Biol Invasions (2018) 20:2343–2361

https://doi.org/10.1007/s10530-018-1705-4 (0123456789().,-volV)
(0123456789().,-volV)

ORIGINAL PAPER

Invasive Lionfish (Pterois volitans/miles) feeding ecology

in Biscayne National Park, Florida, USA
Gorka Sancho . Peter R. Kingsley-Smith . James A. Morris Jr. .
C. Anna Toline . Vanessa McDonough . Sarah M. Doty

Received: 28 August 2017 / Accepted: 13 March 2018 / Published online: 20 April 2018
Ó Springer International Publishing AG, part of Springer Nature 2018

Abstract 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, opportunis-
tic 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 con-
tents, supplemented with DNA barcoding for identi-
fication of highly-digested prey items, Lionfish diet

G. Sancho (&) V. McDonough

Department of Biology, College of Charleston, 205 Fort Biscayne National Park, 9700 SW 328th Street,
Johnson Road, Charleston, SC 29412, USA Homestead, FL 33033, USA
e-mail: sanchog@cofc.edu e-mail: vanessa_mcdonough@nps.gov

P. R. Kingsley-Smith S. M. Doty
SCDNR Marine Resources Research Institute, 217 Fort Department of Science and Mathematics, Trident
Johnson Rd, Charleston, SC 29422-2559, USA Technical College, 7000 N Rivers Ave, North Charleston,
e-mail: kingsleysmithp@dnr.sc.gov SC 29406, USA
e-mail: sarah.doty@tridenttech.edu
J. A. Morris Jr.
National Centers for Coastal Ocean Science, National
Ocean Service, NOAA, 101 Pivers Island Rd, Beaufort,
NC 28516, USA
e-mail: james.morris@noaa.gov

C. A. Toline
Hollings Marine Laboratory, National Park Service,
Southeast Region, 331 Fort Johnson Road, Charleston,
SC 29412, USA
e-mail: catherine_toline@nps.gov

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was analyzed and compared among fish sizes (imma-
ture, 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 crus-
taceans, 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
(C 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 eco-
nomic consequences for BNP and south Florida
coastal habitats.
Keywords Invasive species
Lionfish
Coral reef

Diet
Stomach contents
Introduction
Lionfishes are native to the tropical Indo-Pacific, but
over the last three decades, a complex of two sister
species, Pterois volitans (Linnaeus, 1758) and P. miles
(Bennett, 1828) (hereafter referred to collectively as
‘‘Lionfish’’), have become established in the temperate
to tropical regions of the western North Atlantic
Ocean. The Lionfish’s geographic range now includes
the southeastern coast of the U.S., Bermuda, the Gulf
of Mexico, the Caribbean Sea, and the northern coast
of South America (Whitfield et al. 2002; Meister et al.
2005; Schofield 2010; Santander-Monsalvo et al.
2012; Fortunato and Avigliano 2014). Additionally,
Lionfish have the potential to spread along the eastern
coast of South America into Brazil and Uruguay
(Kimball et al. 2004; Morris and Whitfield 2009;
Schofield 2010; Luiz et al. 2013). Individuals have
been recorded as far north as Rhode Island, however,
due to intolerance of temperatures below 10 °C, the
predominant range for Lionfish is generally consid-
ered to be south of Cape Hatteras, North Carolina. In
addition to a broad geographic distribution, Lionfish
are habitat generalists and have been documented in
estuaries (Jud et al. 2011), mangroves (Barbour et al.
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G. Sancho et al.
2010), seagrass beds (Claydon et al. 2012), anthro-
pogenic structures (e.g., Smith 2010; Jud et al. 2011;
Toledo-Hernández et al. 2014), hard bottom habitats
(Whitfield et al. 2002), coral reefs (Green and Côté
2009), and artificial reefs (Dahl et al. 2016), at depths
from \ 1 to [ 300 m (Morris and Green 2012).
The invasive Lionfish population has increased
rapidly in the last decade. Densities in the Atlantic, on
average, are 15 times higher than in the Indo-Pacific
(Hackerott et al. 2013), causing significant declines
([ 90%) in reef fish recruitment rates and biomass on
some reefs (Albins and Hixon 2008; Green et al. 2012;
Albins 2013). Bioenergetics modeling that considers
temperature-specific Lionfish growth rates and
observed densities has provided estimates of prey
biomass reductions of over 900 kg of prey per hectare
per year on reefs in the Bahamas (Cerino et al. 2013).
Further, a shift from coral to algal dominated
communities in the Bahamas has been correlated with
the introduction of Lionfish and attributed to their
predation on herbivorous fishes (Lesser and Slattery
2011).
Invasive Lionfish in the Atlantic exhibit higher diet
diversity than in the Pacific Ocean, even though
Pacific reefs generally have higher reef fish diversity
than Atlantic reefs (Cure et al. 2012). Lionfish in the
Atlantic are generalist, opportunistic predators, con-
suming multiple families of small-bodied fishes and
crustaceans (Morris and Akins 2009; Muñoz et al.
2011; Valdez-Moreno et al. 2012; Côté et al. 2013).
Previous invasive Lionfish diet studies have cumula-
tively identified more than 80 different species of
fishes in 29 different families as Lionfish prey,
consisting mostly of small-bodied cryptic benthic
fishes (e.g., gobies and blennies) and small demersal
reef fishes (e.g., wrasses, damselfishes, and parrot-
fishes) (Morris and Akins 2009; Muñoz et al. 2011;
Valdez-Moreno et al. 2012; Côté et al. 2013). Juve-
niles of economically-important fish species, such as
Yellowtail Snapper Ocyurus chrysurus (Bloch, 1791),
Nassau Grouper Epinephelus striatus (Bloch, 1792),
and Graysby Grouper Cephalopholis cruentata (La-
cepède, 1802), have been found in Lionfish stomach
contents, albeit in low numbers and infrequently
(Morris and Akins 2009; Valdez-Moreno et al. 2012;
Côté et al. 2013). Examination of trait-based prey
selection revealed that small, shallow-bodied, solitary,
demersal fishes are most vulnerable to Lionfish
predation (Green and Côté 2014). Although
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park
invertebrate prey have received far less attention, ten
different families in three separate phyla (Echinoder-
mata, Mollusca, and Arthropoda) have been identified
as Lionfish prey, the majority comprising decapod
crustaceans (Morris and Akins 2009; Muñoz et al.
2011; Valdez-Moreno et al. 2012). Consumption of
invertebrates by invasive Lionfish is inversely related
to predator size, suggesting a possible ontogenetic diet
shift from invertebrate prey to fish prey with increas-
ing predator size (Morris and Akins 2009; Muñoz et al.
2011).
This study assessed Lionfish diet in Biscayne
National Park (hereafter BNP) located on the Atlantic
coast of Florida. BNP is located in southeast Florida,
extending nearly 22 miles from Key Biscayne, just
south of Miami, to near Key Largo (Leynes and
Cullison 1998; National Park Service 2011). The Park
was established in 1980 to preserve and protect the
natural resources of the diverse area of Biscayne Bay
and the northern Florida Keys (Leynes and Cullison
1998; McCreedy et al. 2012). Nearly 95% of BNP’s
more than 170,000 acres is comprised of four main
marine habitats: (1) shoreline mangrove swamps; (2)
shallow waters and seagrass beds; (3) coral limestone
key islands; and (4) the northern portion of the Florida
Reef, the world’s third largest coral reef system
(Kellison et al. 2012; McCreedy et al. 2012). Accord-
ingly, BNP supports commercial and recreational
fishing and marine tourism. BNP hosted more than
514,709 visitors in 2012 that generated $29 million of
revenue and supported 375 jobs in the surrounding
communities (Cullinane Thomas et al. 2014). There-
fore, the introduction of invasive Lionfish is not only
of concern in the context of the ecology of BNP, but
also in terms of the surrounding south Florida
economy.
Despite BNP’s proximity (\ 30 nautical miles) to
the first Lionfish sighting in Dania Beach, FL in 1985,
invasive Lionfish were not reported in BNP until 2009
(McCreedy et al. 2012). Extensive Lionfish collection
by BNP staff and volunteers began in 2010 (McCreedy
et al. 2012). Between July 2010 and April 2017, a total
of 6994 Lionfish were removed from the Park, ranging
in size from 32 mm to 444 mm total length. The
National Park Service (NPS) developed a Lionfish
Response Plan, aimed at addressing its impacts in
multiple marine parks, that sets forth guidelines for the
management of Lionfish within the various NPS units,
including collection of Lionfish specimens,
2345
cooperation with other entities (e.g., other govern-
mental agencies, non-governmental organizations,
and universities) for research, and education of visitors
and the public (McCreedy et al. 2012). To better
understand the direct impacts that invasive Lionfish
are having within the boundaries of BNP, the current
study’s objectives were to: (1) describe the overall diet
composition of Lionfish in BNP; (2) define variations
in diet explained by Lionfish size, park region, and
season; and (3) estimate annual consumption of prey
by Lionfish.
Materials and methods
Lionfish collection and morphological prey
identification
Lionfish were collected every month for 2 years (June
2010–June 2012) within BNP (Fig. 1) by National
Park Service personnel and interns. Specimens were
collected from multiple natural and artificial habitats
(e.g., coral reefs, patch reefs, seagrass beds, shipwreck
sites, seawalls, dock pilings) ranging in depth from 3
to 50 m while snorkeling or SCUBA diving using
hand nets and pole spears. Captured Lionfish were
stored in individual bags with collection information
(i.e., site, date, habitat type, and depth), placed on ice
for the duration of the field day (08:00–15:00), and
either immediately processed in the field or frozen for
later processing. During processing, Lionfish speci-
mens were weighed (total wet weight, measured to
nearest 0.01 g) and measured (total length, TL,
measured to nearest 1 mm) and stomachs were
removed. The total content of each stomach was
weighed and preserved in 95% EtOH in individual
specimen vials. The preserved stomach contents of
513 Lionfish were analyzed. Prey items were visually
identified to the lowest possible taxa using a dissecting
microscope and published taxonomic keys and iden-
tification guides. Individual prey items were counted,
measured (to nearest 0.1 mm TL for fishes, carapace
length for crustaceans), lightly blotted dry, and
weighed (to nearest 0.01 g). No adjustments of prey
lengths due to digestion were performed.
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Fig. 1 Map of Biscayne National Park (BNP) outlining the bay, shelf, and edge regions, and marking Lionfish collection sites
DNA barcoding prey identification
DNA barcoding analysis was performed on a subset of
prey items with the following goals in mind: (1) to
verify visual morphological identifications (‘‘vou-
cher’’ specimens, n = 12; 10 fishes and 2 shrimp);
(2) to validate and identify to species level econom-
ically-important prey items (‘‘validation’’ specimens,
n = 24; 20 fishes, 3 crabs, and 1 shrimp); and (3) to
reduce the percentage of unidentified fish prey items
(‘‘identification’’ specimens, n = 207). The latter
‘‘identification’’ specimens were randomly chosen
from the pool of unidentified fish prey items that
individually weighed at least 0.01 g and collectively
comprised * 34% of the total unidentified fish prey
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G. Sancho et al.
items. Cumulative prey curves were calculated to
examine the effect of genetic barcoding identification
on sample size sufficiency (Colwell and Coddington
1994; Bizzarro et al. 2007; Colwell 2013), and as
expected curves including barcoding and morpholog-
ical identifications were farther away from reaching
asymptotes than curves including only morphological
identifications.
If the prey item was sufficiently large, 0.1–0.25 mg
of muscle tissue was removed. If the item was small
(\ 30 mg), the entire specimen was used. Dissection
tools were cleaned between samples by removing all
residual tissue, soaking the tools in 95% EtOH, and
heat sterilizing over a Bunsen burner flame. DNA was
extracted from collected tissues using NucleoSpin
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park
Tissue extraction kits (Clontech Laboratories, Inc.,
Mountain View, CA) and the 650 base-pair (bp)
barcode region of COI was amplified using PCR
following the thermal cycle and PCR reagents outlined
by Ivanova et al. (2007). The 20 ll PCR mixes
included 8.8 ll ultra-pure water, 4.0 ll 59 Taq Flexi
DNA Polymerase buffer (Promega, Madison, WI),
3.0 ll MgCl2 (25 mM; Promega, Madison, WI),
1.0 ll bovine serum albumin (Sigma-Aldrich, Inc.,
St. Louis, MO), 1.0 ll dNTP mix (25 mM; Promega,
Madison, WI), 0.5 ll each primer cocktail (10 mM),
0.2 ll Go Taq Flexi DNA Polymerase (Promega,
Madison, WI), and 1.0 ll of DNA template. PCR
products were visualized via electrophoresis on a 1.5%
agarose gel, purified with ExoSAP-IT (Affymetrix,
Santa Clara, CA), and sent to Macrogen USA (Rock-
ville, MD) for bi-directional sequencing using the
universal sequencing primers M13F and M13R for fish
specimens (see Ivanova et al. 2007) or LCO1490 and
HCO2198 for crustacean specimens (see Costa et al.
2007). Resultant sequences were trimmed, amended as
necessary, and aligned using SequencherÒ version
5.2.3 sequence analysis software (Gene Codes Cor-
poration, Ann Arbor, MI, USA, www.genecodes.
com). Final sequences were identified using the
‘‘Species Level Barcode Records’’ animal identifica-
tion function in BOLD (www.boldsystems.org; Rat-
nasingham and Herbert 2007). To avoid false
positives, a sequence similarity of greater than 99%
was used to definitively determine species identifica-
tion (Côté et al. 2013). Sequences returning no mat-
ches via the BOLD species-level database were
identified as best as possible using the ‘‘All Barcode
Records on BOLD’’ database or NCBI GenBank’s
Basic Local Alignment Search Tool (BLAST, http://
www.ncbi.nlm.nih.gov).
Influences on Lionfish diet
To examine Lionfish diet differences based on size,
categories were developed based on size-at-maturity
data from Morris (2009) in which male Lionfish were
shown to mature at approximately 100 mm TL and
female Lionfish to mature at approximately 180 mm
TL. For the present study three size classes of Lionfish
were defined: \ 100 mm TL (n = 30), 100–179 mm
TL (n = 216), and C 180 mm TL (n = 267), hereafter
referred to as ‘‘immature’’, ‘‘transitional’’, and ‘‘ma-
ture’’, respectively.
2347
To examine differences in Lionfish diet among
regions within BNP that differ in physical features and
habitat type, the Park was divided into three regions
based on geography, hydrography, and benthic habitat
type (Fig. 1). The ‘‘bay’’ area is generally shallow
(\ 4 m) and partially protected from currents and
wave action due to the presence of barrier islands
(Bernal 2014). The primary substrates in the bay are
seagrass and hard bottom, with multiple areas of
artificial, anthropogenic substrate (e.g., wrecks, house
stilts, dock pilings) distributed throughout (Ault et al.
2001). The ‘‘shelf’’ area refers to the deeper waters
(2–30 m), immediately east of the barrier keys along
the West Florida continental shelf. Currents and wave
action are stronger on the shelf due to a lack of
physical protection, and the benthos is dominated by
seagrass, but with patch reefs dispersed across areas of
unconsolidated substrate (Ault et al. 2001). The
‘‘edge’’ area refers to the waters east of the barrier
island keys along the BNP boundary and West Florida
continental shelf escarpment. Within the edge region,
depths increase dramatically as the continental shelf
slopes steeply, currents and wave action are intense,
and the benthic substrate is primarily composed of
continuous coral reef habitat, contributing to the
Florida Reef tract, and includes a number of wreck
sites (Ault et al. 2001; Kellison et al. 2012). Bernal
(2014) suggested that the western-most third of
Biscayne Bay would be the most suitable for Lionfish
occupancy according to benthic rugosity and Lionfish
physiological preferences limits (i.e., temperature,
salinity, and dissolved oxygen).
Although Lionfish were captured in every month of
the year, seasonality was defined based on wet and dry
seasons, an ecologically-relevant scale for southern
Florida and BNP. Though variable, the wet season in
south Florida is generally considered to extend from
May through October (average total precipita-
tion * 1140 mm, Florida State University 2014),
while the dry season extends from November through
April (average total precipitation * 360 mm, Florida
State University 2014).
Quantification of prey
Three relative measures of prey quantity were calcu-
lated for each prey category. Percent frequency of
occurrence (%FO) was calculated as the total number
of stomachs containing at least one item of a particular
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prey category, divided by the total number of stom-
achs containing prey, multiplied by 100 (Hyslop 1980;
Bowen 1996). The mean percent number (%MN) and
mean percent weight (%MW) for a prey category were
calculated by first calculating the percent number or
percent weight, respectively, of a prey category for
each individual stomach (by dividing the number or
weight of a prey item by the total number or weight of
all prey items in that stomach and multiplying by 100)
and then averaging these values across all stomachs
(Graham et al. 2007). This method of calculating
%MN and %MW per individual stomach avoids
sacrificial pseudoreplication from pooling across all
stomachs and allows for calculation of variation around
each metric (Ferry and Cailliet 1996; Muñoz et al.
2011). The index of relative importance (IRI) was
calculated as: IRIa = %FOa * (%MNa ? %MWa),
where a is a given prey category (Pinkas et al. 1971).
To facilitate comparison to other diet studies, IRI was
converted to percent IRI (%IRI) by dividing the IRIa for
a given prey category by the sum of the individual IRIa
values for each prey category and multiplying by 100
(Cortés 1997). Additionally, to facilitate comparison
among a wider selection of historical dietary studies,
cumulative percent number (%N) and cumulative
percent weight (%W) were also calculated by dividing
the total number or weight, respectively, of items of a
prey category by the total number or weight of all prey
items (Hyslop 1980; Bowen 1996).
Analysis of similarities (ANOSIM) tests using
%MN of total fishes and total invertebrates were
performed to examine differences in diet based on
Lionfish size, region, and season. Differences were
considered significant with p values \ 0.05 for overall
diet-by-size, diet-by-region, and diet-by-season. If
significant differences were found in overall diet-by-
size and overall diet-by-region, pairwise ANOSIMs
were performed (and significance of p values Bonfer-
roni-corrected to 0.0167) to determine which size
classes or regions differed from one another with
respect to diet.
Estimation of annual Lionfish consumption
Annual consumption of prey by Lionfish in BNP was
estimated using the Lionfish bioenergetics model
parameters provided by Cerino et al. (2013), mean
monthly sea surface temperature data for Fowey
Rocks in BNP from the National Weather Service
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G. Sancho et al.
National Data Buoy Center (National Oceanic and
Atmospheric Administration 2009), Lionfish size class
distribution observed in the current study (e.g.,
immature = 6%, transitional = 42%, mature = 52%),
and mean weights of prey types in this study derived
from the most intact specimens of each prey type, as
determined by a numerical digestive index given to
each prey item upon identification. Recruitment,
growth, and mortality information for Lionfish in
BNP was unavailable, and therefore was not incorpo-
rated, so only ‘‘instantaneous’’ annual consumption
was estimated using the consumption equation
(C = Cmax * p * f (T)) and parameters provided by
Cerino et al. (2013). Likewise, Lionfish density
estimates in BNP were also unavailable; to compen-
sate for this data gap, three hypothetical densities were
modeled based on published densities from the
invaded range: low = 30 Lionfish/ha (Bonaire; de
León et al. 2013), average = 440 Lionfish/ha (Ba-
hamas; Hackerott et al. 2013), and high = 5200
Lionfish/ha (Bahamas; Hackerott et al. 2013). While
other bioenergetics models are available, Cerino et al.
(2013) provides the most conservative results.
Results
Lionfish diet
A total of 2676 prey items were identified from the
stomach contents of Lionfish examined in this study.
These prey items were sampled from 513 Lionfish
ranging in size from 41 to 372 mm TL (mean
TL = 198 mm) and captured from 123 separate loca-
tions (Fig. 1).
Fish prey items were found in 78.9% (%FO) of the
stomachs (n = 405) and accounted for 57.5% ± 1.8
SE of the total prey by number (%MN) and
66.2% ± 1.9 SE by weight (%MW), for a combined
total of 68.2% IRI of the total prey composition
(Table 1). Forty-six fish species in 20 families were
identified. Gobiidae was the single most consumed
fish family, comprising 84% of the identified fishes by
family (all identified fish families = 19.6% IRI,
Gobiidae = 16.5% IRI) (Table 1). Other frequently
ingested fish families (by %IRI) included Pomacen-
tridae (5%), Labridae (3%), Labrisomidae (2.5%),
Blenniidae (2%), and Haemulidae (2%). The order is
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park 2349

Table 1 Identified prey categories in Lionfish stomach contents from Biscayne National Park and corresponding relative measures
of prey quantity in the overall diet
% FO %N % MN SE %W % MW SE % Total IRI % Mean IRI

Fishes 78.9 37.6 57.5 1.8 91.4 66.2 1.9 70.1 68.2
Benthopelagic fishes 21.6 6.0 11.7 1.2 50.8 16.4 1.5 19.8 7.6
Apogonidae 1.6 0.4 0.5 0.2 1.3 0.9 0.4 0.1 0.0
Carangidae 1.9 0.4 0.8 0.3 0.9 1.2 0.4 0.1 0.1
Engraulidae* 0.2 0.0 0.1 0.1 0.1 0.2 0.2 0.0 0.0
Haemulidae 3.7 1.3 2.0 0.5 5.7 2.4 0.6 0.7 0.3
Labridae 4.7 1.3 2.2 0.5 9.6 3.4 0.7 1.4 0.5
Lutjanidae 1.0 0.2 0.5 0.3 0.5 0.4 0.3 0.0 0.0
Monacanthidae 0.6 0.1 0.1 0.1 0.9 0.4 0.2 0.0 0.0
Pomacentridae 5.7 1.2 3.4 0.7 16.2 4.7 0.9 2.7 0.9
Scaridae 2.5 0.6 1.2 0.4 2.5 1.7 0.5 0.2 0.1
Sciaenidae* 0.2 0.0 0.2 0.2 0.1 0.2 0.2 0.0 0.0
Serranidae 1.4 0.3 0.6 0.3 12.9 0.9 0.4 0.5 0.0
Sparidae* 0.2 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0
Cryptobenthic fishes 33.9 12.0 18.2 1.4 22.6 24.3 1.7 18.9 18.0
Blenniidae 4.3 1.0 2.1 0.5 3.3 2.7 0.6 0.5 0.4
Callionymidae* 0.2 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0
Chaenopsidae 0.2 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0
Gobiidae 26.1 9.4 13.3 1.2 15.3 17.9 1.5 17.5 16.5
Labrisomidae 4.7 1.0 2.2 0.5 3.5 3.1 0.7 0.6 0.5
Synodontidae 0.2 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0
Tripterygiidae 1.6 0.5 0.5 0.2 0.5 0.5 0.2 0.0 0.0
Unidentified fishes 52.8 19.5 27.7 1.5 18.1 25.5 1.7 32.0 35.1
Invertebrates 61.2 62.4 42.5 1.8 8.6 31.8 1.9 29.9 31.8
Shrimp 52.8 23.2 31.3 1.6 6.4 26.1 1.8 25.1 37.9
Suborder Pleocyemata 27.1 9.6 12.4 1.1 2.2 11.4 1.3 9.2 14.1
Alpheidae 2.5 0.5 1.1 0.4 0.2 1.3 0.4 0.1 0.1
Hippolytidae 2.9 0.8 1.1 0.4 0.1 0.9 0.4 0.1 0.1
Palaemonidae 6.4 1.5 2.0 0.4 0.5 2.5 0.6 0.3 0.6
Stenopodidae 1.4 0.3 0.5 0.3 0.4 0.7 0.3 0.0 0.0
Other Pleocyemata 1.2 0.2 0.4 0.2 0.1 0.4 0.2 0.0 0.0
Unidentified Pleocyemata 18.7 6.3 7.2 0.8 0.9 5.7 0.8 3.6 4.9
Suborder Dendrobranchiata 12.9 3.7 6.0 0.9 2.7 6.0 0.9 2.3 3.4
Penaeidae* 5.7 1.3 2.5 0.6 1.5 3.2 0.7 0.4 0.7
Other Dendrobranchiata 0.4 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0
Unidentified Dendrobranchiata 9.0 2.3 3.5 0.6 1.1 2.8 0.6 0.8 1.1
Unidentified shrimp 31.6 9.8 12.9 1.0 1.5 8.7 1.1 9.7 13.8
Other invertebrates 25.0 39.2 11.2 1.1 2.2 5.7 0.9 14.8 1.4
Crabs 4.9 1.1 2.0 0.5 0.9 2.1 0.6 0.2 0.2
Identified crabs 2.3 0.5 1.0 0.4 0.7 1.3 0.5 0.1 0.1
Unidentified crabs 2.7 0.6 1.0 0.3 0.2 0.8 0.3 0.1 0.1
Unidentified decapods 0.8 0.1 0.4 0.2 0.0 0.3 0.2 0.0 0.0
Mysida 5.7 33.5 4.3 0.8 0.9 2.4 0.6 3.1 0.5
Other crustaceans 2.1 0.5 0.7 0.2 0.3 0.4 0.2 0.0 0.0

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Table 1 continued
% FO %N % MN SE %W % MW SE % Total IRI % Mean IRI

Amphipoda 0.8 0.2 0.3 0.1 0.0 0.0 0.0 0.0 0.0
Isopoda 0.6 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0
Stomatopoda 0.8 0.2 0.3 0.2 0.3 0.4 0.2 0.0 0.0
Unidentified crustaceans 12.7 3.9 3.8 0.5 0.1 0.5 0.3 0.8 0.7
Gastropoda 0.6 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0
Since percent frequency of occurrence (%FO) is a non-additive index (i.e., totals can equal [ 100%), category levels were calculated
independently (i.e., larger categories are not sums of sub-categories) (Cortés 1997). Families followed by an asterisk (*) are
previously unreported in the Lionfish diet literature. %N = percent by number, % MN = percent by mean number per stomach,
SE = standard error, %W = percent by weight, %MW = percent by mean weight per stomach, IRI = Index of Relative Importance

the same when ranking their occurrence by %FO, constituted 73% IRI of the diet of immature Lionfish
%MN, and %MW (Table 1). while fish prey constituted 83% IRI of the diet of
The identified fish prey were grouped into two main mature Lionfish (Fig. 2; ANOSIM R = 0.097,
ecological categories: (1) benthopelagic fishes (com- p \ 0.001). Benthopelagic fishes were absent from
monly found swimming in the water column just
above the seafloor or other structure); and (2) crypto- Other invertebrates Shrimps
benthic fishes (bottom-dwelling fishes that are found Unidentified fishes Cryptobenthic fishes
in direct contact with the substrate and are often Benthopelagic fishes
cryptically colored). Cryptobenthic fishes comprised A A B
18.0% IRI of the total prey composition and 70% of 100%

the identified fishes, whereas benthopelagic fishes 90%

comprised 7.6% IRI of the total prey composition and
30% of the identified fishes (Table 1). 80%

Invertebrate prey items were found in 61.2% 70%

(%FO) of stomachs (n = 314) and accounted for
42.5% ± 1.8 SE of the total prey numerically (%MN) 60%

and 31.8% ± 1.9 SE gravimetrically (%MW) for a

% IRI

50%
total of 31.8% IRI (Table 1). Other than three
gastropods, invertebrate prey items identified were 40%

crustaceans, encompassing 20 families in five orders: 30%

Amphipoda, Decapoda, Isopoda, Mysida, and Stoma-
topoda (Table 1). Decapod crustaceans comprised the 20%

majority of all invertebrate prey, and shrimp com- 10%

prised more than 96% IRI of all identified invertebrate
prey items. Twenty-eight species of shrimp in ten 0%
Immature Transitional Mature
families spanning two suborders (Dendrobranchiata < 100 mm TL 100 - 179.9 mm TL ≥ 180 mm TL
and Pleocyemata) were identified. (n=30) (n=216) (n=267)

Fig. 2 Percent Index of Relative Importance (%IRI) of main

Diet by Lionfish size
A clear transition in Lionfish diet from mainly
invertebrate prey to mainly fish prey was observed
as Lionfish size increased. Invertebrate prey
prey categories in the diet of Lionfish of different size classes
(immature, transitional, and mature) in Biscayne National Park
from June 2010–June 2012. Letter labels above columns
indicate significant differences in %MN of fish versus inverte-
brate prey as determined by pairwise ANOSIM tests (Bonfer-
roni-adjusted significant p B 0.0167)

123
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park
the diet of immature Lionfish, but comprised 12% IRI
of the identified fishes in the diet of transitional
Lionfish and 45% IRI of the identified fishes in the diet
of mature Lionfish (Fig. 2).
Diet by BNP region
The majority of immature Lionfish (83%) were caught
in the bay region of BNP, while transitional and
mature Lionfish were caught in multiple regions
across BNP, i.e., in the bay, shelf and edge reef
regions (Fig. 3). Significant differences in diet by
region (Fig. 4, ANOSIM R = 0.1095, p \ 0.001)
were observed for transitional fish, which fed mostly
on invertebrates in the bay and mostly on fishes in the
shelf and edge regions. Mature size class diet did not
differ significantly among park regions (Fig. 4,
ANOSIM R = - 0.021, p = 0.800). Fish comprised
the majority of the diet in the shelf and edge regions,
irrespective of Lionfish size class (Fig. 4).
Diet by season
Nearly equal proportions of Lionfish stomach samples
of all three size classes were examined in each of the
two seasons (immature: wet season n = 16, dry season
n = 14; transitional: wet season n = 130, dry season
n = 86; mature: wet season n = 138, dry season
n = 129). Collectively across all Lionfish size classes,
Edge Shelf Bay
100%
90%
80%
% Lionfish samples
70%
60%
50%
40%
30%
20%
10%
0%
Immature Transitional
< 100 mm TL 100 - 179.9 mm TL
(n=30) (n=216)
Fig. 3 Proportion of Lionfish samples by size class (immature,
transitional, and mature) in each Biscayne National Park (BNP)
region (i.e., bay, shelf, and edge)
2351
diet differed between seasons, with fishes being more
common in the diet during the dry season (89% IRI)
than the wet season (60% IRI; ANOSIM R = 0.014,
p = 0.011). This seasonal trend was observed in
mature size class (Fig. 5, ANOSIM R = 0.017,
p = 0.017), with immature (ANOSIM R = 0.013,
p = 0.261) and transitional (ANOSIM R = 0.001,
p = 0.323) Lionfish displaying no significant seasonal
dietary differences (Fig. 5). Prey items sampled from
mature Lionfish indicated that benthopelagic fishes
were consumed in nearly equal proportions between
the two seasons, but cryptobenthic fishes were
consumed nearly three times more in the dry season
(Fig. 5), which can largely be attributed to the more
than twofold increase in consumption of gobiids.
Conversely, dendrobranchiate shrimp were mainly
observed in stomachs during the wet season (4.0%IRI
vs. 1.1%IRI in dry season), but pleocyemate shrimp
were consumed in nearly equal proportions between
dry (1.0%IRI) and wet (1.2%IRI) seasons by mature
fish. Pleocyemate shrimp were the most important
prey item in immature Lionfish, accounting for
26.3%IRI and 37.4%IRI during the dry season and
wet season, respectively.
Genetic barcoding prey identification
After the completion of the genetic barcoding, 48% of
the total fish prey items were identified to at least
family, compared to 40% prior to barcoding. Barcod-
ing revealed seven more fish families than morpho-
logical identification alone. All dietary results
presented are cumulative, including both morpholog-
ically-based and molecular genetic-based taxonomic
identifications.
All 12 voucher specimens returned identifiable
sequences (mean sequence length ± SD = 708 ± 49
bp) and 10 of the sequences matched the morpholog-
ically-identified species (83% success rate). All 24
validation specimens returned identifiable sequences
(mean sequence length ± SD = 638 ± 74 bp).
Twenty-two of the sequences (17 fishes, 3 crabs, and
1 shrimp) matched the morphologically-identified
family and were further identified to species (87.5%
Mature
≥ 180 mm TL success rate). Of the 207 identification specimens
(n=267) sequenced, 180 returned viable sequences (mean
sequence length ± SD = 592 ± 94 bp) and 81 were
positively identified to at least genus via BOLD or
GenBank.
123
2352 G. Sancho et al.

Fig. 4 Percent Index of Other invertebrates Shrimps

Relative Importance (%IRI) Unidentified fishes Cryptobenthic fishes
of main prey categories in Benthopelagic fishes
the diet of transitional
(100–179 mm TL) and A B A A A
B
mature (C 180 mm TL) 100%
Lionfish size classes,
independently, based on
90%
region (bay, shelf, and edge)
in Biscayne National Park
from June 2010–June 2012. 80%
Letter labels above columns
indicate significant 70%
differences in %MN of fish
versus invertebrate prey as
determined by pairwise 60%
ANOSIM tests for each size
class independently
% IRI

50%
(Bonferroni-adjusted
significant p B 0.0167).
Immature Lionfish are not 40%
shown since they were
mainly captured in the bay 30%
region
20%

10%

0%

Bay Shelf Edge Bay Shelf Edge

(n=118) (n=49) (n=49) (n=29) (n=64) (n=174)

Transitional Mature

Two fish voucher specimens morphologically-
identified as Clown Wrasse Halichoeres maculipinna
(Müller & Troschel, 1848) and Peppermint Goby
Coryphopterus lipernes Böhlke & Robins, 1962
returned sequences that matched with [ 99% similar-
ity to Lionfish (P. volitans). Due to the high level of
confidence in morphologically-based identifications
of the voucher specimens, this was presumed to be the
result of contamination from Lionfish. The other 99
specimens barcoded as Lionfish were also considered
contaminated samples and remained as unidentified
fishes in the final analyses.
Estimated annual consumption by BNP Lionfish
When employing a relatively low Lionfish density
(i.e., 30 Lionfish/ha), the consumption model (Cerino
et al. 2013) estimated that the Lionfish population in
BNP would consume 28 kg of fishes and 12 kg of
invertebrates annually (Table 2). Using the mean
weight of the most intact specimens of each prey type
found in the stomach contents and the proportions of
each prey type found in the diet, the model using low
density Lionfish estimates that 55,000 prey items/ha of
the ten most consumed fish families would be
consumed annually (Table 3), including more than
40,000 gobies/ha and more than 84,000 shrimp/ha.
Employing an average density of 440 Lionfish/ha, the
model estimates an annual consumption of 414 kg of
fishes and 179 kg of invertebrates (Table 2). This
equates to Lionfish in BNP consuming more than
819,000 individual fishes/ha and more than 1.2 million
individual shrimp/ha annually (Table 3). A relatively
high density of Lionfish (5200 fish/ha) in BNP would
consume 7007 kg of prey/ha per year, including 9.6
million individual fishes of which 7.1 million would be

123
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park 2353

Fig. 5 Percent Index of Other invertebrates Shrimps

Relative Importance (%IRI)
Unidentified fishes Cryptobenthic fishes
of main prey categories in
the diet of Lionfish of all Benthopelagic fishes
size classes (immature
[\ 100 mm TL], A A A A A B
100%
transitional [100–179 mm
TL], and mature
[C 180 mm TL]), 90%
independently, based on
season (dry and wet) in 80%
Biscayne National Park
from June 2010–June 2012. 70%
Letter labels above columns
indicate significant 60%
differences in %MN of fish
% IRI

versus invertebrate prey as 50%

determined by pairwise
ANOSIM tests for each size 40%
class independently
(significant p B 0.05) 30%

20%

10%

0%
Dry Wet Dry Wet Dry Wet
(Nov-Apr) (May-Oct) (Nov-Apr) (May-Oct) (Nov-Apr) (May-Oct)
(n = 14) (n = 16) (n = 86) (n = 130) (n = 129) (n = 138)

Immature Transitional Mature

Table 2 Annual prey consumption (kg/ha) for Biscayne National Park (BNP) Lionfish modeled with three density treatments (low,
average, and high)
Model Lionfish Density Total annual consumption Annual fish consumption Annual invertebrate
density category (Lionfish/ha) (kg prey/ha) (kg prey/ha) consumption (kg prey/ha)

Low 30 40.4 28.2 12.2

Average 440 592.9 414.1 178.8
High 5200 7007.1 4894.1 2113.1
Total Annual Consumption is the sum of Annual Fish Consumption and Annual Invertebrate Consumption, each calculated using
Cerino et al.’s (2013) temperature- and Lionfish-specific consumption equation, the proportion of each Lionfish size class to the total
BNP Lionfish population, and the proportion of fishes and invertebrates in the diet of each size class (Appendix Table A3)

gobies (Table 3). More than 14.6 million individual Discussion

shrimp would be consumed per hectare annually if
BNP Lionfish density is similar to the high densities
encountered in the Bahamas (Table 3).
This study demonstrates that Lionfish are generalist,
carnivorous predators that consume a variety of small-
bodied fishes and crustaceans within the Biscayne
National Park (BNP). These results are consistent with
other invasive Lionfish diet studies from the Bahamas
(Morris and Akins 2009; Côté et al. 2013), North

123
2354 G. Sancho et al.

Table 3 Annual consumption estimates (N individual prey/ha) (low = 30 Lionfish/ha, average = 440 Lionfish/ha, high =
for the most-consumed Lionfish prey items in Biscayne 5200 Lionfish/ha) (Table 2)
National Park (BNP) modeled for three density treatments
Scientific name Avg Ind Wt (g) % Diet by % MW Annual consumption (N/ha)
Low Average High

Gobiidae 0.30 44 41,411 607,366 7,177,963

Pomacentridae 4.22 11 736 10,794 127,570
Labridae 2.51 8 900 13,199 155,986
Labrisomidae 0.92 8 2455 36,010 425,571
Blenniidae 0.65 7 3041 44,597 527,053
Haemulidae 1.83 6 926 13,577 160,461
Scaridae 0.47 4 2403 35,244 416,516
Carangidae 0.35 3 2420 35,495 419,491
Apogonidae 1.46 2 387 5673 67,042
Tripterygiidae 0.24 1 1176 17,255 203,919
Total fish 55,855 819,210 9,681,574
Pleocyemate shrimp 0.06 36 73,146 1,072,789 12,678,417
Dendrobranchiate shrimp 0.2 19 11,581 169,858 2,007,416
Unidentified shrimp 0.17 27 19,362 283,974 3,356,052
Total shrimp 84,727 1,242,647 14,685,833
Quantities are based on average weight (Avg Ind Wt, in g) of the least-digested prey type specimens found in BNP Lionfish stomach
contents and the proportion of each prey type by percent-mean-weight (% diet by %MW) in the fish or invertebrate component of
BNP Lionfish diet

Carolina (Muñoz et al. 2011), and the Mexican
Caribbean (Valdez-Moreno et al. 2012). The propor-
tion of fishes to invertebrates in the diet identified in
BNP closely resembles the proportions found in the
Bahamas and Mexico (Morris and Akins 2009;
Valdez-Moreno et al. 2012). Analysis of BNP Lionfish
diets, however, revealed previously unrecorded fish
prey, including four new families (Callionymidae,
Engraulidae, Sciaenidae, Sparidae) and 21 new
species, most of which were identified solely through
molecular genetic-based identifications based upon
DNR barcoding (Valdez-Moreno et al. 2012; Côté
et al. 2013). Nine families, five genera, and two
species of crustaceans have been previously recorded
as prey of invasive Lionfish (Morris and Akins 2009;
Valdez-Moreno et al. 2012). The present study
identified an additional 12 families, 23 genera, and
31 species of crustaceans previously unrecorded in
Lionfish diets. The increase in invertebrate identifica-
tion in the current study may be the result of sampling
smaller Lionfish and shallower habitats within BNP as
compared to Morris and Akins (2009), who primarily
sampled larger Lionfish from deeper habitats at higher
latitudes. Valdez-Moreno et al. (2012) used DNA
barcoding as the sole means of prey identification,
which resulted in greater than 60% of their inverte-
brate prey being identified only to the level of order,
possibly due to the lack of invertebrate species
barcoding data available in reference databases. The
morphological identification of invertebrate prey in
this study provides the first evidence of Lionfish
consuming portunid and xanthid crabs, and penaeid
shrimp, some of which are important fisheries species
(e.g., Florida Stone Crab, Menippe mercenaria (Say,
1818), and Pink Shrimp, Farfantepenaeus duorarum
(Burkenroad, 1939) (Florida Fish and Wildlife Con-
servation Commission 2013)).
Genetic barcoding proved to be a valuable tool for
the identification of digested prey items. All of the 7
families identified by barcoding alone were repre-
sented by only one specimen, and four of them were
previously unrecorded Lionfish prey, suggesting that
these prey are relatively rare in the diet of Lionfish and
as such may often be missed via visual analysis

123
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park
(Valdez-Moreno et al. 2012; Côté et al. 2013). Caution
should be used when extrapolating results from diets
obtained through visual identifications alone, which
may suggest more specialized Lionfish diets by
missing rare and unique prey items (Côté et al.
2013). A significant proportion of barcoding genetic
sequences in this study were identified as Lionfish,
which was treated as tissue contamination rather than
Lionfish cannibalism. This decision was made based
on the observation of voucher specimens that were
confidently visually-identified as species other than
Lionfish as well as the lack of Lionfish prey observed
through morphological identification. One potential
cause of contamination could be the small size of some
of the prey items that were barcoded. These specimens
were completely pulverized for DNA extraction since
they were too small to remove the outer layers of tissue
to ensure removal of Lionfish stomach cells. Addi-
tionally, Valdez-Moreno et al. (2012) also noted that
specimens that had been stored in ethanol for a year
prior to DNA extraction yielded poorer results than
samples from fresh specimens. Cannibalism has been
recorded in invasive Lionfish through genetic analysis
of stomach contents (Valdez-Moreno et al. 2012; Côté
et al. 2013), albeit at low frequencies (13 and 3%,
respectively). To this end, it is possible that a small
portion of the specimens identified as Lionfish but
attributed to sample contamination in our study may
indeed have been Lionfish.
The shift in Lionfish diet observed within BNP,
both in terms of prey type (from crustaceans to fishes)
and prey size (larger Lionfish consume larger prey)
with increasing Lionfish size is equivalent to the diet
shifts observed in the Bahamas and North Carolina
(Morris and Akins 2009; Muñoz et al. 2011). The shift
in prey type and size with increasing Lionfish size
represents an ontogenetic diet shift, a common pattern
among piscivorous reef fishes (e.g., Eggleston et al.
1998; Cocheret de la Morinière et al. 2003). Smaller,
immature Lionfish consumed proportionately more
shrimp than larger, mature Lionfish. Large Lionfish
consumed proportionately more benthopelagic fishes.
This dietary shift could be related to benthopelagic
fishes being generally larger in size than cryptobenthic
fishes (average prey weight = 1.31 g and 0.29 g,
respectively) and therefore exceeding the gape-size
limitation of small Lionfish, but being available and
more energy-efficient for larger Lionfish to consume.
Similarly, dendrobranchiate shrimp were only
2355
consumed by larger Lionfish, possibly due to their
larger size compared to pleocyemate shrimp (average
weight = 0.12 and 0.08 g, respectively), which were
mainly consumed only by the smaller size classes of
Lionfish.
Lionfish diet within BNP differed by park region.
Lionfish in the shallow bay region consumed more
shrimp than fishes, whereas Lionfish in the shelf and
edge regions consumed more fishes. This is likely due
to an ontogenetic shift in habitat preference, since
smaller Lionfish were mainly found in the shallow,
more protected bay region of BNP, while larger
Lionfish were mostly found in the deeper, higher
energy shelf and edge regions. Several studies have
suggested that Lionfish exhibit an ontogenetic shift in
habitat preference, settling in shallow, sheltered
mangroves and seagrass beds and later moving onto
deeper reefs once they reach larger sizes (Barbour
et al. 2010, 2011; Biggs and Olden 2011; Claydon
et al. 2012). This type of protected bay-to-reef habitat
shift is common among other reef fishes in BNP such
as Schoolmaster, Lutjanus apodus (Walbaum, 1792),
Gray Snapper, L. griseus (Linnaeus, 1758) and Great
Barracuda, Sphyraena barracuda (Edwards, 1771)
(Ault et al. 2001; Jones et al. 2010). It is unclear,
however, whether this shift is due to a change in prey
preference (e.g., shrimp to fish) or habitat preference
(e.g., shallow to deep, seagrass to reef). In the
particular case of BNP, region and habitat type are
tightly associated, with artificial habitats present in the
bay, patch reefs common in the shelf, and continuous
reefs dominating in the edge.
Adult Lionfish at BNP displayed dietary differ-
ences between wet and dry seasons, a phenomenon not
previously described at other locations within their
invasive range. Though fishes collectively dominate
adult Lionfish diet throughout the year, their propor-
tion in the diet changed seasonally, comprising 90% of
total diet in the dry season versus 60% in the wet
season. This seasonal pattern of relatively more fish
being consumed in the dry season was observed in
Lionfish of all sizes, but was most evident in larger,
mature Lionfish. Consumption of cryptobenthic fishes
(mostly gobies) in the dry season was higher than in
the wet season. By contrast, dendrobranchiate shrimp
were consumed more often by all Lionfish sizes during
the wet season than during the dry season. These
patterns of prey consumption are negatively correlated
with local seasonal recruitment and settlement of these
123
2356
prey organisms. Goby recruitment tends to peak
during the wet season (June–August) (Sponaugle
et al. 2012) and dendrobranchiate shrimp recruitment
typically peaks in the dry season (December–April)
(Criales et al. 2000). The offset in peak consumption
seasons from peak recruitment seasons may represent
a time lag, required for new recruits to grow and reach
an optimal size for Lionfish consumption.
Implications of Lionfish predation
Using the Cerino et al. (2013) Lionfish-specific
consumption model, it is estimated that Lionfish at
BNP could consume 593 kg of prey/ha, if their
densities resemble those of the Bahamas (440 Lion-
fish/ha; Hackerott et al. 2013). Our consumption
model did not include Lionfish growth throughout the
year, and is therefore a conservative estimate. In fact,
when accounting for growth, Cerino et al. (2013)
estimates that a Bahamian Lionfish population would
consume 929 kg/ha of prey annually at a density of
393 Lionfish/ha—almost 60% more than the amount
estimated without growth for a similar average density
within BNP. When converted to numbers of individual
fishes consumed, annual consumption estimates
within BNP could be more than 819,000 individual
prey fishes consumed per hectare, which would be
even higher if Lionfish growth estimates were avail-
able and taken into account.
Gobiidae was the most prevalent family of finfish
observed within Lionfish stomachs in the samples
from BNP, which reflects the patterns seen in the diet
of Lionfish in the Bahamas (Morris and Akins 2009;
Côté et al. 2013). The Bridled Goby, Coryphopterus
glaucofraenum Gill, 1863, was the single most
consumed species followed by the Glass Goby,
Coryphopterus hyalinus Böhlke & Robins, 1962, and
Peppermint Goby, Coryphopterus lipernes Böhlke &
Robins, 1962, and together they outnumbered the next
ten most consumed species combined. According to a
BNP cryptic reef fish inventory performed in
2005–2006, before Lionfish were reported in BNP,
C. glaucofraenum was a ubiquitous species within the
Park; it was documented in eight out of nine habitat
types with an estimated abundance of more than 21
individuals/100 m2 (Akins 2006). Data indicate that
C. glaucofraenum may be consumed by Lionfish more
often than other species (Morris and Akins 2009;
Albins 2013; Marsh-Hunkin et al. 2013; Pusack 2013).
123
G. Sancho et al.
In fact, Lionfish predation may even lead to local
extirpation of C. glaucofraenum (Pusack 2013).
Marsh-Hunkin et al. (2013) suggested that C. glau-
cofraenum may be more prone to the effects of
invasive Lionfish predation than an alternative simi-
larly-sized goby species such as the Goldspot Goby
Gnatholepis thompsoni Jordan, 1904 due to the lack of
an adaptive response to Lionfish behavior in C.
glaucofraenum. Additionally, C. glaucofraenum
embodies many of the traits that Green and Côté
(2014) found to be particularly vulnerable to Lionfish
predation, such as small, shallow body type and
solitary, demersal life history. Coryphopterus hyalinus
was also a fairly prevalent species within BNP and was
recorded in five of nine habitat types, with an
estimated abundance of more than 58 individuals/m2
(Akins 2006). The abundance and prevalence of these
species, and gobies in general, support the premise that
invasive Lionfish are opportunistic predators that
consume whichever prey are most available. We
recommend future periodic monitoring of gobies and
other small demersal fishes in the Park. A decline in
the abundance of the vulnerable C. glaucofraenum
would indicate an increase in the Lionfish predation
pressure, which could lead to a prey-switch to other
gobies and demersal species of similar characteristics.
Certain Lionfish prey items were colorful reef
species (e.g., damselfishes, wrasses) that many recre-
ational snorkelers and divers seek out when visiting
BNP. The 2005–2006 fish survey recorded a total of
seven species of damselfishes (Pomacentridae) and
eight species of wrasses (Labridae), of which three and
four species, respectively, were recorded in BNP
Lionfish diet (Akins 2006). The Bicolor Damselfish,
Stegastes partitus (Poey, 1868), Yellowhead Wrasse,
Halichoeres garnoti (Valenciennes, 1839) and Blue-
head Wrasse, Thalassoma bifasciatum (Bloch, 1791)
were often-consumed species by Lionfish in BNP.
Based upon the proportions of these fishes found in
their stomach contents, BNP Lionfish at the average
Bahamian Lionfish density (440 Lionfish/ha; Hack-
erott et al. 2013) can be conservatively estimated to
remove more than 10,700 damselfishes, 13,000
wrasses, and 35,000 parrotfishes per hectare annually
at BNP (Table 3). While most of these species are
ubiquitous and abundant, excessive predation by
Lionfish may drastically decrease their numbers,
which could lead to both ecological and economic
Invasive Lionfish (Pterois volitans/miles) feeding ecology in Biscayne National Park
impacts within BNP through trophic disruption and
decreased ecotourism, respectively.
By consuming benthic algae, herbivorous fishes,
such as parrotfishes (Scaridae) help to control algal
growth on coral reefs (Mumby 2006; Nemeth and
Appeldoorn 2009; Lesser and Slattery 2011). A
prominent concern associated with the Lionfish inva-
sion of the Atlantic is the potential for excessive
predation on herbivorous fishes to lead to algal
overgrowth on reefs, thereby smothering coral species,
a phenomenon that has previously been documented
by Lesser and Slattery (2011). Fourteen species of
parrotfishes and three species of surgeonfishes (Acan-
thuridae) were documented in BNP prior to the
invasion of Lionfish (BNP unpublished data), and
exhibited relatively high estimated abundances (aver-
age * 20 individuals/100 m2). Although parrotfishes
represented \ 1% of the prey fishes documented, and
no surgeonfishes were identified in Lionfish stomach
contents in this study, both of these herbivorous
families have been found in Lionfish stomach contents
in other regions of the invasive range (Morris and
Akins 2009; Muñoz et al. 2011). It would be prudent to
monitor the effects of Lionfish predation on herbiv-
orous fishes in BNP to maintain the overall health and
integrity of the reefs.
Several federally-regulated commercial fisheries
species were recorded in BNP Lionfish diets in this
study, including Bluestriped Grunt, Haemulon sciurus
(Shaw, 1803), Lane Snapper, Lutjanus synagris (Lin-
naeus, 1758), Yellowtail Snapper, Ocyurus chrysurus
(Bloch, 1791), Florida Stone Crab (Menippe merce-
naria), and Pink Shrimp (Farfantepenaeus duorarum).
These species were observed in low numbers and
frequencies, similar to what has been observed in other
Lionfish diet studies (Morris and Akins 2009; Valdez-
Moreno et al. 2012; Côté et al. 2013). This may be a
reflection of the low population sizes of many fisheries
species due to overharvest (Ault et al. 2014), as
Lionfish have been shown to have a generalist,
opportunistic diet, feeding on the most available prey
(Morris and Akins 2009; Muñoz et al. 2011; Valdez-
Moreno et al. 2012; Côté et al. 2013).
Despite the majority of species in Lionfish diets not
being defined as economically-important fisheries
species (e.g., gobies, grass shrimp), many of those
species are common prey for species that are com-
mercially important, such as snappers and groupers
(Morris and Akins 2009; Muñoz et al. 2011). Many
2357
fisheries-targeted snappers and groupers that are mid-
or top-trophic level piscivores could serve as natural
Lionfish predators in the Atlantic (Albins and Hixon
2013; Leung et al. 2011; Mumby et al. 2011). Some
predatory species present in BNP that potentially fill
this role include Lane Snapper Lutjanus synagris
(Linnaeus, 1758), Mutton Snapper Lutjanus analis
(Cuvier, 1828), Black Grouper Mycteroperca bonanci
(Poey, 1860), Red Grouper, Epinephelus morio (Va-
lenciennes, 1828), Gag Grouper Mycteroperca micro-
lepis (Goode & Bean, 1879), and Nassau Grouper
Epinephelus striatus (Bloch, 1792) (Arreguı́n-Sán-
chez and Valero 1996; Mullaney and Gale 1996;
Rivera-Arriaga et al. 1996; Eggleston et al. 1998;
Duarte and Garcia 1999; Brulé et al. 2005). Many of
these snapper and grouper species are, however,
overfished almost to the point of population collapse
in southern Florida (Ault et al. 2014), and recovery of
these stocks may be further impeded by prey compe-
tition for juveniles with Lionfish (Morris and Whitfield
2009; Paddack et al. 2009; Muñoz et al. 2011).
Lionfish also have the capacity of affecting small
predatory species through ecological competition. The
presence of Lionfish in reef areas of BNP has been
shown to alter the diet of co-existing small Graysby
Grouper, Cephalopholis cruentata (Lacepede, 1802)
though competition (Curtis et al. 2017). Several
studies have suggested that the presence of Lionfish
within shallow nursery areas (e.g., mangroves, sea-
grass beds) may be a source of early juvenile mortality
for native fish species, via either direct predation or
competition for food and/or space (Morris and Whit-
field 2009; Barbour et al. 2010, 2011; Green et al.
2012). The results of the present study suggest that
direct predation on native juvenile fishes may be
relatively low in the bay region of BNP, since smaller
Lionfish that tend to be found in the bay preyed mostly
on invertebrates. Most large adult, primarily piscivo-
rous Lionfish, were found in the shelf and edge
regions. Competition between juvenile Lionfish and
juvenile native fishes has yet to be examined, however,
and should be made a future research priority due to
low population sizes of some important fishery stocks
that utilize Biscayne Bay as nursery habitat (Ault et al.
2001; 2014).
Reductions in Lionfish density to below site-
specific threshold levels, as determined by prey
biomass, can be just as effective at minimizing
Lionfish impacts as complete eradication, which is
123
2358
likely impossible at this stage of the invasion (Côté
et al. 2014; Green et al. 2014). Site-specific culling to
threshold Lionfish densities could be applied in BNP.
Site-specific Lionfish density estimates would be
needed to determine which sites are the most heavily
populated by Lionfish, and prey biomass at Lionfish
‘‘hotspots’’ could be estimated through visual cen-
suses. Finally, threshold Lionfish densities could be
calculated on a per-site basis (Green et al. 2014).
Maintaining Lionfish densities below certain thresh-
olds can allow prey biomass to recover by at least 50%
(Green et al. 2014). This would be more practical than
complete eradication over a large geographic area
such as BNP.
Acknowledgements This study would have not been possible
without the support of the National Park Service and Lionfish
collections made by Biscayne National Park personnel. We
would also like to thank Erik Sotka and the Grice Marine
Laboratory Molecular Core Facility at the College of Charleston
for assistance with genetic barcoding, Anthony Harold from
College of Charleston for helpful guidance and Gary Sundin
from the SCDNR Shellfish Research Section for the preparation
of Fig. 1. This work was part of Sarah Doty’s graduate thesis at
the Graduate Program of Marine Biology at the College of
Charleston. The research was funded by an American Museum
of Natural History’s Lerner-Gray Grant for Marine Research, a
Joanna Deepwater Fund Fellowship and a College of Charleston
Student Assistantship to Sarah Doty, as well as internal funds
from the National Park Service and the Biology Department of
College of Charleston. This research followed the College of
Charleston Institutional Animal Care and Use Committee
(IACUC) permit number 2012–014. This publication
represents the College of Charleston Grice Marine Laboratory
contribution 506 and the South Carolina Department of Natural
Resources Marine Resources Research Institute contribution
781.
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