Diet of invasive lionfish on hard bottom reefs of the Southeast USA: insights from stomach contents and stable isotopes

In the paper

“Native to the subtropical and tropical regions of the South Pacific, Indian Ocean, and the Red Sea, lionfish are venomous predators whose popularity in the aquarium trade may have contributed to their introduction to Atlantic waters (Whitfield et al. 2002, Semmens et al. 2004, Ruiz-Carus et al. 2006).”

What the sources say

• Supported

  The Introduction describes lionfish as having venomous spines and a distinctive style of predation. The Method of introduction calls them popular aquarium fish and identifies aquarium release as the most likely introduction mechanism, while acknowledging ballast water as another possibility. (LLM, conf=0.93)

  cites Whitfield et al. 2002 source
• Partly supported

  The Results identify Indo-Pacific lionfish among non-native fishes sighted in the western Atlantic and say the observed species were imported for aquaria. The Discussion finds aquarium releases consistent with the introductions, supporting the claim's hedged pathway. Our copy does not describe lionfish as venomous predators or establish their species-specific popularity in the aquarium trade. (LLM, conf=0.86)

  cites Semmens et al. 2004 sourceExperimental — accuracy not yet measured
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Ruiz-Carus et al. 2006

In the paper

“The Indo-Pacific lionfish (Pterois volitans and P. miles complex) has become widely established along the United States Southeast coast and continues to colonize the Caribbean, yet its biology and ecology is only beginning to be understood.”

What the sources say

• ?Not checked — no full text of the sourcecites Morris et al. 2009 source
• Supported

  The Schofield entry in the indexed copy (pp. 473–479) says lionfish were established along the U.S. Atlantic coast from the Florida Keys to Cape Hatteras and in several Caribbean locations. It also reports sightings in additional Caribbean localities where establishment had not yet occurred, consistent with continued colonization. The cited geographical claim does not include the sentence’s separate statement about how well lionfish biology is understood. (LLM, conf=0.89)

  cites Schofield 2009 source

In the paper

“Along the United States Southeast Atlantic coast, and in Bermuda and the Bahamas, red lionfish Pterois volitans and devil firefish P. miles are now established and are continuing to expand their range in the Caribbean (Morris et al. 2009, Schofield 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Morris et al. 2009 source
• Supported

  The Schofield entry (pp. 473–479) identifies both Pterois volitans and P. miles and reports lionfish established along the southeastern U.S. Atlantic coast, in Bermuda and the Bahamas, and in several Caribbean locations. It also describes reports from additional Caribbean locations where establishment was considered likely imminent, supporting the paraphrase about range expansion. (LLM, conf=0.87)

  cites Schofield 2009 source

In the paper

“Stomach contents provided information on ingested prey, representing the animal's diet over the last few hours to days, while stable isotopes offered additional information on the trophic status of lionfish by providing a long-term integration of the diet over previous weeks to months (Cocheret de la Morinière et al. 2003).”

What the sources say

• Partly supported

  The paper’s introductory discussion (page not available in our copy) says stomach-content analysis identifies ingested prey and usually represents diet over the last few hours. It also says stable isotopes reflect diet over previous weeks to months. It does not support extending the stomach-content window to days. (LLM, conf=0.89)

  cites Cocheret de la Morinière et al. 2003 sourceExperimental — accuracy not yet measured

In the paper

“Bahamian studies documented lionfish at 5 to 177 times higher densities than in their native range (Grubich et al. 2009), capable of reducing reef fish recruitment by 79% from experimental patch reefs, and preying mostly on fishes (78% by volume), including 41 species from 21 families (Albins & Hixon 2008, Green & Cote 2009, Morris & Akins 2009).”

What the sources say

• Partly supported

  The main text reports Western Atlantic densities of about 21.2–390 lionfish per hectare and much lower estimates at Palau’s Helen Reef Atoll; those figures can yield roughly 5- to 177-fold comparisons. It does not identify the full Western Atlantic range as Bahamian or state the 5–177 figure as a finding of Bahamian studies. (LLM, conf=0.84)

  cites Grubich et al. 2009 sourceExperimental — accuracy not yet measured

In the paper

“These hard bottom reefs are well recognized as essential fish habitats that support economically valuable commercial and sport fisheries and an increasingly popular sport diving industry (Struhsaker 1969, Parker & Ross 1986), and lionfish have become widespread residents in the region (Whitfield et al. 2007, Morris et al. 2009).”

What the sources say

• ?Could not verify

  Our copy contains a correction discussing lionfish abundance in surveyed areas and the head-boat fishery, but gives no geographic distribution or evidence that lionfish were widespread residents of the Southeast US Atlantic region. Searches for established populations and geographic range returned only that correction; the relevant distribution discussion may be missing from our copy. (LLM, conf=0.74)

  cites Whitfield et al. 2007 sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites Morris et al. 2009 source

In the paper

“Bahamian studies documented lionfish at 5 to 177 times higher densities than in their native range (Grubich et al. 2009), capable of reducing reef fish recruitment by 79% from experimental patch reefs, and preying mostly on fishes (78% by volume), including 41 species from 21 families (Albins & Hixon 2008, Green & Cote 2009, Morris & Akins 2009).”

What the sources say

• ?Could not verify

  The paper’s main text reports lionfish densities in the Bahamas and cites a 79% reduction in native-fish recruitment on experimental reefs. It does not address prey composition, the 78% by-volume figure, or the count of 41 species from 21 families (searched for those figures and lionfish diet). (LLM, conf=0.76)

  cites Green & Cote 2009 sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites Morris & Akins 2009 source

In the paper

“For example, the narrower depth distribution of lionfish in northern parts of the invaded range may be related to winter water temperatures (Whitfield et al. 2002, Kimball et al. 2004, Ruiz-Carus et al. 2006).”

What the sources say

• Partly supported

  The Fig. 4 caption identifies a February 16°C isotherm as the likely inshore limit of lionfish habitat, and the Potential survival section says cold winter water likely prevents overwintering on the northeastern shelf. These passages support a possible temperature-related limit, but the paper does not establish that the depth distribution is narrower in northern than in southern parts of the invaded range. (LLM, conf=0.82)

  cites Whitfield et al. 2002 sourceExperimental — accuracy not yet measured
• Supported

  The ‘Distribution and potential overwintering range’ section (page unavailable) predicts lionfish habitat from the coast to the shelf edge farther south, but only on the mid- and outer shelf in the mid- and northern portions. It attributes the inshore and northern limits to winter temperatures, supporting the proposed explanation for the narrower northern depth range. (LLM, conf=0.91)

  cites Kimball et al. 2004 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Ruiz-Carus et al. 2006

In the paper

“Onslow Bay reef fishes include those utilizing a benthic pathway (filled symbols) and pelagic pathway (open symbols).”

What the sources say

• Supported

  The Discussion (page unavailable) states that the study distinguished three benthic-feeding species from two pelagic-feeding species. The Introduction places these reef communities in the North Carolina shelf ecosystem, including Onslow Bay. The filled and open symbols describe the citing paper’s figure, not a separate finding attributed to this source. (LLM, conf=0.91)

  cites Thomas & Cahoon (1993) source

In the paper

“Analyses of stomach contents (n = 183) and stable isotopes (n = 115) suggest a generalist carnivorous diet, and prey categories were predominately fish (~99% of total volume) from 16 families.”

In the paper

“Serranidae and Scaridae dominated the diet in 2004 while Haemulidae, Carangidae (scad and jacks), and Crustacea were important in 2006”

In the paper

“mean number of prey per stomach in 2004 vs. 2006 was 4.0 ± 0.36, n = 104 vs. 5.5 ± 1.0, n = 79 (Fig. 5C; 1-way ANOSIM, R = 0.015, p = 0.035).”

In the paper

“Unidentifiable prey fishes accounted for 45% by both volume and weight, and 57% by number.”

In the paper

“The range in δ 15 N values was greater (9.61 to 12.07 ‰), with an overall mean of 11.00 ± 0.04 ‰ (Fig. 7).”

In the paper

“We found that the smallest lionfish contained the greatest proportion by volume of crustacean prey, and higher proportions of small bodied benthic fishes such as gobies (Gobiidae) and blennies (Blenniidae)”

In the paper

“We found that prey categories that were important (sea basses, parrotfishes) in 2004 were less important in 2006 (when grunts, crustaceans, and scad were the dominant prey items).”

In the paper

“a diverse array of prey fishes from 16 different families was observed in the stomachs (3.77 ± 0.37 ml, n = 176; Table 1, Fig. 2).”

In the paper

“With increasing size, the percent volume of larger-bodied prey such as sea basses, parrotfishes, and grunts increased, along with the variety of different prey categories consumed”

In the paper

“Visual surveys of prey availability compared with prey from stomach contents revealed that grunts were observed most frequently in the environment and also in the stomachs of lionfish (Fig. 6).”

In the paper

“The δ 13 C values of lionfish muscle tissue exhibited little variability between samples, with an overall mean of -16.60 ± 0.02 ‰ (range -16.0 to -17.2 ‰).”

In the paper

“In the case of number of prey, the low global R statistic indicates that the overall distribution of prey numbers consumed between years, although statistically significant (because of large sample sizes; Warwick 1993, Warwick & Clarke 2001), was not biologically significant.”

What the sources say

• ?Not checked — no full text of the sourcecites Warwick 1993 source
• ?Not checked — no full text of the sourcecites Warwick & Clarke 2001

In the paper

“The mean prey/predator TL ratio for 2004 was 0.18 ± 0.01 (max.0.41, n = 41), compared with 0.16 ± 0.01 (max.0.25, n = 35) for 2006 and the difference was not significant (t-test, t 74 = 1.84, p = 0.07).”

In the paper

“a 47 cm (1435 g) lionfish was landed off North Carolina (J.Morris unpubl.)”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites J.Morris unpubl.

In the paper

“In addition, the average prey/predator TL ratio was slightly higher off North Carolina (0.16-0.18 vs. 0.15 in the Bahamas), though the maximum prey/predator TL ratio was higher in the Bahamas (0.48 vs. 0.25-0.41 in North Carolina) (Albins & Hixon 2008, Morris & Akins 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Albins & Hixon 2008 source
• ?Not checked — no full text of the sourcecites Morris & Akins 2009 source

In the paper

“Smaller-bodied fishes such as gobies, basslets (Grammatidae), and wrasses were ranked as the most important prey items in the Bahamas; whereas, typically larger-bodied prey (sea basses, grunts, parrotfishes) were most important off North Carolina.”

In the paper

“Classified as demersal mesocarnivores together with groupers (Serranidae) and snappers (Lutjanidae) (Caillet et al. 1986), lionfish are believed to be opportunistic predators that consume fish, shrimp, and crabs in their native range (Hiatt & Strasburg 1960, Harmelin-Vivien & Bouchon 1976, Sano et al. 1984).”

What the sources say

• ?Not checked — no full text of the sourcecites Hiatt & Strasburg 1960 source
• ?Not checked — no full text of the sourcecites Harmelin-Vivien & Bouchon 1976 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Sano et al. 1984

In the paper

“Generalist invasive predators that become established might potentially affect a wide range of species, but the risk of local extirpation for native species is thought to be lower than for specialized invasive predators.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rilov 2009

In the paper

“They have the potential to transform marine habitats worldwide, and to cause hundreds of billions of dollars in economic damages (reviews in Strayer et al. 2006, Molnar et al. 2008).”

What the sources say

• ?Not checked — no full text of the sourcecites Strayer et al. 2006 source
• ?Not checked — no full text of the sourcecites Molnar et al. 2008 source

In the paper

“However, until their recent invasion of the Atlantic and Caribbean, their biology and ecology were poorly understood beyond general descriptions because conclusions from these earlier studies were hampered by small sample sizes (n = 1 to 12; Hiatt & Strasburg 1960, Harmelin-Vivien & Bouchon 1976, Sano et al. 1984).”

What the sources say

• ?Not checked — no full text of the sourcecites Hiatt & Strasburg 1960 source
• ?Not checked — no full text of the sourcecites Harmelin-Vivien & Bouchon 1976 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Sano et al. 1984

In the paper

“To determine the ecological impacts of invasive predators, we must understand how they interact with other members of the community and how their arrival and establishment affects community structure (Kalogirou et al. 2007, Rilov 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Kalogirou et al. 2007 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rilov 2009

In the paper

“Invasive species are a major threat to biological diversity.”

What the sources say

• ?Not checked — no full text of the sourcecites Strayer et al. 2006 source
• ?Not checked — no full text of the sourcecites Molnar et al. 2008 source

In the paper

“These, in contrast, would be expected to exert stronger direct impacts on a few species and possibly cause more extensive indirect effects, depending on the ecological status of the targeted prey (Rilov 2009).”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rilov 2009

In the paper

“Predators are known to have profound effects on the structure of marine communities, and have been shown capable of enhancing species diversity (keystone species) and altering community structure (trophic cascades) (Paine 1966, Estes et al. 1998, Rilov 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Paine 1966 source
• ?Not checked — no full text of the sourcecites Estes et al. 1998 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rilov 2009

In the paper

“Classified as demersal mesocarnivores together with groupers (Serranidae) and snappers (Lutjanidae) (Caillet et al. 1986), lionfish are believed to be opportunistic predators that consume fish, shrimp, and crabs in their native range (Hiatt & Strasburg 1960, Harmelin-Vivien & Bouchon 1976, Sano et al. 1984).”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Caillet et al. 1986

In the paper

“Invasive predators have been implicated as one of the most important causes of declines and extinctions of species worldwide (Vitousek et al. 1997).”

What the sources say

• ?Not checked — no full text of the sourcecites Vitousek et al. 1997 source

In the paper

“For example, in the native range, lionfish are reported to have few natural predators (Bernadsky & Goulet 1991), to attain a size of 38 cm total length (TL), and to reach depths of 50 m (Randall et al. 1990).”

What the sources say

• ?Not checked — no full text of the sourcecites Bernadsky & Goulet 1991 source

In the paper

“The size (and economic importance) of prey consumed in the invaded range may increase, however, as increases in body size (relative to the native range) are known for a variety of invasive species, potentially including lionfish (Grosholz & Ruiz 2003).”

What the sources say

• ?Not checked — no full text of the sourcecites Grosholz & Ruiz 2003 source

In the paper

“Because collection of fishes at depth may cause regurgitation (Bowen 1992), we immediately placed speared lionfish in plastic bags to prevent loss of regurgitated stomach contents.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Bowen 1992

In the paper

“Based on laboratory studies, 80 adult lionfish along a 1 km stretch of reef in Eilat (Red Sea) were estimated to cumulatively consume approximately 230 kg of mostly small-bodied prey fishes per year (Fishelson 1997).”

What the sources say

• ?Not checked — no full text of the sourcecites Fishelson 1997 source

In the paper

“Bahamian studies documented lionfish at 5 to 177 times higher densities than in their native range (Grubich et al. 2009), capable of reducing reef fish recruitment by 79% from experimental patch reefs, and preying mostly on fishes (78% by volume), including 41 species from 21 families (Albins & Hixon 2008, Green & Cote 2009, Morris & Akins 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Albins & Hixon 2008 source

In the paper

“These depth and size extensions, as well as novel observations of predation on lionfish in the invaded range, probably reflect the paucity of investigations in the native range.”

In the paper

“These hard bottom reefs are well recognized as essential fish habitats that support economically valuable commercial and sport fisheries and an increasingly popular sport diving industry (Struhsaker 1969, Parker & Ross 1986), and lionfish have become widespread residents in the region (Whitfield et al. 2007, Morris et al. 2009).”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Struhsaker 1969
• ?Not checked — no full text of the sourcecites Parker & Ross 1986 source

In the paper

“In contrast, lionfish in the invaded range have been found in the stomach contents of piscivorous groupers (Maljkoviç et al. 2008) and have been observed to occur at greater depths (down to 304.8 m, R. Gilmore unpubl.)and attain sizes larger (47.6 cm TL; J. Morris unpubl.)than in the native range.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites R. Gilmore unpubl.

In the paper

“For example, in the native range, lionfish are reported to have few natural predators (Bernadsky & Goulet 1991), to attain a size of 38 cm total length (TL), and to reach depths of 50 m (Randall et al. 1990).”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Randall et al. 1990

In the paper

“The cumulative number of prey categories (18) recorded from these specimens had neared an asymptote and additional estimates of prey items indicated that only 1.5 to 3 prey categories may have been missed by sampling”

In the paper

“Despite recent studies of potential impacts of lionfish to tropical reef communities in the Bahamas (Albins & Hixon 2008, Morris & Akins 2009, Cote & Maljkoviç 2010), impacts to warm-temperate hard bottom reefs of the Southeast US Atlantic Ocean remain unknown.”

What the sources say

• ?Not checked — no full text of the sourcecites Albins & Hixon 2008 source
• ?Not checked — no full text of the sourcecites Morris & Akins 2009 source
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Cote & Maljkoviç 2010

In the paper

“In contrast, lionfish in the invaded range have been found in the stomach contents of piscivorous groupers (Maljkoviç et al. 2008) and have been observed to occur at greater depths (down to 304.8 m, R. Gilmore unpubl.)and attain sizes larger (47.6 cm TL; J. Morris unpubl.)than in the native range.”

What the sources say

• ?Not checked — no full text of the sourcecites Maljkoviç et al. 2008 source

In the paper

“The invaded range, and densities, size attained and depth distribution of lionfish in invaded habitats, together with laboratory studies of food consumption, all indicate the possibility for substantial impacts to both native prey communities and trophic competitors.”

In the paper

“For example, in the native range, lionfish are reported to have few natural predators (Bernadsky & Goulet 1991), to attain a size of 38 cm total length (TL), and to reach depths of 50 m (Randall et al. 1990).”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Randall et al. 1990

In the paper

“In contrast, lionfish in the invaded range have been found in the stomach contents of piscivorous groupers (Maljkoviç et al. 2008) and have been observed to occur at greater depths (down to 304.8 m, R. Gilmore unpubl.)and attain sizes larger (47.6 cm TL; J. Morris unpubl.)than in the native range.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites J. Morris unpubl.

In the paper

“A comprehensive understanding of the impacts of invasive species requires investigation of interactions at broad geographic scales (Crooks & Rilov 2009), because impacts may vary as invaders confront and respond to different ecological and physical factors in their invaded range.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Crooks & Rilov 2009

In the paper

“These differences may have been due to the availability of prey or a reflection of the sizes of lionfish collected between years.”

In the paper

“Since lionfish appear to be generalist predators that feed primarily on benthic fishes”

In the paper

“Like lionfish, serranids, haemulids, and carangids are some of the most important food items in the scamp diet”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Matheson et al. 1986

In the paper

“Predators may preferentially attack large schools if school size increases the conspicuousness and detectability of prey (Botham & Krause 2005), and not all predators succumb equally to the confusion effect attributed to schools (Turesson & Brönmark 2004).”

What the sources say

• ?Not checked — no full text of the sourcecites Botham & Krause 2005 source

In the paper

“Such an ontogenetic shift is a general pattern for numerous piscivores and can result in increased growth rates following the shift (Reñones et al. 2002, Scharf et al. 2009).”

What the sources say

• ?Not checked — no full text of the sourcecites Reñones et al. 2002 source
• ?Not checked — no full text of the sourcecites Scharf et al. 2009 source

In the paper

“Available evidence suggests that the most successful fish invaders appear to be piscivorous or omnivorous and ecological generalists (Kolar & Lodge 2001, Unmack & Fagan 2004).”

What the sources say

• ?Not checked — no full text of the sourcecites Kolar & Lodge 2001 source
• ?Not checked — no full text of the sourcecites Unmack & Fagan 2004 source

In the paper

“Predators may preferentially attack large schools if school size increases the conspicuousness and detectability of prey (Botham & Krause 2005), and not all predators succumb equally to the confusion effect attributed to schools (Turesson & Brönmark 2004).”

What the sources say

• ?Not checked — no full text of the sourcecites Turesson & Brönmark 2004 source

In the paper

“Morris & Akins (2009) found a greater number of families of fish prey (21 vs. 16 off North Carolina) in their samples but also had a larger sample size (n = 1069 vs. 183 in the present study).”

What the sources say

• ?Not checked — no full text of the sourcecites Morris & Akins 2009 source

In the paper

“When present, however, mid-water schooling herrings (Clupeidae) and fusiliers (Caesionidae) were selected over reef-associated species, similar to our observations of consumption of mid-water schooling grunts and scad.”

In the paper

“Morris and Akins (2009) examined lionfish stomachs from the Bahamas and also found that the proportion of fishes in the diet increased with lionfish size.”

What the sources say

• ?Not checked — no full text of the sourcecites Morris and Akins (2009) source

In the paper

“Unfortunately, we did not conduct prey censuses in 2004 so we do not know if these differences in prey importance between years were a response to the local availability of prey.”

In the paper

“numerous Atlantic fishes are capable of consuming venomous scorpaenids, including goosefish Lophius americanus and mutton snapper Lutjanus analis, which are known to consume the venomous scorpaenid blackbelly rosefish Helicolenus dactylopterus and spotted scorpionfish Scorpaena plumieri, respectively”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Randall 1967
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Bowman et al. 2000

In the paper

“Sustained control measures may mitigate the eventual extent of lionfish populations, but only on a local scale”

In the paper

“Piscivores are believed most capable of altering the communities which they invade”

What the sources say

• ?Not checked — no full text of the sourcecites Marchetti et al. 2004 source

In the paper

“The combined effects of overfishing, climate change, and multiple invasive species on ecosystems are not well understood but have been implicated in ecosystem state shifts and decline in several marine ecosystems”

What the sources say

• ?Not checked — no full text of the sourcecites Harris & Tyrrell 2001 source
• ?Not checked — no full text of the sourcecites Stachowicz et al. 2002 source
• ?Not checked — no full text of the sourcecites Frank et al. 2005 source