This document summarizes a study examining the effects of spearfishing on lionfish populations in Roatan, Honduras. The study used transect surveys, roving diver surveys, and dive shop surveys to observe 293 lionfish at sites both within and outside the Roatan Marine Park, where spearfishing efforts have focused on controlling lionfish. The results showed no significant differences in lionfish abundance or behavior between the areas. However, there was evidence that spearfishing was excluding lionfish from shallow depths and influencing the habitat use of potential lionfish predators within the marine park. Overall, the study provided preliminary evidence that regular spearfishing of lionfish may be affecting their distribution and the behavior of native predatory fish species

1. 1
Introduction
The invasion of Pterois miles (Devil Firefish) and Pterois volitans (Red Lionfish),
collectively referred to as the Indo-Pacific Lionfish, along the Atlantic Seaboard, among
Caribbean Islands, and in the Mesoamerican Barrier Reef System (MBRS) in the last 20-30 years
has significantly disrupted reef communities and population dynamics (Whitfield et al., 2002;
Morris, 2009; Morris & Whitfield, 2009; Schofield, 2009). The lionfish’s lack of natural
predators and disease, abundance of prey, reproductive qualities and cryptic, venomous nature
account for their rapid spread throughout the Caribbean (Kizer et al., 1985; Whitfield et al.,
2002; Morris & Whitfield, 2009; Aguilar-Perera & Tuz-Sulub, 2010). Furthermore, lionfish lay
large masses of floating eggs approximately every ten days which can survive for nearly a month
before settling on a reef. Together, these characteristics allow for long distance dispersal by
ocean currents and permits lionfish to expand their invasive range efficiently and quickly
(Whitfield et al., 2007; Morris, 2009; Morris et al., 2009; Ahrenholz & Morris, 2010).
Many of the Caribbean islands have had to deal with the disruption of local reef
community structure and dynamics produced by the lionfish (Morris et al., 2009; Morris &
Akins, 2009). Lionfish are voracious feeders that consume many different species of fish (>40)
and crustaceans (Morris & Akins, 2009). Their broad diet includes the juveniles of many
economically important fish (e.g. groupers; snappers; aquarium trade fish) as well as important
reef grazers that maintain reef health (e.g. parrot fish, damselfish, surgeonfish) (Albins & Hixon,
2011). Besides being efficient hunters in a new region, lionfish also appear to be effective
competitors of other predators and are themselves resistant to predation, potentially altering
natural marine systems in a manner similar to over fishing (Albins & Hixon, 2011).

2. 2
Controlling lionfish is both difficult and dangerous. Lionfish cannot be caught effectively
by traditional fishing methods with a line and net as they typically reside on reefs which would
produce high amounts of by-catch. The best known method for their capture is spearfishing
where hunters can actively target lionfish. Although spearfishing is species-specific and avoids
unwanted collateral deaths, it is also time-consuming, expensive, and often has low catch per
unit effort. It is not well known what kinds of natural predators may control lionfish populations
in their native ranges, although in the Caribbean they have been found in the stomach contents of
some upper trophic-level species (large-bodied Caribbean groupers), indicating that local
predatory fish in some areas might be creating a search image for the invaders (Morris &
Whitfield, 2009; Mumby et al., 2011). Divers in the region have also reported that beside
groupers, a few species of snapper, moray eels, and sharks have also accepted speared lionfish
allowing them to be considered as potential lionfish predators (PLPs).
In recent years Roatan, an island off the northern coast of Honduras, located at the
southern end of the MBRS, has begun to recognize the effects of the lionfish invasion on local
reef communities (Morris et al., 2009; Lesser & Slattery, 2011). The dive and conservation
communities on the island have noticed that with the increase in the amount of lionfish in the
park there has been a decrease in small-bodied reef fish as well as juveniles of large-bodied
predators (e.g. wrasse, damselfish, basselets, parrotfish, and grouper). Currently there are efforts
being made to control lionfish populations in the Roatan Marine Park (RMP) through a
spearfishing campaign involving the local dive community. As a result of this campaign, there
have been drastic declines in lionfish sightings in the park compared to when they first invaded
in 2009. Although they are venomous, lionfish are considered to be palatable with a taste similar
to grouper, prompting a slight economic incentive to catch them.

3. 3
In this study we examined abundance and distribution of lionfish in two areas to assess
the effects that spearfishing has had on their populations with respect to their behavior and
habitat choice. The behavior and areas where potential lionfish predators were found was also
taken into account in an attempt to understand how some native predatory reef species (e.g.
groupers, snappers, morays) have reacted to control efforts.
Methods
I and a diving partner surveyed for lionfish at two different sites using two different
survey methods (see below) designed to examine different aspects of lionfish distribution and
abundance. In addition, we made posters that would allow divers at five different dive shops to
record lionfish sightings during the period that we surveyed. I conducted interviews with staff at
dive shops in West End, Roatan to investigate the community perceptions of the effects that
spearfishing is having on lionfish and how spearfishing activity might be affecting reef
community dynamics.
Sites:
The two study areas were located on the northern side on the western end of the island of
Roatan, Honduras (Figure 1). The first area (sites 1-13) was located within the Roatan Marine
Park (RMP) spanning from the western tip of the island to the dive site Wrasse Hole (13). We
designated this area as well-speared for lionfish as culling efforts by both the RMP and local dive
shops have focused on managing lionfish populations here since 2010. The second area was
located northeast of the RMP (sites 14-19) where we had no specific information on past culling
efforts.

4. 4
Transect Survey Method:
We conducted six belt transect surveys (10 X 25 m) in each of the two areas. Survey sites
were between 5-18m deep (15-60 feet), were relatively flat, and had an abundance of aggregate
coral-cover. These areas were chosen because they were thought to offer the best lionfish habitat
(Biggs & Olden 2011). We conducted three surveys per site on two different dives. All transects
were conducted by two divers.
The transect surveys were
performed using a combination of the
fish survey from the Atlantic and Gulf
Rapid Reef Assessment (AGRRA) and
the Zig-Zag method (Figure 2) (AGRRA,
2010). We recorded our data focusing on lionfish and their potential predators on data sheets
based on those of the AGRRA surveys. Each survey took us roughly 20 minutes to complete. For
all of the lionfish sighted we recorded the depth, size, behavior, and characteristics of the area
(eg. out in the open, beneath an overhang, in a cave) where it was found. We also noted potential
predator behavior and the area where they were found. We surveyed areas in a biased eastward
Figure 1: Map showing the Central American region and the island of Roatan, Honduras with dive sites;for
site names see additional information on pg. 17
in red
Figure 2: Transect survey method used combining
AGRRA and Zig-Zag methodologies

5. 5
or westward fashion so that when we returned from our surface interval we wouldn’t survey the
same area twice. As a safety precaution we surveyed the deepest areas first.
Roving Diver Survey Method:
We conducted eight roving diver surveys at eight sites within the RMP randomly chosen
by the dive shop (Ocean Connections) with whom we partnered. Roving surveys were
conducted with groups of recreational divers. The advantages of roving surveys were that we
could visit deeper portions of the reef (>21.5m) than the transect surveys would allow. High
preference was given to sites that could be visited in the morning because they would most likely
be the deepest dives of the day. On roving dives we surveyed for lionfish under overhangs, in
swim-throughs, small caves and openings.
Dive Shop Data Collection Method and Interviews:
I administered a written survey at five dive shops in which divers voluntarily recorded the
following information from their dives: date; dive site; number of lionfish observed; maximum
dive depth; and whether or not any lionfish were caught. The reliability of the survey data was
enhanced by the fact that most entries were by dive instructors/masters rather than tourists.
I also conducted interviews with dive masters and instructors at different dive shops in
West End. I asked each dive master or instructor a series of similar questions in an effort to
substantiate or not trends observed in lionfish/potential predator behavior. We also asked each
interviewee their opinion on how spearfishing had affected lionfish populations in recent years.
Statistical Analysis:
To analyze the numbers of lionfish spotted outside and inside the RMP on transect
surveys, I used a Mann-Whitney non-parametric test to compare abundances per site. I analyzed

6. 6
the behaviors of both lionfish and potential predators using contingency tables to determine any
differences between populations observed in the RMP and outside of it. Lionfish were classified
as having either indifferent or vigilant/avoidant behavior whereas their potential predators were
classified as being either indifferent
or interested in our presence. I
performed a similar contingency test
to compare the number of
individuals found in different
microhabitats in both areas (Table
1).
In my analyses of potential
predators we used anecdotal information from local scientists, dive shop owners, dive masters, as
well as suggestions made by Mumby et al. (2011) to assign fish a potential predatory ranking.
The three families/subfamilies that represented predators were Epinephelinae (groupers),
Lutjanidae (snappers), and Muraenidae (moray Eels). We also classified sites into two different
groups depending on whether or not the ratio of lionfish to potential predators was greater than
or less than one to determine if a predator/prey ratio greater than one significantly affected
lionfish abundance. We performed a nonparametric contingency analysis for these groups to test
for differences between sites. We used a Student’s t-test to determine if there were any
differences in size in relation to depth using compiled data from the transect, roving, and dive
shop sightings. A student’s t-test was used to determine any differences between numbers of
lionfish spotted in shallow dives (<21.5m max depth) and deep dives (>21.5m).
Microhabitat found Criteria
C Found beneath an
overhang or in a coral
nook where it was
protected from 3 or more
sides
N Near the coral not more
than 0.5m away from the
surface
O Out in the open >0.5m
away from the coral
surface swimming about
Table 1: Classification of Microhabitat where lionfish and their
potential predators were found

7. 7
Results:
All three methods resulted in 293 lionfish sightings over a period of 4 weeks. About 16%
of the sightings were made during transect and roving surveys; the remaining were reported in
the dive shop surveys. There was no significant difference in abundance per site in transect
surveys (U(5,5) = 15.5, P > 0.10). According to the Roatan Marine Park and the dive
masters/instructors that I interviewed, areas within the park were determined to have one of the
highest levels of spearfishing effort compared to anywhere else on the island.
Fish Behavior
Preliminary data analysis suggested there were no differences in lionfish behavior
between sites in the RMP to those outside of it (χ2 = 1.16, df = 1, p = 0.2815). Lionfish exhibited
two main types of behaviors: indifferent behavior in which individuals would ignore our
presence or vigilant behavior in which they would actively avoid us. Further analysis grouping
the roving and transect sites into 2 different categories based on relative abundance of lionfish
and potential predators showed that there was no significant difference in lionfish behavior
between sites (χ2 = 1.915, df = 1, P =
0.1664). There was a trend of finding
more avoidant lionfish than expected
at sites that had equal numbers or
more potential predators than lionfish
(X2 = 3.19, df = 1, p = 0.0740)
(Figure 2). Our preliminary analysis
of potential predator behavior
showed no differences between areas (χ2 = 0,
0
5
10
15
20
25
LF>PLP LF≤PLP
NumberofLionfish
Indifferent
Avoidant
Figure 2: Lionfish behavior between sites with
more or less PLPs

8. 8
df = 1, P=1). The potential predators
exhibited two main types of behaviors:
indifferent or interested in our presence. The
fish in both areas that were most interested in
us belonged to the Epinephelinae (Groupers)
and Lutjanidae (Snappers) families and
subfamilies. The two species in both areas that appeared the most interested were the Mutton
Snapper (Lutjanus analis) and the Nassau Grouper (Epinephelus striatus). These two species
made up about half of the potential predators that appeared interested in our presence inside and
out of the park, respectively (Figure 3).
Fish Habitat
We observed lionfish in two main microhabitats on the reef: covered areas (overhangs,
small channel-like formations (swim-throughs), caves, & coral pockets) and non-covered areas.
We found that for both areas, in and out of the park, more than half of the lionfish were found in
covered areas where they were surrounded by three or more sides of coral. There was no
difference between the two main areas in
where the lionfish were found (χ2 = 0.56, df =
1, P = 0.454). Potential predators were mostly
found swimming out in the open with the
exception of some groupers and Moray eels
which commonly inhabit reef pockets and
crevices (J.S. Nelson, 1994). Although nearly
three-quarters of the predators we observed were
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
Frequency in C Frequency in
N
Frequency in
O
Outside RMP
Inside RMP
Figure 4: Frequency of PLPs found in the 3
different microhabitats; C = Covered, N =
Near, O = Open (Refer to Table 1 for complete
classifications)
Black Grouper
11%
Dog Snapper
5%
Mutton
Snapper
26%
Nassau Grouper
21%
Tiger Grouper
26%
Yellow-tail
Snapper
11%
Figure 3: Proportion of PLPs found inside and out of RMP

9. 9
found swimming in the open, we found that there was a significantly higher number of predators
found in covered microhabitats within the RMP compared to the area outside (χ2 = 4.6, df = 1, P
= 0.032) (Figure 4).
Depth Analysis
I found that the average number of lionfish spotted per dive was significantly higher on
dives whose maximum depth exceeded 21.5m (t = 2.19, df = 95, p = 0.0311). For dives that had
a maximum depth <21.5m an average of 2 lionfish were spotted whereas on dives whose max
depths were >21.5m an average of 4 lionfish were observed. All 10 dive masters who were
interviewed agreed that they were encountering more lionfish at depth than on shallower areas of
the reef. Their assertion that bigger lionfish were being found deeper, however, was not
supported by our analysis (χ2 =
0.23, df = 1, P = 0.6315). After
comparing depth data between
sightings during transect and
roving diver surveys there was a
trend of finding them in higher
abundances on deeper portions of
the reef (>10m) (Figure 5).
Discussion:
Spearfishing around Roatan has become a common activity throughout the dive
community. The Roatan Marine Park (RMP) has been subject to the most intense culling efforts
due to the large diver population and widely publicized effort to control lionfish. I found that
there is evidence of regular hunting outside the RMP and that culling efforts may be excluding
Figure 5: Lionfish depth sightings

10. 10
them from the upper 10m of the reef. It is also having an effect on the habitat use of certain
upper trophic level species that I have distinguished as Potential Lionfish Predators (PLPs)
which may serve as a natural form of invasive species control.
Transect surveys at twelve different sites between the well-fished area (RMP) and areas
of unknown culling effort revealed similar numbers in both locales, indicating that spearfishing
was occurring inside and out of the RMP. Anecdotal data from two dive masters associated with
regular hunting of lionfish support our findings, verifying that the area of unknown culling effort
is regularly visited by spearfishing divers. Similar abundances at similar depths between both
areas suggests that culling efforts may be excluding lionfish from some of the most productive
areas of the reef (i.e. upper 10 m of the reef).
Four different categories of behavior was displayed by lionfish and PLPs evenly between
both areas. Such observed differences between behaviors among both groups of fish may be
caused by spearfishing (Côté et al., 2014). Since culling efforts began in 2009, divers in the
community have noticed shifts in the behavior of certain individuals. We found that the majority
of lionfish were indifferent to our presence often allowing us to swim near them to take
measurements. On multiple occasions we took note of individuals that were alerted by our
presence and avoided us by swimming away or taking refuge in the reef were they were not
visible. When asked about this kind of behavior, many of the dive masters claimed that avoidant
individuals had come into previous contact with spearfishing divers. They claimed that if the
lionfish had been previously targeted, it would avoid any sort of diver thereafter. Côté et al.
(2014) suggested that culling efforts are having an effect on lionfish behavior in the Bahamas
causing depressed activity levels during the day on patches of reef with known hunting efforts.

11. 11
Culling efforts have most likely had a similar effect on lionfish populations around Roatan
producing avoidant individuals in and around areas with high amounts of spearing.
Spear fishing divers will occasionally feed part of their catch to reef predators found at or
nearby the kill site. Multiple dive maters interviewed claimed that PLPs would only accept
lionfish if they were wounded or dead. This feeding may prevent PLPs from including lionfish in
their search images and may only teach them that lionfish are only edible if provided by a hunter
(Orams, 2002). Mumby et al. (2011) suggested that grouper in the Exuma Keys, Bahamas are
acting as a natural biocontrol of lionfish populations and may be keeping their numbers low in
shallow depths. Mumby et al. (2011) did not address whether or not these grouper were catching
the lionfish on their own or if they were being fed by spearfishing divers.
According to the interviewed dive masters and dive instructors, groupers, snappers, and
morays have been the most common reef predators to be fed lionfish by spearfishing divers. All
of the interviewees concluded that the two most prominent species of predators to accept lionfish
were the Nassau Grouper (Epinephelus striatus) and Mutton Snapper (Lutjanus analis).
Maljković et al. (2008) and Mumby et al. (2011) found that these local species would also accept
lionfish in other invaded regions of the Caribbean. While surveying we noticed that the majority
of potential predators were indifferent to our presence. On more than one occasion we
encountered potential predators that approached us unprovoked. We speculate that this kind of
behavior may be caused by regular feeding by visiting divers.
The lack of any significant difference in the behaviors of the lionfish or potential
predators between both areas also suggests that lionfish culling efforts are occurring outside the
park. Changes in behavior in both the native PLPs and lionfish at different areas of the reef has
led us to believe that spear fishing is impacting population/community dynamics and fish

12. 12
response to human presence. These findings do not conclude that the PLPs are viable
biocontrols, but it does suggest that they may be used as tools in conservation and control efforts.
After comparing the two areas I hypothesize that spearing of lionfish on the shallower
portions of the reef has changed the distribution of depths at which lionfish are found. I found
that most of the lionfish spotted were at depths exceeding 10 m. I also found that on average
there were twice as many lionfish sighted on dives whose maximum depth exceeded 21.5m. All
of the dive masters and instructors that I interviewed agreed that the lionfish populations were
being found in greater abundances at depth since culling efforts began. As most spearing is
reported to occur in the upper 30 m of the reef the lionfish populations at this depth have been
impacted the most. Most hunting doesn’t extend beyond these depths on a regular basis as most
hunters are not certified to travel this deep. Current research is being conducted in an attempt to
maintain culling efforts that range from developing fish traps with specially designed vents to
establishing fisheries with high lionfish exploitation rates. Such methods would eliminate divers
visiting deeper, more dangerous portions of the reef (Barbour et al., 2011; Olsen & Hill, 2012).
Finding higher abundances of lionfish at deeper portions of the reef may present a risk to
reef health. If populations of lionfish continue to grow at mesophotic reef depths (>30 m) they
may indirectly affect overall reef health by consuming small-bodied reef fish and crustaceans
that maintain reef health (Lesser and Slattery, 2011). Lesser et al. (2009) also suggested that
mesophotic reefs provide an important nursery-like habitat for some corals, sponges, and fish
that inhabit the shallower areas as adults. Lesser and Slattery (2011) speculated that if lionfish
populations remained at mesophotic areas then they could affect the reef at all depths by
removing important algal grazers allowing for coral-algal phase shifts to take place.

13. 13
Another factor that may have attributed to lionfish progressing to deeper areas is
interspecific competition with other predators for shelters. We found that there was a significant
difference between the locations that we were finding predators within the park relative to
outside of it. Within the park PLPs were mostly found in caves whereas outside the protected
area they were most commonly found out in the open. Raymond et al. (2014) suggested that
when Nassau grouper and lionfish share a habitat, the lionfish will dominate the available shelter.
The prevalence of more predators inhabiting caves in the RMP could signify that larger
populations of lionfish remain outside the park and that they are outcompeting predators for
shelter. The prevalence of PLPs in particular microhabitats may also be used as a marker of how
intense and focused spearing efforts may reduce the invasive competition enough to reverse
some of the negative effects of the invasion by providing habitat that native species need to
survive and reproduce (Zavaleta et al. 2001).
Thus far, research on the lionfish invasion has resulted in increased knowledge about the
biology, ecology, and potential impacts of this species. Further data is needed to help understand
and predict what the long term impacts lionfish will have on the Caribbean and if ecosystems can
remain healthy, if invaded. The future of the reef around Roatan is uncertain. Overfishing,
terrestrial development, and lionfish weigh on the conservation of the reef. The location at which
the greatest abundances of lionfish are being found around the island is still unknown. Such an
analysis should be done in as many areas of the island as possible to better understand what
factors influence lionfish distribution. Such information would be invaluable for conservationists
in Roatan and around the Caribbean. Culling efforts around Roatan have had an impact on the
dynamics of different areas of reef on the north-western side of the island. When lionfish first
invaded in 2009 it was common to sight 10-12 individuals on a dive, these days more effort is

14. 14
required to locate them. Individuals are more wary of diver presence and have been cleared in
high numbers from the marine park most likely allowing for their potential predators to find
shelter in the reef. The consensus around the island is that spearfishing is working and our data
suggest something similar. Currently, spearing seems to be the best and most efficient way to
control the lionfish populations, however, more comprehensive, long-term control strategies are
needed to ensure the health of the reef and its native communities.

15. 15
References:
Aguilar-Perera A, Tuz-Sulub A (2010) Non-native, Invasive Red Lionfish (Pterois Volitans
[Linnaeus, 1758]: Scorpaenidae), Is First Recorded in the Southern Gulf of Mexico, off the
Northern Yucatan Peninsula, Mexico. Aquatic Invasions 5 Supplement 1: S9-S12
Ahrenholz DW, Morris Jr JA (2010) Larval duration of the lionfish, Pterois volitans along the
Bahamian Archipelago. Environmental Biology of Fishes 8: 305-309
Albins MA, Hixon MA (2011) Worst Case Scenario: Potential Long-term Effects of Invasive
Predatory Lionfish (Pterois Volitans) on Atlantic and Caribbean Coral-reef Communities.
Environmental Biology of Fishes 96.10-1: 1151-1157
Atlantic and Gulf Rapid Reef Assessment (2010) Method for Assessing Coral Condition.
AGRRA. http://www.agrra.org/
Barbour AB, Allen MS, Frazer TK, Sherman KD (2011) Evaluating the Potential Efficacy of
Invasive Lionfish (Pterois volitans) Removals. PLoS ONE 6: n. pag
Biggs C, Olden J (2011) Multi-scale Habitat Occupancy of Invasive Lionfish (Pterois Volitans)
in Coral Reef Environments of Roatan, Honduras. Aquatic Invasions 6.3: 347-353
Côté IM, Darling ES, Malpica-Cruz L, Smith NS, Green SJ, Curtis-Quick J, Layman C
(2014) What Doesn’t Kill You Makes You Wary? Effect of Repeated Culling on the Behaviour
of an Invasive Predator. PloS ONE 9(4): n. pag
Kizer KW, McKinney HE, Auerbach PS (1985) Scorpaenidae envenomations: A five-year
poison center experience. Journal of the American Medical Association 253:807-810
LesserMP, Slattery M (2011) Phase Shift to Algal Dominated Communities at Mesophotic
Depths Associated with Lionfish (Pterois Volitans) Invasion on a Bahamian Coral Reef.
Biological Invasions 13: 1855-1868
LesserMP, Slattery M, Leichter JJ (2009) Ecology of Mesophotic Coral Reefs. Journal of
Experimental Marine Biology and Ecology 375.1-2: 1-8
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Volitans (Pisces: Scorpaenidae), by Native Groupers in the Bahamas. Coral Reefs 27.3: 501
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thesis. North Carolina State University, Raleigh
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Morris Jr JA, Akins JL (2009) Feeding Ecology of Invasive Lionfish (Pterois Volitans) in the
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17. 17
Additional Information:
(Figure 1 pg. 7)
Sites inside Roatan Marine Park
1) West End Wall
2) Sea Quest Deep
3) Turtle Crossing
4) Octopus Acre
5) The Bight
6) Blue Channel
7) Dixie’s
8) Dive Master’s Choice
9) Hole in the Wall
10) Gibson Bight
11) Overheat Reef
12) Front Porch
13) Wrasse Hole
Sites outside Roatan Marine Park
14) Mila’s Spot
15) Jenny’s Dream
16) Mucky Hole
17) Man O’ War Cay
18) Turtling Bay
19) Palmetto Bay