Exoskeletal Trade-off between Claws and Carapace in Deep-sea Hydrothermal Vent Decapod Crustaceans.

Limitations on energetic resources create evolutionary trade-offs, prompting us to investigate if investment in claw strength remains consistent across crustaceans living in diverse habitats. Decapod crustaceans living in deep-sea hydrothermal vents are ideal for this study due to their extreme environment. In this study, we investigated whether decapods (blind crab Austinograea sp. and the squat lobster Munidopsis lauensis) living in deep-sea hydrothermal vents prioritize investing in strong claws compared to the carapace, like coastal decapods. We analyzed exoskeleton morphology, mechanical properties, structures, and elemental composition in both the carapace and claws of four Decapoda species (two each from Brachyura and Anomura infraorders) in vent and coastal habitats. Coastal decapods had ∼4-9 times more teeth on their claw cutting edge than the vent species. Further, only the coastal species exhibited higher firmness in their claws than in their carapaces. Each infraorder controlled exoskeletal hardness differently: Brachyura changed the stacking height of the Bouligand structure, while Anomura regulated magnesium content in the exoskeleton. The vent decapods may prioritize strengthening their carapace over developing robust claws, allocating resources to adapt to the harsh conditions of deep-sea hydrothermal vents. This choice might enhance their survival in the extreme environment, where carapace strength is crucial for protecting internal organs from environmental factors, rather than relying on the powerful claws seen in coastal decapods for a competitive advantage.

Hot crabs with bold choices: Temperature has little impact on behavioural repeatability in Caribbean hermit crabs.

An animal's boldness is generally considered to be influenced by genetic and developmental factors. However, abiotic factors such as temperature have profound effects on the physiology of ectothermic animals, and thus can influence the expression and measurement of this behavioural trait. We examined the relationship between temperature and behaviour in the Caribbean hermit crab (Coenobita clypeatus) using field and lab experiments. Crabs captured in the sun were bolder than crabs captured in the shade, even when measured at a common temperature, which led to bold crabs experiencing higher microhabitat temperatures. In laboratory housed conditions, crabs demonstrated highly repeatable boldness behaviours at all temperatures, and as temperature increased, the mean behavioural latencies decreased across all individuals. Bolder crabs do not seem to rely on an innately higher thermal preference, since there was no association between boldness behaviours and thermal preference in the laboratory. Instead, bolder crabs seem to exploit more open, riskier habitats than shyer crabs. Our results highlight the complex interplay between physiological and ecological factors influencing the behaviour of a widespread and ecologically important ectothermic animal.

Boldness is not associated with dynamic performance capacity in hermit crabs.

Boldness, the way an individual reacts to risk, is a commonly studied personality trait in animals. Consistent among-individual differences in startle response durations (latency to recover from a startling stimulus) are frequently assumed to reflect variation in boldness. An alternative explanation is that these latencies are not directly driven by variation in responses to information on risk, but by underlying differences in dynamic performance capacities. Here we investigate this possibility by analysing relationships between locomotory speed, a measure of whole-body dynamic performance capacity in hermit crabs, and startle response duration, a repeatable latency measure used as an index of boldness. Individuals differed in mean startle response duration, in the consistency of their startle responses, in their reaction norms across repeated observations, and mean startle responses increased with crab mass. However, there were no relationships between startle responses and locomotory speed. This indicates that startle responses do not reflect underlying performance capacities and suggests that they provide insight into differences in how individuals respond to risky situations. Since similar latencies are used as measures of boldness in other animals, we suggest that potential relationships between apparent boldness and performance capacity should be tested.

Artificial sound impact could put at risk hermit crabs and their symbiont anemones.

The sea anemone Calliactis parasitica, which is found in the East Atlantic (Portugal to Senegal) and the Mediterranean Sea, forms a symbiotic relationship with the red hermit crab, Dardanus calidus, in which the anemone provides protection from predators such as the octopus while it gains mobility, and possibly food scraps, from the hermit crab. Acoustic pollution is recognised by the scientific community as a growing threat to ocean inhabitants. Recent findings on marine invertebrates showed that exposure to artificial sound had direct behavioural, physiological and ultrastructural consequences. In this study we assess the impact of artificial sound (received level 157 ± 5 dB re 1 µPa2 with peak levels up to 175 dB re 1 µPa2) on the red hermit crab and its symbiotic sea anemone. Scanning electron microscopy analyses revealed lesions in the statocyst of the red hermit crab and in the tentacle sensory epithelia of its anemone when exposed to low-intensity, low-frequency sounds. These ultrastructural changes under situations of acoustic stress in symbiotic partners belonging to different phyla is a new issue that may limit their survival capacity, and a new challenge in assessing the effects of acoustic disturbance in the oceanic ecosystem. Despite the lesions found in the red hermit crab, its righting reflex time was not as strongly affected showing only an increase in the range of righting times. Given that low-frequency sound levels in the ocean are increasing and that reliable bioacoustic data on invertebrates is very scarce, in light of the results of the present study, we argue that anthropogenic sound effects on invertebrates species may have direct consequences in the entire ecosystem.

Behavioural adaptation to heat stress: shell lifting of the hermit crab Diogenes deflectomanus.

Behavioural responses to heat and desiccation stress in ectotherms are crucial for their survival in habitats where environmental temperatures are close to or even exceed their upper thermal limits. During low tide periods when pools in intertidal sediments heat up, a novel shell lifting behaviour (when hermit crabs crawl out of pools and lift up their shells) was observed in the hermit crab, Diogenes deflectomanus, on tropical sandy shores. On-shore measurements revealed that the hermit crabs left pools and lifted their shells predominantly when pool water exceeded 35.4 °C. Standing on emersed substrates above the pool water, the hermit crabs maintained their body temperatures at 26 - 29 °C, ∼ 10 °C lower than temperatures at which their physiological performances (as measured using heart rate) reached the maximum. This mismatch between preferred body temperatures and temperatures at maximal physiological performance was also observed under a laboratory controlled thermal gradient, where hermit crabs spent more time at 22 - 26 °C as compared to > 30 °C. These behaviours suggest a thermoregulatory function of the shell lifting behaviour, where the hermit crabs can avoid further increase in body temperatures when pools heat up during low tide periods. Such a behavioural decision allows the hermit crabs to be less prone to the strong temporal fluctuation in temperatures experienced during emersion periods on thermally dynamic tropical sandy shores.

Polarization vision in terrestrial hermit crabs.

Polarization vision is used by a wide range of animals for navigating, orienting, and detecting objects or areas of interest. Shallow marine and semi-terrestrial crustaceans are particularly well known for their abilities to detect predator-like or conspecific-like objects based on their polarization properties. On land, some terrestrial invertebrates use polarization vision for detecting suitable habitats, oviposition sites or conspecifics, but examples of threat detection in the polarization domain are less well known. To test whether this also applies to crustaceans that have evolved to occupy terrestrial habitats, we determined the sensitivity of two species of land and one species of marine hermit crab to predator-like visual stimuli varying in the degree of polarization. All three species showed an ability to detect these cues based on polarization contrasts alone. One terrestrial species, Coenobita rugosus, showed an increased sensitivity to objects with a higher degree of polarization than the background. This is the inverse of most animals studied to date, suggesting that the ecological drivers for polarization vision may be different in the terrestrial environment.

Delayed response of hermit crabs carrying anemones to a benthic impact experiment at the deep-sea nodule fields of the Peru Basin?

The deep Peru Basin is characterised by a unique abyssal scavenging community featuring large numbers of hermit crabs (Probeebei mirabilis, Decapoda, Crustacea). These are atypical hermit crabs, not carrying a shell, but on some occasions carrying an anemone (Actiniaria). The reason why some hermit crabs carry or not carry anemones is thought to be indicative of a changed environment, outweighing the cost/benefit of their relationship. Here we present the temporal variation of abundances of P. mirabilis with and without anemones, spanning more than two decades, following a benthic impact experiment. An overall decrease in hermit crab densities was observed, most noticeable and significant after 26 years and characterised by a loss of Actiniaria on the Probeebei mirabilis' pleon. Whether this is a delayed response to the benthic impact experiment carried out 26 years' prior or a natural variation in the population remains to be corroborated by an extension of the time-series. Attention is drawn to the limitations of our knowledge over time and space of the abyssal community dynamics and the urgent necessity to fill in these gaps prior to any type of deep-sea exploitation.

Caution is required when considering sentience in animals: a response to the commentary by Briffa (2022) on "Hermit crabs, shells, and sentience" (Elwood 2022).

My recent review examined the complex and intimate relationship between hermit crabs and the empty gastropod shells upon which they depend for survival. Because shells come with costs as well as benefits, the crabs are highly selective about which ones they prefer to occupy. Thus, a new shell is investigated and the information that is gathered appears to be compared with their existing shell before a decision is made. This is often prolonged and complex. Crabs also fight for shells and again the information that is gathered and used to inform fight decisions is complex. In my review, I consider these and other situations with reference to the possibility of sentience (including awareness). The excellent commentary from Mark Briffa expands on aspects of sentience and invokes the use of Lloyd Morgan's Canon to avoid suggesting complex abilities when simpler explanations would suffice. I agree with this approach. However, I also suggest that when simpler explanations appear not to explain the data, then it is reasonable to consider if more complex mental abilities might be used. I also like Briffa's suggestion that the use of an apparently higher mental faculty might result in a mechanism that is simpler than a large number of apparently simple decisions used to make complex decisions.

Carcinoecium-Forming Sea Anemone Stylobates calcifer sp. nov. (Cnidaria, Actiniaria, Actiniidae) from the Japanese Deep-Sea Floor: A Taxonomical Description with Its Ecological Observations.

Here we describe Stylobates calcifer sp. nov. (Cnidaria, Actiniaria, Actiniidae), a new carcinoecium-forming sea anemone from the deep-sea floor of Japan. Stylobates produces a carcinoecium that thinly covers the snail shells inhabited by host hermit crabs Pagurodofleinia doederleini. The new species is distinct from other species by the shape of the marginal sphincter muscle, the distribution of cnidae, the direction of the oral disk, and host association. The species' novelty is supported by the data of its mitochondrial genes 12S, 16S, and COIII and nuclear genes 18S and 28S. Also, we conducted behavioral observation of this new species, focusing on the feeding behavior and interaction with the specific host hermit crab. Our observations suggest that this sea anemone potentially feeds on the suspended particulate organic matter from the water column or the food residuals of hermit crabs. When the host's shell changed, intensive manipulation for transference of S. calcifer sp. nov. was recorded. However, although the hermit crab detached and transferred the sea anemone to the new shell after shell change, the sea anemone did not exhibit active or cooperative participation. Our data suggest that the sea anemone may not produce a carcinoecium synchronously to its host's growth, contrary to the anecdotal assumption about carcinoecium-forming sea anemones. Conversely, the host hermit crab's growth may not depend entirely on the carcinoecium produced by the sea anemone. This study is perhaps the first observation of the behavioral interaction of the rarely studied carcinoecium-forming mutualism in the deep sea.

Losers can win: Thermoregulatory advantages of regenerated claws of fiddler crab males for establishment in warmer microhabitats.

Fiddler crab males present a hypertrophied claw, which is used for sexual and aggressive displays, fights with competitors, and has been proposed as an important thermoregulatory organ for heat control. Two claw morphologies can be observed within fiddler crab populations: brachychelous claws (unregenerated) and leptochelous claws (regenerated). The leptochelous morphotype presents less muscle mass and longer fingers, resulting in a less advantageous weapon in fights. Considering their slender shape, we hypothesized that the leptochelous morphotype would present lower thermal inertia and be more efficient at body heat dissipation. We evaluated the role of the fiddler crabs' claw shape as a heat sink and how this influences their distribution between unshaded and shaded microhabitats. We tested in the field whether the proportion of adult male Leptuca uruguayensis with leptochelous claws was higher in unshaded microhabitats than shaded ones. In the laboratory, we tested if heat transfer between the body and claw is higher in leptochelous males than in brachychelous males. Spontaneous waving behaviour and active time above the surface were compared between both morphotypes in the field during the hottest period of the day. Leptuca uruguayensis with regenerated claws comprised more than 60% of the sampled male population of unshaded microhabitats compared to 18% in shaded microhabitats. Leptochelous males showed a mean heat transfer between body to claw 35% higher than that observed for brachychelous males. During the hottest period of the day, brachychelous males waved approximately 28% more than leptochelous males. Moreover, brachychelous males spent less time under the sediment surface since the surface temperature increased, while activity of leptochelous males was not associated with a temperature increase. Therefore, regenerated claws may be advantageous for the establishment of L. uruguayensis males in warmer and unshaded microhabitats because they are more efficient for heat loss and allow crabs to cool down, spending less time waving. Our study shows the relevant context of winners and losers in the face of climate change and highlights the importance of morphological variations in thermoregulatory structures for the occupation of thermal niches.

Ocean Acidification Amplifies the Olfactory Response to 2-Phenylethylamine: Altered Cue Reception as a Mechanistic Pathway?

With carbon dioxide (CO2) levels rising dramatically, climate change threatens marine environments. Due to increasing CO2 concentrations in the ocean, pH levels are expected to drop by 0.4 units by the end of the century. There is an urgent need to understand the impact of ocean acidification on chemical-ecological processes. To date, the extent and mechanisms by which the decreasing ocean pH influences chemical communication are unclear. Combining behaviour assays with computational chemistry, we explore the function of the predator related cue 2-phenylethylamine (PEA) for hermit crabs (Pagurus bernhardus) in current and end-of-the-century oceanic pH. Living in intertidal environments, hermit crabs face large pH fluctuations in their current habitat in addition to climate-change related ocean acidification. We demonstrate that the dietary predator cue PEA for mammals and sea lampreys is an attractant for hermit crabs, with the potency of the cue increasing with decreasing pH levels. In order to explain this increased potency, we assess changes to PEA's conformational and charge-related properties as one potential mechanistic pathway. Using quantum chemical calculations validated by NMR spectroscopy, we characterise the different protonation states of PEA in water. We show how protonation of PEA could affect receptor-ligand binding, using a possible model receptor for PEA (human TAAR1). Investigating potential mechanisms of pH-dependent effects on olfactory perception of PEA and the respective behavioural response, our study advances the understanding of how ocean acidification interferes with the sense of smell and thereby might impact essential ecological interactions in marine ecosystems.

Crustaceans in changing climate: Global warming and invasion of tropical land hermit crabs (Crustacea: Decapoda: Anomura: Coenobitidae) into temperate area in Japan.

There is currently a strong scientific consensus that recent global change of climate, including the global warming, seriously damage ecosystems of both lands and oceans. Here, I review recent drastic northern expansion of distribution of tropical land hermit crabs (Coenobita) into temperate Japan. Seto Marine Biological Laboratory of Kyoto University has a long history (97 years) and has been conducting various programs of long-term monitoring survey on coastal biota. A part of the results is also introduced here. Seventeen species of land hermit crabs are known from tropical regions worldwide, and seven species are known in Japan, which are mainly distributed in tropical region of Japan. Recent intensive studies on coastal areas of Japan have shown that many juveniles and small individuals of land hermit crabs are found during warm season in temperate regions. The finding of the species that were identified by DNA analysis as Coenobita rugosus and C. purpureus might be an indication for the global warming effect. Further, I here introduce the model of northern limit of distribution proposed by Gorodkov (1986) which explains change of detailed structure of population toward the limit of distribution. Moreover, I here propose the model of stages of adaptation of tropical species to temperate zone through ecological and evolutional times. These two models are useful and helpful to understand the things happen in populations in the limit of distribution and, therefore, useful for conservation of species and biological communities.

Ontogenetic changes in energetic reserves, digestive enzymes, amino acid and energy content of Lithodes santolla (Anomura: Lithodidae): Baseline for culture.

The southern king crab (SKC) Lithodes santolla is an important commercial species in southern South America. Fishing pressure has caused the deterioration of its stocks. Currently, culture techniques are being developed for producing SKC juveniles to enhance the natural population and to recover the fishing stock. Therefore, it is necessary to know about physiology, energetic and nutritional requirements for SKC maintenance in hatchery. Thus, this study aims to evaluate the biochemical and physiological changes in the midgut gland, muscle and hemolymph of juveniles, pre-adults and adults of wild SKC. The energetic reserves, digestive enzymes activity, amino acid profile and energy were quantified in twelve juveniles, ten pre-adult, and ten adult crabs. Juveniles showed high glycogen and low lipids in the midgut gland, and low proteins and low lactate in muscle. In the hemolymph, juveniles had high lipids. Pre-adults had high glycogen and lipids in the midgut gland, and both high protein and lactate in muscle. In the hemolymph, pre-adults had high lipids. Adults had low glycogen and high lipids in midgut gland, and both high proteins and high lactate in muscle. In hemolymph, adults had high glucose and lactate. Juveniles and pre-adults had high proteinase activity, whereas adults had high lipase activity. Major essential amino acids of SKC were arginine, methionine, and tryptophan, and the non-essential amino acids were glycine, aspartic acid and glutamic acid. On another hand, SKC had similar energy in the midgut gland and muscle, regardless of the ontogenetic stage. Moreover, we demonstrated that the biochemical energy calculation underestimates the actual measured values by a calorimeter. Thus, our results help to understand the physiological changes, energetic and nutritional requirements of L. santolla, and this study is a baseline for research on diet formulation for maintaining this species under culture conditions.

Origin and evolution of the symbiosis between sea anemones (Cnidaria, Anthozoa, Actiniaria) and hermit crabs, with additional notes on anemone-gastropod associations.

The anemone-crab mutualism is ubiquitous in temperate and tropical marine environments. In this symbiosis, one or more anemones live on a shell inhabited by a hermit crab and reciprocal phoretic, trophic, and defensive benefits are exchanged between the partners. Sea anemone-hermit crab symbionts belong to three families: Hormathiidae (Calliactis and Paracalliactis), Sagartiidae (Carcinactis and Verrillactis), and Actiniidae (Stylobates). Hermit crabs establish most partnerships by detaching anemones and placing them on their shell; sea anemones can also mount shells unaided, triggered by a mollusc-derived substance in the periostracum of the shell. At least partial cooperation by the anemones is necessary for successful establishment of the symbiosis. Here, we expand the evolutionary framework for hormathiid symbionts by generating a phylogeny with at least one member of each actiniarian symbiotic genus with hermit crabs using five molecular markers (16S, 12S, 18S, 28S, CO3). We not only corroborated the results from a previous study by finding two origins of hermit crab symbiosis within Hormathiidae, but also found additional origins for hermit crab symbiosis within Actiniaria. We provide for the first time evidence of a close relationship between symbionts Carcinactis dolosa and V. paguri. The ability to secrete chitin by the ectoderm of the column is inferred to be broadly convergent within Actiniaria whereas the secretion of a chitinous carcinoecium by the pedal disc is a distinct but convergent morphological adaptation of several lineages within Actiniaria. Our finding of multiple origins for both the hermit crab and gastropod symbioses suggests that the shell-mounting behavior might only have been the precursor of the hermit crab association among Calliactis spp.

Acetoin is a key odor for resource location in the giant robber crab Birgus latro.

The terrestrial and omnivorous robber crab Birgus latro inhabits islands of the Indian Ocean and the Pacific Ocean. The animals live solitarily but occasionally gather at freshly opened coconuts or fructiferous arenga palms. By analyzing volatiles of coconuts and arenga fruit, we identified five compounds, including acetoin, which are present in both food sources. In a behavioral screen performed in the crabs' habitat, a beach on Christmas Island, we found that of 15 tested fruit compounds, acetoin was the only volatile eliciting significant attraction. Hence, acetoin might play a key role in governing the crabs' aggregation behavior at both food sources.

Vision is highly sensitive to oxygen availability in marine invertebrate larvae.

For many animals, evolution has selected for complex visual systems despite the high energetic demands associated with maintaining eyes and their processing structures. Therefore, the metabolic demands of visual systems make them highly sensitive to fluctuations in available oxygen. In the marine environment, oxygen changes over daily, seasonal and inter-annual time scales, and there are large gradients of oxygen with depth. Vision is linked to survival in many marine animals, particularly among the crustaceans, cephalopods and fish, and early life stages of these groups rely on vision for prey capture, predator detection and their distribution in the water column. Using in vivo electroretinogram recordings, we show that there is a decrease in retinal sensitivity to light in marine invertebrates when exposed to reduced oxygen availability. We found a 60-100% reduction in retinal responses in the larvae of cephalopods and crustaceans: the market squid (Doryteuthis opalescens), the two-spot octopus (Octopus bimaculatus), the tuna crab (Pleuroncodes planipes) and the graceful rock crab (Metacarcinus gracilis). A decline in oxygen also decreases the temporal resolution of vision in D. opalescens These results are the first demonstration that vision in marine invertebrates is highly sensitive to oxygen availability and that the thresholds for visual impairment from reduced oxygen are species-specific. Oxygen-impaired retinal function may change the visual behaviors crucial to survival in these marine larvae. These findings may impact our understanding of species' vulnerability to ocean oxygen loss and suggest that researchers conducting electrophysiology experiments should monitor oxygen levels, as even small changes in oxygen may affect the results.

Ecological risk assessment of predicted marine invasions in the Canadian Arctic.

Climate change is impacting environmental conditions, especially with respect to temperature and ice cover in high latitude regions. Predictive models and risk assessment are key tools for understanding potential changes associated with such impacts on coastal regions. In this study relative ecological risk assessment was done for future potential introductions of three species in the Canadian Arctic: periwinkle Littorina littorea, soft shell clam Mya arenaria and red king crab Paralithodes camtschaticus. These species occur in locations connected to Canadian Arctic ports through shipping and have the potential to be introduced via ballast water discharge. The methodology proposed in this study is unique in the sense that it considers not only ballast water origin, but also the distribution of the species being assessed and the sensitivity of the receiving habitat. It combines detailed information (ballast water source of each tank, transit time, time of the year when the water is released, environmental suitability of receiving habitat, impact, and habitat sensitivity) in order to assess ecological risk. Through the use of this approach it is highlighted that domestic discharge events pose a higher relative overall risk on a vessel-specific and cumulative annual bases than international discharges. The main ports of Deception Bay and Churchill were classified as being at moderate to high relative risk for L. littorea and M. arenaria, especially from domestic vessels, while relative overall risk for P. camtschaticus was low for international vessels and null for domestic vessels due to few ships transiting from its range of distribution to Canadian Arctic ports. This work can serve as an approach to help build a list of potential high risk species-a "grey" watch list-for the Canadian Arctic, and provides useful information for consideration in future decision making actions such as the identification of high risk pathways, species and ports.

What are the sympatric mechanisms for three species of terrestrial hermit crab (Coenobita rugosus, C. brevimanus, and C. cavipes) in coastal forests?

Terrestrial hermit crabs play a significant role in coastal ecology. For example, as seed dispersers and debris scavengers in coastal forests, they accelerate the decomposition of organic substances. In the Indo-Pacific Ocean, Coenobita rugosus, C. brevimanus, and C. cavipes are the three most common species of terrestrial hermit crab. Because the mechanisms that contribute to the sympatry of these three species of crab have not been identified, this study investigated the three most likely explanations: niche differences, competition, and predation. The results showed that the three species displayed niche differences in terms of seasonal activity, habitat, utilization of shells, and food preference, suggesting that competition for resources is avoided. The habitat of terrestrial hermit crabs in Taiwan is closely associated with that of humans. Our study helps improve our understanding of the ecology of terrestrial hermit crabs and their conservation.

Morphology and Habitats of the Hermit-Crab-Associated Calyptraeid Gastropod Ergaea walshi.

Ergaea walshi, a gastropod with a markedly flat shell, often lives inside empty snail shells occupied by hermit crabs. We investigated its lifestyle, shell growth pattern, and habitat preference for host hermit crabs and host snail shells. Four hundred sixteen snail shells, including 363 shells with hermit crabs and 53 empty shells, were collected from intertidal zones of sandy and muddy flats around Kii Peninsula, Japan. The specimens comprised seven hermit crab species occupying 24 shell species; E. walshi was harbored in 13.2% of snail shells with hermit crabs and 17.0% of those without hermit crabs. Although no preference was detected for particular species of hermit crab or snail shell, E. walshi preferred to live inside of snail shells with wider apertures used by comparatively bigger hermit crabs. This suggests that the occurrence of E. walshi was influenced by host size rather than host species. When looking at growth patterns, we found that the attached shell portion of E. walshi continued to be enlarged horizontally, while growth in shell height slowed at approximately 5.0 mm. The conspicuously flattened shell of E. walshi is considered as a growth pattern for adapting to the narrow space within the snail shell occupied by hermit crabs. Consistent with this idea, our comparison of shell growth patterns in 23 calyptraeid species showed that shell of E. walshi is the flattest in this family.

Stronger positive association between an invasive crab and a native intertidal ecosystem engineer with increasing wave exposure.

Ecosystem engineers are predicted to have stronger facilitative effects when environmental stress is higher. Here we examined whether facilitation of the invasive porcelain crab Petrolisthes elongatus by the ecosystem engineering serpulid tube worm Galeolaria caespitosa increased with wave exposure. Petrolisthes occurs beneath intertidal boulders which often have a high cover of Galeolaria on their underside. Surveys across nine sites demonstrated Petrolisthes abundance beneath boulders increased with wave exposure and Galeolaria cover, although only when the habitat matrix beneath boulders was rock or mixed rock and sand. Moreover, as wave exposure increased, the strength of relationship between Petrolisthes abundance and the surface area of Galeolaria also increased. Experimentally, the presence of Galeolaria on the underside of boulders increased Petrolisthes abundance by 50% compared to boulders lacking Galeolaria. Our findings suggest the facilitative role of Galeolaria is stronger at more wave-exposed sites, which appears to contribute to a higher abundance of invasive Petrolisthes.