Grouper as a natural biocontrol of invasive lionfish

Lionfish (Pterois volitans/miles) have invaded the majority of the Caribbean region within five years. As voracious predators of native fishes with a broad habitat distribution, lionfish are poised to cause an unprecedented disruption to coral reef diversity and function. Controls of lionfish densities within its native range are poorly understood, but they have been recorded in the stomachs of large-bodied Caribbean groupers. Whether grouper predation of lionfish is sufficient to act as a biocontrol of the invasive species is unknown, but pest biocontrol by predatory fishes has been reported in other ecosystems. Groupers were surveyed along a chain of Bahamian reefs, including one of the region's most successful marine reserves which supports the top one percentile of Caribbean grouper biomass. Lionfish biomass exhibited a 7-fold and non-linear reduction in relation to the biomass of grouper. While Caribbean grouper appear to be a biocontrol of invasive lionfish, the overexploitation of their populations by fishers, means that their median biomass on Caribbean reefs is an order of magnitude less than in our study. Thus, chronic overfishing will probably prevent natural biocontrol of lionfishes in the Caribbean.     

Environmental and Biotic Correlates to Lionfish Invasion Success in Bahamian Coral Reefs

Lionfish (Pterois volitans), venomous predators from the Indo-Pacific, are recent invaders of the Caribbean Basin and southeastern coast of North America. Quantification of invasive lionfish abundances, along with potentially important physical and biological environmental characteristics, permitted inferences about the invasion process of reefs on the island of San Salvador in the Bahamas. Environmental wave-exposure had a large influence on lionfish abundance, which was more than 20 and 120 times greater for density and biomass respectively at sheltered sites as compared with wave-exposed environments. Our measurements of topographic complexity of the reefs revealed that lionfish abundance was not driven by habitat rugosity. Lionfish abundance was not negatively affected by the abundance of large native predators (or large native groupers) and was also unrelated to the abundance of medium prey fishes (total length of 5–10 cm). These relationships suggest that (1) higher-energy environments may impose intrinsic resistance against lionfish invasion, (2) habitat complexity may not facilitate the lionfish invasion process, (3) predation or competition by native fishes may not provide biotic resistance against lionfish invasion, and (4) abundant prey fish might not facilitate lionfish invasion success. The relatively low biomass of large grouper on this island could explain our failure to detect suppression of lionfish abundance and we encourage continuing the preservation and restoration of potential lionfish predators in the Caribbean. In addition, energetic environments might exert direct or indirect resistance to the lionfish proliferation, providing native fish populations with essential refuges.     

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.     

Low Susceptibility of Invasive Red Lionfish (Pterois volitans) to a Generalist Ectoparasite in Both Its Introduced and Native Ranges

Escape from parasites in their native range is one of many mechanisms that can contribute to the success of an invasive species. Gnathiid isopods are blood-feeding ectoparasites that infest a wide range of fish hosts, mostly in coral reef habitats. They are ecologically similar to terrestrial ticks, with the ability to transmit blood-borne parasites and cause damage or even death to heavily infected hosts. Therefore, being highly resistant or highly susceptible to gnathiids can have significant fitness consequences for reef-associated fishes. Indo-Pacific red lionfish (Pterois volitans) have invaded coastal habitats of the western tropical and subtropical Atlantic and Caribbean regions. We assessed the susceptibility of red lionfish to parasitic gnathiid isopods in both their native Pacific and introduced Atlantic ranges via experimental field studies during which lionfish and other, ecologically-similar reef fishes were caged and exposed to gnathiid infestation on shallow coral reefs. Lionfish in both ranges had very few gnathiids when compared with other species, suggesting that lionfish are not highly susceptible to infestation by generalist ectoparasitic gnathiids. While this pattern implies that release from gnathiid infestation is unlikely to contribute to the success of lionfish as invaders, it does suggest that in environments with high gnathiid densities, lionfish may have an advantage over species that are more susceptible to gnathiids. Also, because lionfish are not completely resistant to gnathiids, our results suggest that lionfish could possibly have transported blood parasites between their native Pacific and invaded Atlantic ranges.     

Habitat complexity and fish size affect the detection of Indo-Pacific lionfish on invaded coral reefs

A standard approach to improving the accuracy of reef fish population estimates derived from underwater visual censuses (UVCs) is the application of species-specific correction factors, which assumes that a species’ detectability is constant under all conditions. To test this assumption, we quantified detection rates for invasive Indo-Pacific lionfish (Pterois volitans and P. miles), which are now a primary threat to coral reef conservation throughout the Caribbean. Estimates of lionfish population density and distribution, which are essential for managing the invasion, are currently obtained through standard UVCs. Using two conventional UVC methods, the belt transect and stationary visual census (SVC), we assessed how lionfish detection rates vary with lionfish body size and habitat complexity (measured as rugosity) on invaded continuous and patch reefs off Cape Eleuthera, the Bahamas. Belt transect and SVC surveys performed equally poorly, with both methods failing to detect the presence of lionfish in >50 % of surveys where thorough, lionfish-focussed searches yielded one or more individuals. Conventional methods underestimated lionfish biomass by ~200 %. Crucially, detection rate varied significantly with both lionfish size and reef rugosity, indicating that the application of a single correction factor across habitats and stages of invasion is unlikely to accurately characterize local populations. Applying variable correction factors that account for site-specific lionfish size and rugosity to conventional survey data increased estimates of lionfish biomass, but these remained significantly lower than actual biomass. To increase the accuracy and reliability of estimates of lionfish density and distribution, monitoring programs should use detailed area searches rather than standard visual survey methods. Our study highlights the importance of accounting for sources of spatial and temporal variation in detection to increase the accuracy of survey data from coral reef systems.     

What Doesn't Kill You Makes You Wary? Effect of Repeated Culling on the Behaviour of an Invasive Predator

As a result of being hunted, animals often alter their behaviour in ways that make future encounters with predators less likely. When hunting is carried out for conservation, for example to control invasive species, these behavioural changes can inadvertently impede the success of future efforts. We examined the effects of repeated culling by spearing on the behaviour of invasive predatory lionfish (Pterois volitans/miles) on Bahamian coral reef patches. We compared the extent of concealment and activity levels of lionfish at dawn and midday on 16 coral reef patches off Eleuthera, The Bahamas. Eight of the patches had been subjected to regular daytime removals of lionfish by spearing for two years. We also estimated the distance at which lionfish became alert to slowly approaching divers on culled and unculled reef patches. Lionfish on culled reefs were less active and hid deeper within the reef during the day than lionfish on patches where no culling had occurred. There were no differences at dawn when removals do not take place. Lionfish on culled reefs also adopted an alert posture at a greater distance from divers than lionfish on unculled reefs. More crepuscular activity likely leads to greater encounter rates by lionfish with more native fish species because the abundance of reef fish outside of shelters typically peaks at dawn and dusk. Hiding deeper within the reef could also make remaining lionfish less likely to be encountered and more difficult to catch by spearfishers during culling efforts. Shifts in the behaviour of hunted invasive animals might be common and they have implications both for the impact of invasive species and for the design and success of invasive control programs.     

Invasive Lionfish (<Emphasis Type="Italic">Pterois volitans/miles</Emphasis>) feeding ecology in Biscayne National Park,Florida, USA

Indo-Pacific Lionfish (Pterois volitans and P. miles) are venomous marine fishes in the family Scorpaenidae that invaded the Caribbean Sea, Gulf of Mexico, and western North Atlantic Ocean beginning in the mid-1980s. Lionfish are generalist, opportunistic predators that consume a variety of invertebrates and small reef fishes, such that the presence of Lionfish can significantly reduce reef fish abundance, diversity, and recruitment on invaded reefs. This study focused on the feeding ecology of Lionfish in Biscayne National Park (BNP), located in southeast Florida, USA. BNP consists of multiple marine habitats, including mangroves, seagrass beds, coral reefs, and limestone keys that support a diverse array of species resulting in multi-million dollar fishing and tourism industries. These habitats within BNP are at risk from the predatory impacts of invasive Lionfish. Through morphological prey identification of stomach contents, supplemented with DNA barcoding for identification of highly-digested prey items, Lionfish diet was analyzed and compared among fish sizes (immature, transitional and mature), BNP region (bay, shelf, and edge), and seasons (wet and dry). A total of 513 stomachs, containing more than 2600 prey items, were examined. We report that Lionfish in BNP fed predominantly on small reef fishes and small crustaceans, with a dietary shift from crustaceans to fishes occurring with increasing Lionfish size. Diets differed among BNP regions for medium-sized (100–179 mm) transitional Lionfish but not for large-sized (≥?180 mm) mature individuals. Furthermore, dietary differences between seasons were observed in mature Lionfish, but no seasonal differences were detected for smaller Lionfish (i.e., immature and transitional Lionfish). Based on the diet habits observed, Lionfish in BNP could have significant ecological and economic consequences for BNP and south Florida coastal habitats.     

Phase shift to algal dominated communities at mesophotic depths associated with lionfish (<Emphasis Type="Italic">Pterois volitans</Emphasis>) invasion on a Bahamian coral reef

Mesophotic coral reefs (30–150 m) have been assumed to be physically and biologically connected to their shallow-water counterparts, and thus may serve as refugia for important taxonomic groups such as corals, sponges, and fish. The recent invasion of the Indo–Pacific lionfish (Pterois volitans) onto shallow reefs of the Caribbean and Bahamas has had significant, negative, effects on shallow coral reef fish populations. In the Bahamas, lionfish have extended their habitat range into mesophotic depths down to 91 m where they have reduced the diversity of several important fish guilds, including herbivores. A phase shift to an algal dominated (>50% benthic cover) community occurred simultaneously with the loss of herbivores to a depth of 61 m and caused a significant decline in corals and sponges at mesophotic depths. The effects of this invasive lionfish on mesophotic coral reefs and the subsequent changes in benthic community structure could not be explained by coral bleaching, overfishing, hurricanes, or disease independently or in combination. The significant ecological effects of the lionfish invasion into mesophotic depths of coral reefs casts doubt on whether these communities have the resilience to recover themselves or contribute to the recovery of their shallow water counterparts as refugia for key coral reef taxa.     

Habitat-Specific Density and Diet of Rapidly Expanding Invasive Red Lionfish,Pterois volitans,Populations in the Northern Gulf of Mexico

Invasive Indo-Pacific red lionfish, Pterois volitans, were first reported in the northern Gulf of Mexico (nGOM) in summer 2010. To examine potential impacts on native reef fish communities, lionfish density and size distributions were estimated from fall 2010 to fall 2013 with a remotely operated vehicle at natural (n = 16) and artificial (n = 22) reef sites. Lionfish (n = 934) also were sampled via spearfishing to examine effects of habitat type, season, and fish size on their diet and trophic ecology. There was an exponential increase in lionfish density at both natural and artificial reefs over the study period. By fall 2013, mean lionfish density at artificial reefs (14.7 fish 100 m⁻²) was two orders of magnitude higher than at natural reefs (0.49 fish 100 m⁻²), and already was among the highest reported in the western Atlantic. Lionfish diet was significantly different among habitats, seasons, and size classes, with smaller (<250 mm total length) fish consuming more benthic invertebrates and the diet of lionfish sampled from artificial reefs being composed predominantly of non-reef associated prey. The ontogenetic shift in lionfish feeding ecology was consistent with δ¹⁵N values of white muscle tissue that were positively related to total length. Overall, diet results indicate lionfish are generalist mesopredators in the nGOM that become more piscivorous at larger size. However, lionfish diet was much more varied at artificial reef sites where they clearly were foraging on open substrates away from reef structure. These results have important implications for tracking the lionfish invasion in the nGOM, as well as estimating potential direct and indirect impacts on native reef fish communities in this region.     

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.     

Larval duration of the lionfish, <Emphasis Type="Italic">Pterois volitans</Emphasis> along the Bahamian Archipelago

Larval duration of the non-native Indo-Pacific lionfish was estimated from samples collected along the Bahamian Archipelago using sagittal otoliths. Settlement marks, characterized by daily growth increments with reduced coloration, less definitive margins, and a re-orientation of the growth axes and otolith shape, were determined for 28 individuals. Settlement age was between 20 and 35 days with a mean of 26.2 days. Comparisons of settlement age to other littoral and reef fish species suggest that lionfish settlement age is moderate to relatively low. Lionfish pelagic larval duration is apparently sufficient to allow their rapid establishment and wide geographic range in the western Atlantic and Caribbean.     

Simple ecological trade-offs give rise to emergent cross-ecosystem distributions of a coral reef fish

Ecosystems are intricately linked by the flow of organisms across their boundaries, and such connectivity can be essential to the structure and function of the linked ecosystems. For example, many coral reef fish populations are maintained by the movement of individuals from spatially segregated juvenile habitats (i.e., nurseries, such as mangroves and seagrass beds) to areas preferred by adults. It is presumed that nursery habitats provide for faster growth (higher food availability) and/or low predation risk for juveniles, but empirical data supporting this hypothesis is surprisingly lacking for coral reef fishes. Here, we investigate potential mechanisms (growth, predation risk, and reproductive investment) that give rise to the distribution patterns of a common Caribbean reef fish species, Haemulon flavolineatum (French grunt). Adults were primarily found on coral reefs, whereas juvenile fish only occurred in non-reef habitats. Contrary to our initial expectations, analysis of length-at-age revealed that growth rates were highest on coral reefs and not within nursery habitats. Survival rates in tethering trials were 0% for small juvenile fish transplanted to coral reefs and 24-47% in the nurseries. As fish grew, survival rates on coral reefs approached those in non-reef habitats (56 vs. 77-100%, respectively). As such, predation seems to be the primary factor driving across-ecosystem distributions of this fish, and thus the primary reason why mangrove and seagrass habitats function as nursery habitat. Identifying the mechanisms that lead to such distributions is critical to develop appropriate conservation initiatives, identify essential fish habitat, and predict impacts associated with environmental change.     

Mangrove and coral reef sponge faunas: untold stories about shallow water Porifera in the Caribbean

Sponge faunas from coral reefs and mangrove ecosystems in the Caribbean have mostly been studied from an ecological perspective, with researchers considering the effects of physical and biological factors on their species distribution. To discern evolutionary patterns, this study analyzed the systematic composition, taxonomic diversity, and ecological properties (reproductive strategies, size, shape, endosymbiosis) of mangrove and reef sponge assemblages from seven distant Caribbean localities. Species composition was compared by use of cluster analysis (Sørensen’s), and taxonomic diversity by use of the biodiversity index average taxonomic distinctness (AvTD). Mangrove and reef-associated sponge faunas were found to be statistically dissimilar, with the AvTD values suggesting stronger taxonomic bias toward specific groups in mangroves, irrespective of geographic distance. Most Demospongiae orders have 30–50% more species in coral reefs than in mangroves. The richest reef genera (Agelas, Aplysina, Callyspongia, Petrosia, and Xestospongia) rarely colonize contiguous mangrove formations. The distribution and diversity of suprageneric taxa suggest that coral reef sponge assemblages might represent an older fauna. This historical interpretation would place mangrove subtidal habitats as the youngest marine ecosystem, rather than a below-optimum ecosystem. Life history traits support a biological split discussed here from the perspective of distinct evolutionary histories and different environmental conditions.     

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.     

Ecological impacts of an invasive mesopredator do not differ from those of a native mesopredator: lionfish in Caribbean Panama

Coral Reefs - The impacts of invasive lionfish (Pterois volitans/miles) on native coral reef populations in the Western Atlantic Ocean and Caribbean Sea can be enormous. However, how much lionfish...     

Responses of coral reef fishes to past climate changes are related to life‐history traits

Coral reefs and their associated fauna are largely impacted by ongoing climate change. Unravelling species responses to past climatic variations might provide clues on the consequence of ongoing changes. Here, we tested the relationship between changes in sea surface temperature and sea levels during the Quaternary and present‐day distributions of coral reef fish species. We investigated whether species‐specific responses are associated with life‐history traits. We collected a database of coral reef fish distribution together with life‐history traits for the Indo‐Pacific Ocean. We ran species distribution models (SDMs) on 3,725 tropical reef fish species using contemporary environmental factors together with a variable describing isolation from stable coral reef areas during the Quaternary. We quantified the variance explained independently by isolation from stable areas in the SDMs and related it to a set of species traits including body size and mobility. The variance purely explained by isolation from stable coral reef areas on the distribution of extant coral reef fish species largely varied across species. We observed a triangular relationship between the contribution of isolation from stable areas in the SDMs and body size. Species, whose distribution is more associated with historical changes, occurred predominantly in the Indo‐Australian archipelago, where the mean size of fish assemblages is the lowest. Our results suggest that the legacy of habitat changes of the Quaternary is still detectable in the extant distribution of many fish species, especially those with small body size and the most sedentary. Because they were the least able to colonize distant habitats in the past, fish species with smaller body size might have the most pronounced lags in tracking ongoing climate change.     

Life history and habitat use of Norwegian brown trout (Salmo trutta)

SUMMARY. 1. Brown trout ( Salmo trutta) life history and habitat use were studied in two Norwegian rivers: the Vosso river system, western Norway, and the Søre Osa, eastern Norway.
2. Age-groups were partly segregated in feeding habitats, the youngest fish living mainly in running water and in the littoral zone of lakes, the older fish also exploiting pelagic waters and deeper epibenthic habitats. In a population with free access to and from the sea, some individuals smoltified and became sea-run migrants, performing yearly migrations to the coastal sea, whereas others stayed as freshwater residents throughout their entire life span.
3. Within local populations, females were larger and less variable in size than males. This was partly because females matured at an older age than males, partly because the sexes tended to exploit feeding habitats with different food and growth Conditions. Within age-groups, females were more pelagic and migrated more than males, whereas males were more confined to running water and epibenthic areas than females. In the pelagic zone, males were more abundant in near-surface water, and females more abundant in deeper areas. When exploiting the same feeding areas, the two sexes grew at the same rate. There therefore appears to be a connection between feeding habitat and the reproductive ecology of brown trout.     

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.     

Lionfish in the eastern Pacific: a cellular automaton approach to assessing invasion risk

The lionfish invasion in the Atlantic and Caribbean has proceeded with vigor since their introduction in the 1980s or early 1990s. Lionfish affect recruitment of juvenile fish to reefs due to predation and are found in densities far surpassing that of their native Indo-Pacific. There is concern that the lionfish may become introduced and proliferate (through aquarium releases, transport on floating debris, or passage through the Panama Canal in ship ballast water) in the eastern tropical and north Pacific. This study presents the first known prediction of the potential for establishment of lionfish in the eastern Pacific Ocean. Through computational modeling, we compare and contrast the dynamics of random hypothetical introductions of lionfish into the eastern Pacific and Atlantic Oceans in order to highlight the different potentials for invasion in both basins. Connectivity between discrete regions (precincts) in both the Atlantic and eastern Pacific are examined and settlement densities are calculated to indicate possible locations of establishment of breeding lionfish populations. Our results suggest that lionfish, which are successful invaders in the Atlantic, may not be as successful in the eastern Pacific due to weak mesoscale connectivity which reduces the rapid spread of lionfish larvae.     

Invasive lionfish drive Atlantic coral reef fish declines

Indo-Pacific lionfish (Pterois volitans and P. miles) have spread swiftly across the Western Atlantic, producing a marine predator invasion of unparalleled speed and magnitude. There is growing concern that lionfish will affect the structure and function of invaded marine ecosystems, however detrimental impacts on natural communities have yet to be measured. Here we document the response of native fish communities to predation by lionfish populations on nine coral reefs off New Providence Island, Bahamas. We assessed lionfish diet through stomach contents analysis, and quantified changes in fish biomass through visual surveys of lionfish and native fishes at the sites over time. Lionfish abundance increased rapidly between 2004 and 2010, by which time lionfish comprised nearly 40% of the total predator biomass in the system. The increase in lionfish abundance coincided with a 65% decline in the biomass of the lionfish's 42 Atlantic prey fishes in just two years. Without prompt action to control increasing lionfish populations, similar effects across the region may have long-term negative implications for the structure of Atlantic marine communities, as well as the societies and economies that depend on them.