Effects of invasive Pacific red lionfish Pterois volitans versus a native predator on Bahamian coral-reef fish communities

The recent irruption of Pacific red lionfish (Pterois volitans) on Caribbean and Atlantic coral reefs could prove to be one of the most damaging marine invasions to date. Invasive lionfish are reaching densities much higher than those reported from their native range, and they have a strong negative effect on the recruitment and abundance of a broad diversity of native coral-reef fishes. Otherwise, little is known about how lionfish affect native coral-reef communities, especially compared to ecologically similar native predators. A controlled field experiment conducted on small patch-reefs in the Bahamas over an 8-week-period demonstrated that (1) lionfish caused a reduction in the abundance of small native coral-reef fishes that was 2.5?±?0.5 times (mean?±?SEM) greater than that caused by a similarly sized native piscivore, the coney grouper Cephalopholis fulva (93.7 vs. 36.3?% reduction); (2) lionfish caused a reduction in the species richness of small coral-reef fishes (loss of 4.6?±?1.6 species), whereas the native piscivore did not have a significant effect on prey richness; (3) the greatest effects on the reef-fish community, in terms of both abundance and richness, occurred when both native and invasive predators were present; and (4) lionfish grew significantly faster (>6 times) than the native predator under the same field conditions. These results suggest that invasive lionfish have stronger ecological effects than similarly sized native piscivores, and may pose a substantial threat to native coral-reef fish communities.     

Population dynamics of coral-reef fishes: Controversial concepts and hypotheses

Abstract Knowledge of processes that drive the local population dynamics of coral-reef fishes is important for managing reef fisheries, and for using these species as models for understanding the ecology of demersal marine fishes in general. However, the reef-fish literature is replete with poorly defined concepts and vague hypotheses regarding the issue of population dynamics. Dichotomous arguments, such as whether or not recruitment drives population dynamics, are misdirected because they fail to incorporate several important concepts. First, changes in local population size are driven by four demographic rates (birth, death, immigration and emigration), all of which must be studied to understand population dynamics. Second, all populations that persist do so because at least one of these demographic rates operates in a density-dependent way that is both sufficiently strong and appropriately time-lagged. Therefore, identifying the source(s) of direct density dependence is critical for understanding the limits to variation in population size (i.e. population regulation). Third, regulation does not imply a simple point equilibrium in population size; density dependence in populations of reef fishes is bound to lie within a field of stochastic variation, and thus be difficult to detect. Since its formal origin in 1981, the ‘recruitment limitation’ hypothesis for explaining local population dynamics in reef fishes has undergone ambiguous changes in definition that threaten its usefulness. ‘Recruitment, ‘originally defined as the appearance of newly settled fish on a reef, more recently is often measured months after settlement, thus confounding pre- and post-settlement processes. ‘Limitation, ‘ which originally referred to recruitment being so low as to preclude local populations from reaching densities where resources were limiting, is more recently defined as an absence of any form of density dependence after settlement. The most effective means of testing whether post-settlement mortality is in fact density-independent is to examine patterns of mortality directly, rather than indirectly by interpreting the shape of the relationship between initial recruit density and subsequent adult density within a cohort (the recruit-adult function). Understanding the population dynamics of coral-reef fishes will require a more equitable focus on all four demographic rates, be they density dependent or not, as well as greater attention to identifying sources of density dependence. Such a pluralistic focus necessitates integrated studies of both pre- and post-settlement processes conducted at multiple spatial and temporal scales. For example, recent evidence suggests that density-dependent pre-dation on new recruits that have settled among reefs at different densities may prove to be an important source of local population regulation, especially via the aggregative response of transient piscivores.     

Consumptive and non-consumptive effects of an invasive marine predator on native coral-reef herbivores

Invasive predators typically have larger effects on native prey populations than native predators, yet the potential roles of their consumptive versus non-consumptive effects (CEs vs. NCEs) in structuring invaded systems remains unclear. Invasive lionfish (Pterois volitans) may have ecosystem-level effects by altering native fish grazing on benthic algae that could otherwise displace corals. Lionfish could reduce grazing by decreasing the abundance of herbivorous fishes (CEs), and/or the predation risk posed by lionfish could alter grazing behavior of fishes (NCEs). To test for these CEs, we manipulated lionfish densities on large reefs in The Bahamas and surveyed fish populations throughout June 2009–2011. In July 2011, NCEs of lionfish were measured by observing fish grazing behavior on algal-covered substrata placed in microhabitats varying in lionfish presence at different spatial scales, and quantifying any resulting algal loss. Lionfish reduced small herbivorous fish density by the end of the 2010 summer recruitment season. Grazing by small and large fishes was reduced on high-lionfish-density reefs, and small fish grazing further decreased when in the immediate presence of lionfish within-reefs. Lionfish had a negative indirect effect on algal loss, with 66–80 % less algae removed from substrata in high-lionfish-density reefs. Parrotfishes were likely driving the response of herbivorous fishes to both CEs and NCEs of lionfish. These results demonstrate the importance of considering NCEs in addition to CEs of invasive predators when assessing the effects of invasions.     

Recruitment limitation: A theoretical perspective

Abstract A theoretical analysis of the concept of recruitment limitation leads to the conclusion that most populations should he regarded as jointly limited by recruitment and interactions between individuals after recruitment. The open nature of local marine systems does not permit avoidance of density-dependent interactions; it simply may make them more difficult to detect. Local populations consisting of settled organisms may not experience density-dependent interactions under some circumstances, but the entire species population consisting of the collection of local populations and their planktonic larvae must have density-dependent dynamics. Any local population of settled individuals can escape density dependence if sufficient density dependence occurs among planktonic larvae or within other local populations. Common conceptions of density dependence are too narrow, leading too often to the conclusion that it is absent from a system. It is equally wrong to expect that density-dependent interactions after settlement determine local population densities independently of recruitment. Special circumstances allowing density dependence to act strongly and quickly are needed before density dependence can neutralize the effects of recruitment. Recruitment limitation and density-dependent interactions therefore should not be regarded as alternatives but as jointly acting to determine the densities of marine benthic populations. Moreover, the interaction between fluctuating recruitment and density dependence is potentially the most interesting feature of recruitment limitation. For example, this interaction may be an important diversity-maintaining mechanism for marine systems.     

Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.     

Parasite-mediated enemy release and low biotic resistance may facilitate invasion of Atlantic coral reefs by Pacific red lionfish (<Emphasis Type="Italic">Pterois volitans</Emphasis>)

Successful invasions are largely explained by some combination of enemy release, where the invader escapes its natural enemies from its native range, and low biotic resistance, where native species in the introduced range fail to control the invader. We examined the extent to which parasites may mediate both release and resistance in the introduction of Pacific red lionfish (Pterois volitans) to Atlantic coral reefs. We found that fewer lionfish were parasitized at two regions in their introduced Atlantic range (The Bahamas and the Cayman Islands) than at two regions in their native Pacific range (the Northern Marianas Islands and the Philippines). This pattern was largely driven by relatively high infection rates of lionfish by didymozoan fluke worms and parasitic copepods (which may be host-specific to Pterois lionfishes) in the Marianas and the Philippines, respectively. When compared with sympatric, native fishes in the Atlantic, invasive lionfish were at least 18 times less likely to host a parasite in The Bahamas and at least 40 times less likely to host a parasite in the Cayman Islands. We found no indication that lionfish introduced Pacific parasites into the Atlantic. In conjunction with demographic signs of enemy release such as increased density, fish size, and growth of invasive lionfish, it is possible that escape from parasites may have contributed to the success of lionfish. This is especially true if future studies reveal that such a loss of parasites has led to more energy available for lionfish growth, reproduction, and/or immunity.     

DNA barcoding significantly improves resolution of invasive lionfish diet in the Northern Gulf of Mexico

Invasive Indo-Pacific red lionfish (Pterois volitans) have become well-established residents within reef communities across the western Atlantic Ocean where they pose substantial threats to native fish communities and reef ecosystems. Species-specific identification of prey is necessary to elucidate predator–prey interactions, but can be challenging with traditional visual identification methods given prey are often highly digested, thus not identifiable visually. To supplement visual diet analysis of lionfish (n = 934) sampled in the northern Gulf of Mexico, we applied DNA barcoding to identify otherwise unidentifiable fish prey (n = 696) via amplification of the cytochrome c oxidase subunit I (COI) of the mitochondrial genome. Barcoding nearly doubled the number of identifiable fish prey, thereby greatly enhancing our ability to describe lionfish diet. Thirty-three fish prey species were identified via barcoding, twenty-four of which were not previously detected by traditional methods. Some exploited reef fishes were newly reported (e.g., red snapper, Lutjanus campechanus) or found to constitute higher proportions of lionfish diet than previously reported (e.g., vermilion snapper, Rhomboplites aurorubens). Barcoding added a significant amount of new dietary information, and we observed the highest prey diversity reported to date for invasive lionfish. Potential cannibalism on juveniles also was identified via DNA barcoding, with the highest incidence corresponding to high lionfish densities, thus suggesting density-dependent prey demand may have driven this response. Overall, DNA barcoding greatly enhanced our ability to describe invasive lionfish diet in this study, suggesting that even studies with relatively large diet sample sizes could benefit from barcoding analysis.     

Native Predators Do Not Influence Invasion Success of Pacific Lionfish on Caribbean Reefs

Biotic resistance, the process by which new colonists are excluded from a community by predation from and/or competition with resident species, can prevent or limit species invasions. We examined whether biotic resistance by native predators on Caribbean coral reefs has influenced the invasion success of red lionfishes (Pterois volitans and Pterois miles), piscivores from the Indo-Pacific. Specifically, we surveyed the abundance (density and biomass) of lionfish and native predatory fishes that could interact with lionfish (either through predation or competition) on 71 reefs in three biogeographic regions of the Caribbean. We recorded protection status of the reefs, and abiotic variables including depth, habitat type, and wind/wave exposure at each site. We found no relationship between the density or biomass of lionfish and that of native predators. However, lionfish densities were significantly lower on windward sites, potentially because of habitat preferences, and in marine protected areas, most likely because of ongoing removal efforts by reserve managers. Our results suggest that interactions with native predators do not influence the colonization or post-establishment population density of invasive lionfish on Caribbean reefs.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Density-dependent recruitment in grassland small mammals

1. Density dependence has an important influence on the dynamics of many species of small mammals. To regulate population growth, density must affect negatively a vital rate (e.g. fecundity); however, little is known about which vital rates are most affected by density. 2. We used a long-term data set for five species of rodents from north-eastern Kansas, USA to test for relationships between density and the proportion of pregnant females, per capita fecundity and recruitment. We estimated proportion of pregnant females using data collected in the field and fecundity using data on survival, proportion of pregnant females and literature-based density-dependent litter size for each species. We used reverse capture histories to estimate per-capita recruitment. 3. The proportion of pregnant females was related positively to density in most species. Fecundity was related negatively to density in the cotton rat (Sigmodon hispidus Say & Ord, 1825) and not related to density in the four remaining species. Recruitment was related negatively to density in all five species, although not all relationships were statistically significant. 4. The signature of density-dependent recruitment was strongest in the prairie vole [Microtus ochrogaster (Wagner, 1842)] and cotton rat and less so in the remaining species. 5. Our analyses indicate that density affects recruitment negatively in grassland small mammals either through a reduction in immigration or reduced survival of nestlings. Models that seek to include empirical estimates of density dependence may need to include immigration in addition to survival and fecundity.     

Density-dependent colonization and natural disturbance limit the effectiveness of invasive lionfish culling efforts

Culling can be an effective management tool for reducing populations of invasive species to levels that minimize ecological effects. However, culling is labour-intensive, costly, and may have unintended ecological consequences. In the Caribbean, culling is widely used to control invasive Indo-Pacific lionfish, Pterois volitans and P. miles, but the effectiveness of infrequent culling in terms of reducing lionfish abundance and halting native prey decline is unclear. In a 21-month-long field experiment on natural reefs, we found that culling effectiveness changed after the passage of a hurricane part-way through the experiment. Before the hurricane, infrequent culling resulted in substantial reductions in lionfish density (60–79%, on average, albeit with large uncertainty) and slight increases in native prey species richness, but was insufficient to stem the decline in native prey biomass. Culling every 3 months (i.e., quarterly) and every 6 months (i.e., biannually) had similar effects on lionfish density and native prey fishes because of high rates of lionfish colonization among reefs. After the hurricane, lionfish densities were greater on all culled reefs compared to non-culled reefs, and prey biomass declined by 92%, and species richness by 71%, on biannually culled reefs. The two culling frequencies we examined therefore seem to offer a poor trade-off between the demonstrated conservation gains that can be achieved with frequent culling and the economy of time and money realized by infrequent culling. Moreover, stochastic events such as hurricanes can drastically limit the effectiveness of culling efforts.     

Genotyping confirms significant cannibalism in northern Gulf of Mexico invasive red lionfish, <Emphasis Type="Italic">Pterois volitans</Emphasis>

DNA barcoding is used in a variety of ecological applications to identify organisms, including partially digested prey items from diet samples. That particular application can enhance the ability to characterize diet and predator–prey dynamics but is problematic when genetic sequences of prey match those of consumer species (i.e., self-DNA). Such a result may indicate cannibalism, but false positives can result from contamination of degraded prey samples with consumer DNA. Here, nuclear-encoded microsatellite markers were used to genotype invasive lionfish, Pterois volitans, consumers and their prey (n?=?80 pairs) previously barcoded as lionfish. Cannibalism was confirmed when samples exhibited two or more different alleles between lionfish and prey DNA across multiple microsatellite loci. This occurred in 26.2% of all samples and in 42% of samples for which the data were considered conclusive. These estimates should be considered conservative given rigorous assignment criteria and low allelic diversity in invasive lionfish populations. The highest incidence of cannibalism corresponded to larger sized consumers from areas with high lionfish densities, suggesting cannibalism in northern Gulf of Mexico lionfish is size- and density-dependent. Cannibalism has the potential to influence population dynamics of lionfish which lack native western Atlantic predators. These results also have important implications for interpreting DNA barcoding analysis of diet in other predatory species where cannibalism may be underreported.     

Indo-Pacific lionfish are larger and more abundant on invaded reefs: a comparison of Kenyan and Bahamian lionfish populations

The invasion by Indo-Pacific lionfish (Pterois volitans and P. miles) of the western Atlantic, Caribbean and Gulf of Mexico is emerging as a major threat to coral reef communities across the region. Comparing native and introduced populations of invasive species can reveal shifts in ecology and behaviour that can accompany successful invasions. Using standardized field surveys replicated at multiple sites in Kenya and the Bahamas, we present the first direct comparisons of lionfish density, body size, biomass and behaviour between native and invaded coral reefs. We found that lionfish occur at higher densities with larger body sizes and total biomass on invaded Bahamian coral reefs than the ecologically equivalent species (P. miles) does on native Kenyan reefs. However, the combined average density of the five lionfish species (Pterois miles, P. antennata, P. radiata, Dendrochirus brachypterus and D. zebra) on Kenyan reefs was similar to the density of invasive lionfish in the Bahamas. Understanding the ecological processes that drive these differences can help inform the management and control of invasive lionfish.     

The history of the minnow Phoxinus phoxinus (L.) in Norway: from harmless species to pest

The causes, effects and extent of minnow Phoxinus phoxinus introductions in Norway are reviewed to assess why the introductions have had severe effects, especially where brown trout Salmo trutta is the only fish species present. The natural distribution of minnow in Norway was mainly restricted to low altitude localities in the south-eastern part of the country and in some northern areas. The distribution area expanded considerably throughout the 1900s, especially in mountain areas, due in part to the use of minnows as live bait for angling. Although minnow densities do not seem unusually high in the relatively complex fish communities of its native range, the species can achieve very high population densities when introduced to communities with few fish species, such as in the numerous recently invaded lakes where brown trout was the only fish species present. The dense minnow populations in these lakes appear to have led to reduced recruitment and growth rates in the brown trout, with abundances on average 35% lower in lakes where minnow has been introduced. The success of minnow in harsh habitats demonstrates their phenotypic and ecological plasticity, but also implies that their original distribution in Norway was restricted by early immigration history and not by environmental limitations. This suggests that human-assisted spread of the species could have strong adverse effects in Scandinavia lakes of low fish species richness.     

Density-dependent processes in the life history of fishes: evidence from laboratory populations of zebrafish Danio rerio

Population regulation is fundamental to the long-term persistence of populations and their responses to harvesting, habitat modification, and exposure to toxic chemicals. In fish and other organisms with complex life histories, regulation may involve density dependence in different life-stages and vital rates. We studied density dependence in body growth and mortality through the life-cycle of laboratory populations of zebrafish Danio rerio. When feed input was held constant at population-level (leading to resource limitation), body growth was strongly density-dependent in the late juvenile and adult phases of the life-cycle. Density dependence in mortality was strong during the early juvenile phase but declined thereafter and virtually ceased prior to maturation. Provision of feed in proportion to individual requirements (easing resource limitation) removed density dependence in growth and substantially reduced density dependence in mortality, thus indicating that 'bottom-up' effects act on growth as well as mortality, but most strongly on growth. Both growth and mortality played an important role in population regulation, with density-dependent growth having the greater impact on population biomass while mortality had the greatest impact on numbers. We demonstrate a clear ontogenic pattern of change in density-dependent processes within populations of a very small (maximum length 5 mm) fish, maintained in constant homogeneous laboratory conditions. The patterns are consistent with those distilled from studies on wild fish populations, indicating the presence of broad ontogenic patterns in density-dependent processes that are invariant to maximum body size and hold in homogeneous laboratory, as well as complex natural environments.     

Stochastic population dynamics and life-history variation in marine fish species

Abstract We examined whether differences in life-history characteristics can explain interspecific variation in stochastic population dynamics in nine marine fish species living in the Barents Sea system. After observation errors in population estimates were accounted for, temporal variability in natural mortality rate, annual recruitment, and population growth rate was negatively related to generation time. Mean natural mortality rate, annual recruitment, and population growth rate were lower in long-lived species than in short-lived species. Thus, important species-specific characteristics of the population dynamics were related to the species position along the slow-fast continuum of life-history variation. These relationships were further associated with interspecific differences in ecology: species at the fast end were mainly pelagic, with short generation times and high natural mortality, annual recruitment, and population growth rates, and also showed high temporal variability in those demographic traits. In contrast, species at the slow end were long-lived, deepwater species with low rates and reduced temporal variability in the same demographic traits. These interspecific relationships show that the life-history characteristics of a species can predict basic features of interspecific variation in population dynamical characteristics of marine fish, which should have implications for the choice of harvest strategy to facilitate sustainable yields.     

Enemy release and plant invasion: patterns of defensive traits and leaf damage in Hawaii

Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.     

They are here to stay: the biology and ecology of lionfish (Pterois miles) in the Mediterranean Sea

The lionfish, Pterois miles, is one of the most recent Lessepsian immigrants into the Mediterranean Sea, and it poses a serious threat to marine ecosystems in the region. This study assesses the basic biology and ecology of lionfish in the Mediterranean, examining morphometrics, reproduction and diet as well as population structure and distribution. The population density of lionfish has increased dramatically in Cyprus since the first sighting in late 2012; by 2018 aggregations of up to 70 lionfish were found on rocky grounds with complex reefs and artificial reefs in depths of 0–50 m. Lionfish in Cyprus become mature within a year, and adults are capable of spawning year-round, with peak spawning in summer when the sea-surface temperature reaches 28.4°C. The Cypriot lionfish grow faster and bigger than in their native range, and females are more common than males. Lionfish are generalist predators in these waters, as also found in their native range, consuming a range of teleost and crustacean prey, some of which are of high economic value (e.g., Spicara smaris and Sparisoma cretense) or have an important role in local trophic webs (e.g., Chromis chromis). Overall, the reproductive patterns, the presence of juveniles and adults throughout the year, the rapid growth rates and the generalist diet indicate that lionfish are thriving and are now already well established in the region and could potentially become the serious nuisance that they are in their temperate and tropical western Atlantic–invasive range.     

What are the characteristics of lionfish and other fishes that influence their association in diurnal refuges?

In Caribbean reefs, the lionfish Pterois volitans is an invasive species that causes severe negative ecological effects, especially as this crepuscular predator consumes very diverse prey. Lionfish are not active during the day and stay in their refuges, sharing these spaces with various other fishes. The aim of this study is to determine which fishes are associated with the lionfish in their shelters, and what characteristics of both the invasive and native species may influence and explain such coexistence between a predator and its potential prey. Through diving and snorkelling, we visited 141 lionfish refuges, mostly caves, where we observed 204 lionfish and 494 other fish from 16 native species. We recorded species and abundance, as well as lionfish size and abundance. Half of the lionfish were observed in groups and the majority were large-sized. The association with most fish species seems fortuitous, but three species, Gramma loreto, Chromis cyanea and Canthigaster rostrata, were frequently observed in association with lionfish. Numerous fish juveniles, most likely Scarus coeruleus, were also observed together with the invasive predator. The more commonly associated fishes, particularly G. loreto, are mostly associated with large-sized lionfish that were found in groups. The associated fishes are also generally found in groups. Gramma loreto is a potential cleaner of the lionfish; the reasons for the association between these fish species and the invasive lionfish may be more complex than a simple predator-prey relationship and are discussed based on their biological traits and previously reported lionfish trophic ecology and predation behaviour.