer-reviewed literature, occurrence records, and misidentified basis of record reports dating back to the early twentieth century indicate that it is unlikely that S. gigantea occurs in the Red Sea.ConclusionsIn sum, we present a new guide for the host anemones of the Red Sea, revise the host specificity of A. bicinctus, and question whether S. gigantea occurs in the central and western Indian Ocean.

Large host anemones can be shelters of a diverse assemblage of fish species, not just anemonefish.

The mutualistic relationships between anemonefish (Amphiprion; Pomacentridae) and host sea anemones are well known, but host anemones are also used as shelter by damselfish (Pomacentridae), wrasses (Labridae) and cardinalfish (Apogonidae). The threespot dascyllus Dascyllus trimaculatus (Pomacentridae) is known to live on or near host anemones in their immature phase. Nonetheless, detailed studies on the use of host anemones by other fish species have not yet been conducted. To understand the factors that influence the use of host anemones by other fish species, this study investigated the fish biota around host anemones in the Ryukyu Archipelago. Other than Amphiprion spp. and D. trimaculatus, 10 additional species of fish (9 species of damselfish and 1 species of cardinalfish) were confirmed to temporarily use host anemones as shelter, and five species of wrasse including Labroides dimidiatus came to clean anemonefish. Logistic regression analyses (independent variable: presence or absence of other species of fish; dependent variables: anemonefish aggressiveness, size of host anemone, number of D. trimaculatus) indicated that the size of host anemones is important for the presence of fish, both in species using the anemone as temporary shelter and in cleaner species. Large host anemones can provide shelter and food resources for other species of coral reef fish as well as for anemonefish.

The Importance of Home Cleaning: Sediment Transport by Alpheid Shrimps Provides a Competitive Advantage to Their Host Anemones

Bartholomea annulata is a facultative host of the A. armatus species complex. In the Mexican Caribbean it is commonly found in cracks and crevices located where the vertical walls meet the sandy bottom or on large coral patches away from the sand. To protect themselves from predators, anemones often contract their hydraulic body into a cavernous den and extend the stinging tentacles toward the entrance. The high sediment dynamics of the region, however, result in a permanent risk of animal shelters to be obstructed by sand. By both analysing field data and conducting laboratory experiments with artificial shelters, the present study explored the den cleaning behaviour widely extended amongst alpheid shrimp, and its role in the alpheid-anemone symbiotic interaction. Videorecordings showed that den cleaning was composed of three main behaviours: digging, tossing and tamping. It commenced as soon as 7.2 ± 10.5 min after anemones were recognised by alpheids, and behaviours were displayed systematically amongst all 12 replicates. Despite being completely burrowed in sand, Alpheus spp. were capable of finding the anemone and liberating the entrance of the artificial shelters in less than 2.5 h. In addition, manipulative experiments showed that anemones confronted with shelters that were obstructed with sediment had a 25% probability of fully retracting when Alpheus spp. were absent, compared to a 75% probability when shrimps had cleaned the shelter’s entry and internal passage. The analysis of field data indicated that the presence of alpheid shrimps as symbionts of B. annulata was 30% higher amongst anemones in close contact with sandy bottoms than when inhabiting crevices on the top or lateral walls of hard substrates, away from the sediment. Overall, our study concludes that den cleaning constitutes a quick and effective mechanism to assure the anemone’s full retraction into their den, and by keeping the sediment away, alpheids provide the necessary conditions that serve both the anemone’s and the shrimp’s higher chances of acquiring maximum protection from predators. This advantageous exchange in protection partially explains why alpheids are more frequently present in B. annulata in interface microhabitats where the benefit of the interaction is maximised.

Microbiomes of clownfish and their symbiotic host anemone converge before their first physical contact

BackgroundOne of the most charismatic, and yet not completely resolved example of mutualistic interaction is the partnership of clownfish and its symbiotic sea anemone. The mechanism explaining this tolerance currently relies on the molecular mimicry of clownfish epithelial mucus, which could serve as camouflage, preventing the anemone's nematocysts' discharge. Resident bacteria are known as key drivers of epithelial mucus chemical signature in vertebrates. A recent study has proposed a restructuration of the skin microbiota in a generalist clown fish when first contacting its symbiotic anemone. We explored a novel hypothesis by testing the effect of remote interaction on epithelial microbiota restructuration in both partners.MethodsWith metataxonomics, we investigated the epithelial microbiota dynamic of 18 pairs of percula clownfish (Amphiprion percula) and their symbiotic anemone Heteractis magnifica in remote interaction, physical interaction and control groups for both partners during a 4-week trial.ResultsThe Physical and Remote Interaction groups’ results evidence gradual epithelial microbiota convergence between both partners when fish and anemone were placed in the same water system. This convergence occurred preceding any physical contact between partners, and was maintained during the 2-week interaction period in both contact groups. After the interaction period, community structure of both fish and anemone’s epthelial community structures maintained the interaction signature 2 weeks after fish–anemone pairs’ separation. Furthermore, the interaction signature persistence was observed both in the Physical and Remote Interaction groups, thus suggesting that water-mediated chemical communication between symbiotic partners was strong enough to shift the skin microbiota durably, even after the separation of fish–anemone pairs. Finally, our results suggest that fish–anemone convergent microbiota restructuration was increasingly associated with the parallel recruitment of three Flavobacteriaceae strains closely related to a tyrosinase-producing Cellulophaga tyrosinoxydans.ConclusionsOur study shows that bacterial community restructuration, in the acclimation process, does not only rely on direct physical contact. Furthermore, our results challenge, for the first time, the traditional unidirectional chemical camouflage hypothesis, as we argue that convergence of the epithelial microbiota of both partners may play essential roles in establishing mutual acceptance.9p4RZDq8XFt4wx9Qb_GP5vVideo abstract Fish−anemone symbiotic relationship.

Finding Nemo\u2019s clock reveals switch from nocturnal to diurnal activity

Timing mechanisms play a key role in the biology of coral reef fish. Typically, fish larvae leave their reef after hatching, stay for a period in the open ocean before returning to the reef for settlement. During this dispersal, larvae use a time-compensated sun compass for orientation. However, the timing of settlement and how coral reef fish keep track of time via endogenous timing mechanisms is poorly understood. Here, we have studied the behavioural and genetic basis of diel rhythms in the clown anemonefish Amphiprion ocellaris. We document a behavioural shift from nocturnal larvae to diurnal adults, while juveniles show an intermediate pattern of activity which potentially indicates flexibility in the timing of settlement on a host anemone. qRTPCR analysis of six core circadian clock genes (bmal1, clocka, cry1b, per1b, per2, per3) reveals rhythmic gene expression patterns that are comparable in larvae and juveniles, and so do not reflect the corresponding activity changes. By establishing an embryonic cell line, we demonstrate that clown anemonefish possess an endogenous clock with similar properties to that of the zebrafish circadian clock. Furthermore, our study provides a first basis to study the multi-layered interaction of clocks from fish, anemones and their zooxanthellae endosymbionts.

Impact of Menthol on Growth and Photosynthetic Function of Breviolum Minutum (Dinoflagellata, Dinophyceae, Symbiodiniaceae) and Interactions with its Aiptasia Host.

Environmental change, including global warming and chemical pollution, can compromise cnidarian-(e.g., coral-) dinoflagellate symbioses and cause coral bleaching. Understanding the mechanisms that regulate these symbioses will inform strategies for sustaining healthy coral-reef communities. A model system for corals is the symbiosis between the sea anemone Exaiptasia pallida (common name Aiptasia) and its dinoflagellate partners (family Symbiodiniaceae). To complement existing studies of the interactions between these organisms, we examined the impact of menthol, a reagent often used to render cnidarians aposymbiotic, on the dinoflagellate Breviolum minutum, both in culture and in hospite. In both environments, the growth and photosynthesis of this alga were compromised at either 100 or 300µM menthol. We observed reduction in PSII and PSI functions, the abundances of reaction-center proteins, and, at 300µM menthol, of total cellular proteins. Interestingly, for free-living algae exposed to 100µM menthol, an initial decline in growth, photosynthetic activities, pigmentation, and protein abundances reversed after 5-15 d, eventually approaching control levels. This behavior was observed in cells maintained in continuous light, but not in cells experiencing a light-dark regimen, suggesting that B. minutum can detoxify menthol or acclimate and repair damaged photosynthetic complexes in a light- and/or energy-dependent manner. Extended exposures of cultured algae to 300µM menthol ultimately resulted in algal death. Most symbiotic anemones were also unable to survive this menthol concentration for 30 d. Additionally, cells impaired for photosynthesis by pre-treatment with 300µM menthol exhibited reduced efficiency in re-populating the anemone host.

Physiological and behavioural effects of anemone bleaching on symbiont anemonefish in the wild

Abstract Climate change causes extreme heat waves that have induced worldwide mass coral bleaching. The impacts of temperature‐induced bleaching events on the loss of algal endosymbionts in both corals and anemones are well documented. However, the cascading impacts of bleaching on animals that live in association with corals and anemones are understudied. We performed a field‐based experiment to investigate how host anemone bleaching affected the metabolic rate, growth, behaviour and survival of wild juvenile orange‐fin anemonefish Amphiprion chrysopterus over 1, 2 and (for survival) 9 months. We found that the standard metabolic rate of anemonefish residing in bleached anemones decreased over time but was unaffected in fish from healthy anemones. Despite the reduced metabolic cost, the growth rate of fish from bleached anemones was significantly lower compared to fish from healthy anemones, suggesting that animals residing in bleached hosts are at an energetic disadvantage. This was corroborated by our finding that fish from bleached anemones spent more time out of their anemones, suggestive of a greater need to forage in the water column. However, fish from bleached anemones were overall less active and used less space around the anemone, resulting in a negative correlation between space use and survival after 4 weeks. Our results provide insight into the physiological and behavioural effects of host bleaching on juvenile fish in the wild, and highlight how relatively short‐term thermal anomalies can have long‐lasting impacts beyond the bleached anemones or corals themselves. A free Plain Language Summary can be found within the Supporting Information of this article.

Manganese Uptake and Partitioning between the Tissue of the Anemone Host Exaiptasia Pallida and Symbiodinium Sp., Including Assessment of Stress and Recovery

Manganese Uptake and Partitioning between the Tissue of the Anemone Host Exaiptasia Pallida and Symbiodinium Sp., Including Assessment of Stress and Recovery