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LIONFISH PAPERS AND PRESENTATIONS FROM THE GULF AND CARIBBEAN FISHERIES INSTITUTE 2008‐2010 Picture from Acero et al. INTRODUCTION
The documents that comprise this compendium
represent papers and presentations from lionfish special
sessions held at the annual meeting of the Gulf and
Caribbean Fisheries Institute. Starting in 2008, the GCFI,
together with partners from NOAA, Florida Sea Grant,
and REEF, have conducted sessions comprised of oral
presentations on topics ranging from lionfish control
strategies in the Wider Caribbean to new research results
on lionfish biology and ecology. Lionfish ecological
impacts was a common and urgently expressed theme
throughout the sessions as presenters described extreme
changes to local biodiversity and native reef fish biomass
owing to lionfish consumption. New insights were
provided on lionfish age and growth, habitat correlations,
bioenergetics, genetics, and local removal programs.
GCFI expresses grateful appreciation for for financial
support to Florida Sea Grant.
PAPERS
Table of Contents
Papers
2008
Smith and Sullivan Sealey – The Lionfish Invasion in the Bahamas: What do We Know and What
to do About It?
2009
Claydon et al – The Red Lionfish Invasion of South Caicos, Turks & Caicos Islands
Green and Côté – Abundance of Invasive Lionfish (Pterois volitans) on Bahamian Coral Reefs
Morris et al – Biology and Ecology of the Invasive Lionfishes, Pterois miles and Pterois volitans
Sullivan Sealey et al. – The Invasion of Indo-Pacific Lionfish in the Bahamas: Challenges for a
National Response Plan
2010
Green and Côté – Consumption Potential of Invasive Lionfish (Pterois volitans) On Caribbean
Coral Reefs
Presentations
2010
Acero et al – Reconstructing the Western Atlantic Lionfish Invasion
Albins – Effects of invasive Pacific red lionfish on Bahamian coral-reef fish communities:
preliminary results from a large-scale, long-term experiment
Bernal et al – The Impacts of the Indo-Pacific Lionfish (P. volitans and P. miles) on Fish
Assemblages in Near Shore Benthic Reefal Habitats of the Central and Southern Bahamas
Claydon et al – Invasive red lionfish in shallow habitats of the Turks & Caicos Islands
Donaldson et al – Why Are Lionfishes (Pterois, Scorpaenidae) So Rare In Their Native Ranges?
Johnson et al. – Red Lionfish control strategies in the Caribbean UK Overseas Territories
(Cayman Islands, British Virgin Islands and Turks and Caicos Island)
Meléndez – Implementation of the Management Plan for the Control of the Lionfish in Puerto
Rico
Molina Ureña – Lionfish in Costa Rica: threats, actions, and opportunities
Santos-Martínez – Trophic and Reproductive Aspects of the Lion Fish Pterois volitans, in San
Sandrés Island, Biosphere Reserve- Seaflower, Colombian Caribbean
The Lionfish Invasion in the Bahamas: What do We Know and What to do About It?
NICOLA S. SMITH1,2 and KATHLEEN SULLIVAN SEALEY1
Marine and Environmental Studies Institute, College of The Bahamas, Oakes Field Campus,
P.O. Box N4912 Nassau, Bahamas
2
Graduate Department of Zoology, University of British Columbia, Biological Sciences Building,
6270 University Boulevard, Vancouver, BC, Canada V6T 1Z4
1
ABSTRACT
Biological invasions include both human and non-human mediated forms of dispersal in which an exotic or non-native species
successfully arrives, survives and reproduces in a novel locality and then proliferates and spreads throughout a region (Carlton,
1989). The recent invasion of the Indo-Pacific lionfish (Pterois volitans) throughout the western Atlantic Ocean, including The
Bahamas, is generally considered to be the result of several species introductions associated with both the intentional and unintentional release of specimens from private aquariums. Small Island Developing States (SIDS) like The Bahamas are particularly
vulnerable to bioinvasions due to our: 1) import-driven economy; 2) heavy reliance on tourism; and, 3) biological fragility inherent
in island ecosystems. A review of the pattern of invasion by habitat, island group and size distribution is presented from recent
surveys throughout the archipelago.
KEY WORDS: Lionfish, reef fish, Bahamas, invasive species
La Invasión del Pes Escorpion en las Bahamas
Las invasiones biológicas comprenden tanto las formas de dispersión naturales como las generadas por el Hombre, donde una
especie exótica o no nativa arriba, sobrevive y reproduce en una nueva localidad, y prolifera y se propaga por toda la región
(Carlton, 1989). La reciente invasión del pez escorpión del Indopacífico (Pterois volitans) a través del Atlántico occidental,
incluyendo Las Bahamas, se considera el resultado de la liberación intencional y no intencional de ejemplares por acuarios privados.
Los Estados Pequeños Insulares en Desarrollo (SIDS) como Las Bahamas son particularmente vulnerables a las invasiones
biológicas debido a 1) una economía sustentada por la importación y la exportación; 2) a la gran dependencia del turismo; y 3) la
fragilidad biológica inherente a los ecosistemas insulares. Se presenta una revisión del patrón de invasión por hábitat, grupo de islas
y amplitud de la distribución, a partir de los resultados de prospecciones recientes a lo largo del archipiélago
PALABRAS CLAVES: Pez escorpion, arrecifes, Bahamas, invasión
INTRODUCTION
Biological invasions include both human and nonhuman mediated forms of dispersal in which an exotic or
non-native species successfully arrives, survives, and
reproduces in a novel locality and then rapidly spreads
throughout a region. On the one hand, species that disperse
without the aid of humans into an area where they were not
previously found are referred to as range expansions. On
the other hand, species that have been released outside of
their native range and have spread due to human activity
are referred to as species introductions (Carlton 1989). The
recent invasion of the Indo-Pacific lionfish (Pterois
volitans*) throughout the western Atlantic Ocean, including
The Bahamas, is generally considered to be the result of
several species introductions associated with both the
intentional and unintentional release of specimens from
private aquariums followed by the natural dispersal process
(Hare and Whitfield 2003, Ruiz-Carus et. al. 2006).
Biological invasions in the form of species introductions, however, are not new. The introduction of nonnative species has occurred since the dawn of early human
migrations. Historically, flora and fauna were intentionally
introduced to a new region usually to satisfy food demands
or social needs while other non-native species would have
been accidentally brought to an area in the form of
hitchhikers. What distinguishes invasions today from those
Proceedings of the 60th Gulf and Caribbean Fisheries Institute
experienced in the past is that the current rate and magnitude of human-caused invasions is unprecedented (Lowe
et. al. 2000).
Today’s worldwide trend in successful species
introductions can be partly explained by the exponential
increase in global trade, transport, tourism, and travel,
which have all served to transfer species to places that
would have otherwise been virtually impossible to access
due to natural physical barriers (BEST 2003, Lowe et. al.
2000). Furthermore, invasion theory predicts that increased disturbance to an environment should result in
increased invader success (Altman and Whitlatch 2007,
Lozon and MacIsaac 1997). Interpreted from this perspective, Carlton and Ruiz (2005) suggest that current worldwide increases in urban and other disturbed habitats may
have created an excess of modified ecosystems that are
more susceptible to invasion.
Small Island Developing States (SIDS) like The
Bahamas are particularly vulnerable to bioinvasions due to
our:
i) Import-driven economy;
ii) Heavy reliance on tourism; and,
iii) Biological fragility inherent in island ecosystems
(BEST, 2003).
November 5 - 9, 2007 Punta Cana, Dominican Republic
Page 420
60th Gulf and Caribbean Fisheries Institute
Explanations for The Bahamas’ first two vulnerabilities to
invasion are self-evident: a high level in the global traffic
of people and goods leads to increased opportunities for
non-native species that were once far-removed from The
Bahamas to come in contact with our shores as hitchhikers.
Evidently a small proportion of hitchhikers prove to be
successful invaders.
The third vulnerability of The Bahamas to invasion is
a bit more complicated but is probably best explained by
the “empty niche hypothesis”. Ecological models predict
that the likelihood of establishment of an exotic species is
increased when the functional differences between the nonnative and the resident/native species are great (Tilman
2004, Von Holle and Simberloff 2004). Small islands like
those of The Bahamas tend to be missing entire functional
groups of organisms – an example being the absence of
native mammalian top predators. Introduced species that
are able to occupy these missing functional groups may
therefore be more likely to become successful invaders due
to their ability both to use unexploited resources and to
compete for other resources with inexperienced natives
(Alpert 2006).
All invaders impact the environment in which they
invade because every organism must utilize resources such
as space and food to survive. However, not all invaders
have readily discernible effects on the invaded community
or ecosystem. In cases in which impacts are substantial,
and thus detectable, biological invasions have resulted in:
declines, extirpations, and extinctions of native species
(Goldschmidt et. al. 1993, Witte et. al. 2000, Lowe et. al.
2000), alterations of natural disturbance regimes
(D’Antonio and Vitousek 1992), habitat structure (Daehler
and Strong 1996), and nutrient cycling (Vitousek et. al.
1987) which have all in turn, changed the ecology of
natural systems; changes in food web structure (Vander
Zanden et. al. 1999), morphological and behavioural
changes in native species (Vermeij 1982, Trussell and
Smith 2000), and hybridization of native species with the
invader (Rhymer and Simberloff 1996). Indeed, biological
invasions are considered to be one of the leading threats to
biodiversity worldwide (Lowe et. al. 2000).
Nevertheless, it is important to note that there have
been some, though few, documented beneficial effects of
invasions. For example, Crooks et. al. (1998, 1999) found
that the invasive Asian mussel, Musculista senhousia,
created new habitat via producing mats of byssal threads in
the predominantly unstructured mudflats of Mission Bay,
San Diego. This novel habitat subsequently allowed for
the development of a unique community assemblage with a
higher diversity and abundance of taxa than the neighboring mudflats. Similarly, King and colleagues (2006) point
out that the invasive round goby, Neogobius melanostomus, now constitutes more than 92% of prey consumed by
the resident Lake Erie Water Snake, Nerodia sipedon
insularum, which is threatened in the US and endangered
in Canada. This shift in diet by the water snake following
the invasion of the round goby has resulted in more rapid
growth and attainment of a larger body size in the water
snake - which the scientists assert may in turn, reduce
predation, speed reproductive maturity, increase offspring
production and ultimately, promote population growth in
this threatened/endangered species.
LIONFISH IN THEIR NATIVE RANGE
Lionfish (Pterois volitans and P. miles) are tropical
reef fish native to the Indian and South Pacific Oceans,
including the Red Sea, where they inhabit coral reefs,
rocky outcrops and sandy substrates at depths ranging from
the surface (< 1 m) to about 50 m (Schultz 1986). Both P.
volitans and P. miles are variable in color but tend to be
either red-, maroon-, or black-and-white striped (Figure 1).
They are a source of food in their native range and are
highly sought after globally as a high priced aquarium fish
(FishBase 2007).
Figure 1. An adult lionfish on a near shore patch reef in its
introduced range of The Bahamas. Photo credit: COBMESI, 2007.
The maximum size record for adult lionfish in their
native range varies according to the source with a conservative estimate of about 380 mm TL (FishBase, 2007). P.
volitans become sexually mature between 140 - 160 g body
weight and 180 - 190 mm TL (Fishelson 1997). Based on
spawning information and the collection of larvae from the
water column, it is likely that lionfish have a pelagic larval
stage (Hare and Whitfield 2003).
Lionfish are usually solitary as adults and will defend
their home range against conspecifics. Nevertheless, they
tend to congregate in small groups during mating and as
juveniles (Fishelson 1975). Juveniles have also been
observed to gather together in groups of up to 40 individuals (Fishelson 1997).
Relatively stationary, top-level predators, lionfish feed
on a wide variety of smaller fishes and crustaceans
Smith, N.S. and K. Sullivan-Sealy GCFI:60 (2008)
(Fishelson 1997). There are few known, if any, natural
predators of lionfish, most likely due to the venomous
nature of the species (Allen and Eschmeyer 1973).
However, the literature reports an isolated case of a single
pacific cornetfish, Fistularia commersonii, (94 cm SL)
with a P. miles in its stomach (10 cm SL) (Bernadsky and
Goulet 1991). Furthermore, it has been speculated that
some sharks may consume lionfish with no apparent illeffect (Moyer and Zaiser 1981).
Schultz (1986) concluded that Pterois volitans and P.
miles are allopatric, sibling species. Kochzius et. al.
(2003) showed that there are genetic differences between
P. volitans and P. miles, but they were inconclusive as to
whether they are two separate species or two populations
of a single species. This paper recognizes P. volitans and
P. miles as two separate species as determined by Schultz
(1986) – although keeping with convention, both are herein
commonly referred to as lionfish - and acknowledges the
possibility that a cryptic invasion is occurring in The
Bahamas similar to the situation in the US (Hamner and
Freshwater 2007) in which both P. volitans and P. miles
are found along the southeast continental shelf.
LIONFISH ENVENOMATION
Members of the family Scorpaenidae, lionfish possess
venomous dorsal, anal, and pelvic spines and have been
known to sting humans when threatened or harassed (RuizCarus et. al. 2006, Vetrano et. al. 2002). Envenomation
may also occur due to reckless handling of recently dead
specimens (Pulce et. al. 1991). Lionfish venom contains
both acetylcholine and a toxin affecting neuromuscular
transmission, but the major component is an antigenic, heat
labile protein (Vetrano et. al. 2002). No fatalities have
resulted due to lionfish envenomations, and the majority of
stings occurred on the hands of victims who attempted to
clean the aquarium of fish kept as pets (Patel and Wells
1993).
The predominant symptom of lionfish envenomation is
severe pain at the wound site, which is usually responsive
to hot water immersion therapy. Rare but more serious
symptoms include: chills, headache, nausea, vomiting,
abdominal pain or cramping, delirium, seizures, limb
paralysis, hyper- or hypotension, respiratory distress,
congestive heart failure, and pulmonary edema (Vetrano et.
al. 2002). Victims may develop a hypersensitivity to
lionfish venom and experience anaphylactic reactions upon
subsequent envenomation (Auerbach 1991, Patel and Wells
1993).
THE INTRODUCTION OF LIONFISH TO THE
WESTERN ATLANTIC OCEAN AND
THE BAHAMAS
The first documented release of lionfish in US waters
occurred in 1992 in Biscayne Bay, Florida when six
lionfish escaped from a private aquarium following its
destruction by Hurricane Andrew (Courtenay 1995). Since
Page 421
then, adult lionfish have been observed along the southeast
United States coast from Miami, Florida to as far north as
Cape Hatteras, North Carolina - in addition to Bermuda. In
comparison, juveniles have been sighted off North
Carolina, Long Island, New York and Bermuda (Whitfield
et. al. 2002). However, juveniles remaining in US waters
farther north than Cape Hatteras in the fall are predicted to
perish due to an inability to survive winter bottom temperatures there (Kimball et. al. 2004).
In contrast, the first documented report of lionfish in
The Bahamas did not occur until 2004 (Department of
Marine Resources, Pers. comm.). Nevertheless, by late
2006, lionfish had already been reported on a variety of
habitat types throughout much of The Bahamian archipelago by local scientists, environmentalists, fishermen,
recreational divers, and beach goers alike. Many of the
reports were anecdotal and consisted of brief emails or
phone calls directed to various Government Ministries or to
The Marine and Environmental Studies Institute at The
College of The Bahamas.
At the start of 2007, the Reef Environmental Education Foundation (REEF) – a Florida based environmental
not-for-profit organization – teamed up with local dive
operators and the College of The Bahamas to conduct
lionfish surveys throughout the archipelago at popular dive
sites. Furthermore, in August 2007, the College of The
Bahamas Marine and Environmental Studies Institute
(COB-MESI) in collaboration with the Department of
Marine Resources established an on-line lionfish sightings
questionnaire in order to consolidate information on
lionfish occurrences throughout the country and subsequently follow-up by verifying reports.
Preliminary surveys of lionfish around New Providence were conducted in summer of 2007 by The COBMESI. Their findings revealed, among other things, that a
substantial number of lionfish are being found in highly
disturbed, near shore, shallow waters of The Bahamas
(between 1 – 4 m) (Sullivan Sealey and Smith In prep.) as
opposed to the deeper, offshore waters (the majority of
lionfish observed between 35 and 45 m) of the lionfish
introduced range along the southeast coast of the US
(Kimball et. al. 2004). This suggests that the pattern of
invasion of lionfish in The Bahamas may more closely
resemble lionfish occurrences and distributions in their
native range of the Indo-Pacific than their introduced range
in neighboring US waters. Evidently, further research is
needed to fully characterize the nature of the invasion of
lionfish in The Bahamas.
Nevertheless, this initial
difference highlights the need for The Bahamas to invest
substantial resources into closely evaluating the lionfish
invasion in its own waters, as reliance on the findings and
subsequent management policies developed to address the
invasion in the US may not necessarily be applicable here.
Page 422
60th Gulf and Caribbean Fisheries Institute
A PLANNED RESPONSE FOR THE BAHAMAS
The scale and scope of the lionfish invasion in The
Bahamas requires innovative approaches and partnerships
to: protect public health and interests, assess the potential
impacts of lionfish, and effectively manage the invasion.
Biological invasion management is a multi-year endeavor
for both marine and terrestrial species; and a draft of a
National Invasive Species Policy for The Bahamas has
already been developed, which among other things, calls
for the country:
i) To prepare a strategic management plan for
individual species of high priority;
ii) To facilitate research on the occurrence, distribution and impacts of invasive alien species, and
iii) To monitor invasive species populations in The
Bahamas (BEST, 2003).
The College of The Bahamas Marine and Environmental Studies Institute (COB-MESI) in collaboration with
the Department of Marine Resources is launching a multiyear project to develop a National Lionfish Response Plan
(NLRP) that entails a partnership between both local and
regional government and non-governmental agencies. The
plan focuses on:
i) Ecological research,
ii) Invasion management and policy development,
and
iii) Educational initiatives to understand the implications of the establishment of the Indo-Pacific
lionfish on fisheries resources and the ecology of
coastal systems in The Bahamas. The project will
ultimately build a body of stakeholders that can
contribute to the long-term strategic management
of lionfish in our waters.
Preliminary research will address questions surrounding which types of near-shore habitats are more susceptible
to invasion and lionfish diet niches. A few permanent
monitoring sites will also be established around New
Providence to examine lionfish movement, habitat
utilization and recruitment. Longer-term efforts include
the creation of a National Lionfish Specimen Library for
future investigations into genetics, ageing, growth and
reproduction of the species in The Bahamas (Figure 2).
Initial invasion management and policy development
includes the creation of a national online lionfish information network that serves both to compile sightings over the
entire archipelago and to coordinate efforts with the
regional lionfish invasion work being done by REEF and
the U.S. National Marine Fisheries Service Laboratory
based in Beaufort, North Carolina (NMFS-NOAA).
Specifically, the online information network will include: a
national lionfish reporting system that is linked to a spatial
dataset of marine habitats; a specimen cataloging system
for the tracking of lionfish collected in the country and
housed either within The Bahamas at the National Lionfish
Specimen Library or abroad; and, a contact and project
database related to on-going lionfish research permitted in
the country that is linked to two the preceding network
components.
Figure 2. COB-MESI researchers record various lionfish
morphological characteristics in addition to specific habitat
information before housing specimens within the country in
the National Lionfish Specimen Library. Photo credit: COBMESI, 2007.
Future educational efforts of the NLRP involve raising
awareness among beach goers, as well as the local fishing
and recreational diving communities, about lionfish
invasion management options and first-aid response to
envenomation.
The National Lionfish Response Plan (NLRP) is a
costly and ambitious long-term endeavor. The project
requires significant financial, technical, and logistical
support from multiple government and non-governmental
agencies at both the national and the regional level. The
Indo-Pacific lionfish is now found widely throughout The
Bahamas, and the effects of their invasion are expected to
become more apparent as their numbers continue to
increase. As a concerned individual, you can best contribute to the National Lionfish Response Plan by: reporting
lionfish sightings on the national online questionnaire;
urging others to report sightings; and, donating money or
other much needed resources to the National Lionfish
Response Team.
ACKNOWLEDGEMENTS
We would like to thank the Disney Wildlife Conservation Fund,
Department of Marine Resources and the College of The Bahamas Marine
and Environmental Studies Institute for support and funding for this
project.
Smith, N.S. and K. Sullivan-Sealy GCFI:60 (2008)
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The Red Lionfish Invasion of South Caicos, Turks & Caicos Islands
JOHN ALEXANDER BRIGHTMAN CLAYDON, MARTA CATERINA CALOSSO, and SIRI ELIZABETH JACOB
The School for Field Studies – Center for Marine Resource Studies, 1 West Street,
South Caicos, Turks & Caicos Islands, British West Indies
ABSTRACT
The first observation of red lionfish (Pterois volitans/miles) in the waters around South Caicos, Turks & Caicos Islands was
made in December 2007. From this time until the end of July 2008, lionfish sightings were recorded by staff and students from The
School for Field Studies Center for Marine Resource Studies in South Caicos. Twenty-five individuals were observed. Although
effort was made to capture all specimens seen (with 23 individuals captured), sightings represent opportunistic observations made
during other activities. All except two were recorded in waters shallower than 2.5m, and specimens have been found in patch reef (n
= 15), seagrass (n = 6), mangrove (n = 2), and deep reef (25m; n = 1). Although individuals captured ranged in size from 4.1 to
27.7cm TL, all but two individuals were < 15 cm TL. This study documents the invasion of South Caicos by red lionfish, and
although the effects of this invasion are unknown, the exponential increase of sightings per month is worrying. Future monitoring
will include targeted searches for red lionfish.
KEY WORDS: Red lionfish, invasión, Turks & Caicos Islands
La Invasión de South Caicos, Turks & Caicos Islands, por los Peces León Rojo
La primera observación de pez león rojo (Pterois volitans/miles) en las aguas alrededor de South Caicos, Turks & Caicos
Islands fue hecha en Diciembre de 2007. Desde esa fecha hasta fines de Julio de 2008, avistamientos de peces león han sido
registrados por estudiantes y miembros de The School for Field Studies Center for Marine Resource Studies en South Caicos.
Veinticinco individuos han sido observados. Aunque se realizaron esfuerzos para capturar todos los especímenes observados (con 23
individuos capturados), los avistamientos representan observaciones oportunistas hechas durante otras actividades. Todos, excepto
dos, fueron registrados en aguas de profundidad menor a 2.5 m y especímenes han sido encontrados en manchas de arrecife (n = 15),
hierba marina (n = 6), manglares (n = 2) y arrecife profundo (25 m; n = 1). Todos los individuos capturados iban de una gama de
4.1 a 2.27 cm TL, excepto 2 que eran de <15cm TL. Este estudio documenta la invasión de South Caicos por los peces león rojo y,
aunque los efectos de esta invasión son desconocidos, el crecimiento exponencial de avistamientos mensuales es preocupante.
Futuros monitoreos incluirán búsquedas específicamente orientadas al pez león rojo.
PALABRAS CLAVES: Pez león rojo, invasión, Turks & Caicos Islands
Invasion de South Caicos, Îles Turks & Caicos, par le Poisson Scorpion
La première obseration d’un poisson scorpion (Pterois volitans/miles) dans les eaux autour de South Caicos a été faite en
décembre 2007. Depuis lors et jusqu’à la fin juillet 2008, le staff et les étudiants de The School For Field Studies Center for Marine
Resource Studies localisée à South Caicos ont pris note de toutes les observations de poissons scorpions. Vingt-cinq individus ont
été observés. Tandis qu’un effort a été fourni pour capturer tous les individus observés (avec 23 capturés), toutes les observations
représentent des observations opportunistes, réalisées pendant d’autres activités. Toutes, sauf deux, ont été enregistrées dans des
eaux peu profondes de moins de 2.5m et les spécimens ont été trouvés dans des récifs en patch (n = 15), herbiers (n = 6), mangroves
(n = 2), et récifs profonds (25 m ; n = 1). Bien que les individus capturés aient une taille de 4.1 à 27.7 cm TL, tous les individus
étaient < 15 cm TL sauf deux. Cette étude documente l’invasion de South Caicos par le poisson scorpion et, bien que les effets de
cette invasion ne soient pas connus, l’augmentation exponentielle des observations est inquiétante. Futures monitorings incluront
des recherches spécifiques pour les poissons scorpions.
MOTS CLES: Poisson scorpion, invasion, Îles Turks & Caicos
INTRODUCTION
The red lionfish, Pterois volitans/miles, invasion of the
western Atlantic is well documented (Whitfield et al.
2007). However, very little information has been collected
in the Turks & Caicos Islands (TCI). Whilst the first
sightings in the TCI were made off West Caicos and
Providenciales in 2005, in South Caicos no individuals had
been documented until December 2007 (Figure 1). This
study documents the invasion of red lionfish in the waters
around South Caicos following the initial sighting.
Proceedings of the 61st Gulf and Caribbean Fisheries Institute
MATERIALS & METHODS
Lionfish sightings around South Caicos were recorded
by staff and students from The School for Field Studies,
Center for Marine Resource Studies (SFS-CMRS). These
sightings represent opportunistic observations made during
snorkeling and SCUBA diving activities. Depth and
habitat were recorded, and specimens were captured to
prevent multiple sightings of the same individual. Total
lengths were measured and maturity assessed through
gonad analysis.
November 10 - 14, 2008 Gosier, Guadeloupe, French West Indies
Claydon, J.A.B., et al.
GCFI:61 (2009)
Page 401
research will include a more systematic and representative
monitoring protocol. In addition, potential threats of the
lionfish invasion will be investigated with the aim of
identifying appropriate strategies of management and
mitigation.
Figure 1. Location of South Caicos within the Turks &
Caicos Islands.
RESULTS
A total of 25 lionfish were observed, of which 23 were
captured and measured. The number of sightings per
month increased exponentially during the study period
(Figure 2). Most lionfish (n = 24) were recorded at less
than 2.5m depth. The majority were found in reef habitats
(15 on patch reefs, 1 on fringing reef, and 1 on the edge of
the drop-off at 25 m). Six individuals were seen in
seagrass habitats of which five were sheltering in blow-out
ledges (Figure 3). A further two lionfish were found in
mangroves (Figure 2). Specimens ranged in size from 4.7
to 27.7 cm TL, with 21 individuals < 15 cm TL (Figure 3).
The largest specimen caught was the only sexually mature
individual recorded.
DISCUSSION
The lack of sightings prior to December 2007, the
exponential increase in sightings over time, and the
predominance of immature and small individuals suggest
that the lionfish invasion of South Caicos is in its initial
stages. Although the effects of the invasion are unknown,
some results are of concern. Lionfish were observed in
blow-out ledges in seagrass habitats (5 during the study
and 11 thereafter). These structures serve as important
microhabitat for juvenile Nassau groupers (Epinephelus
striatus) around South Caicos (Claydon & Kroetz 2007;
see Figure 3D). A continued exponential increase of
lionfish may lead to inter-specific competition for space in
blow-outs. This could impact populations of the regionally
endangered Nassau grouper. Although the majority of
lionfish were found on reefs and at shallow depths, this
may partly reflect disparity in sampling effort. Future
Figure 2. A. Lionfish sightings per month from August
2007 to July 2008. B. Frequency of lionfish observed in
reef, seagrass and mangrove habitats (n = 25). C. Size
frequency distribution of lionfish caught around South
Caicos (n = 23).
Figure 3. A. The smallest (4.7cm TL) and B. the largest
(27.7cm TL) lionfish captured. C. Lionfish in blowout ledge in
seagrass habitat. D. Juvenile Nassau grouper in blowout
ledge.
Page 402
61st Gulf and Caribbean Fisheries Institute
ACKNOWLEDGEMENTS
We would like to acknowledge: The School for Field Studies –
Center for Marine Resource Studies for financial and logistical support;
TCI Government Department of Environment and Coastal Resources for
supporting the project; and for assistance in the field provided by staff and
students of School for Field Studies – Center for Marine Resource Studies
and the staff of the East Bay Development.
LITERATURE CITED
Claydon, J.A.B. and A. Kroetz. 2008. The distribution of
early juvenile groupers around South Caicos,
Turks & Caicos Islands. Proceedings of the Gulf
and Caribbean Fisheries Institute 60:345-350.
Whitfield PE, J.A. Hare, A.W.David, S.L. Harter, R.C.
Muñoz, and C.M. Addison. 2007. Abundance
estimates of the Indo-Pacific lionfish. Biological
Invasions 9:53-64.
Abundance of Invasive Lionfish (Pterois volitans) on Bahamian Coral Reefs
S.J. GREEN and I.M. Côté
Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, BC, V5A 1S6 Canada
EXTENDED ABSTRACT
Indo-Pacific lionfish (Pterois volitans and P. miles) have recently invaded and rapidly spread throughout temperate and
tropical Western Atlantic and Caribbean habitats. The invasion is thought to be the result of accidental or intentional
releases from aquaria off the southeast coast of Florida, with recent genetic work suggesting a small founding population
(Freshwater et al. 2009). Lionfish abundances have been increasing exponentially at several locations in the Caribbean over
the past few years, and individuals have now been sighted in the Caribbean as far south as Panama. These venomous,
invasive predators use an ambush strategy to consume whole prey fish and have few predators in their introduced range.
Lionfish were first reported from Western Atlantic coral reef habitats in 2004, off the southwest coast of New Providence Island, Bahamas, and are now abundant throughout the archipelago (REEF 2008). Despite their popularity in the
aquarium trade, there is little scientific information about the abundance of lionfish from coral reefs in their native range,
and no information from coral reef habitats in Caribbean.
We set out to assess the population abundance and size distribution of lionfish inhabiting sites along a continuous coral
reef off the southwest coast of New Providence, Bahamas in 2008. At each of the 13 sites, we assessed lionfish abundance
along six 50 m x 10 m transects. Transects were conducted at depths of between 10 and 20 m, and total length of each
lionfish encountered was estimated to the nearest cm by trained observers.
At several sites, densities of P. volitans exceeded those reported anecdotally from their native range in the Red Sea by
Fishelson (1997), as well as density estimates from their invaded range along the eastern coast of the US by Whitfield et al.
(2007) (Table 1). The impacts of the lionfish invasion are generating great concern, with individual lionfish having been
shown to reduce recruitment of native reef fish by 79% on small experimental reefs in the Bahamas (Albins and Hixon,
2008). Given the high densities we documented across our study sites, the impacts of lionfish on natural reefs in the invaded
range may be severe.
These abundance estimates can be used in conjunction with data on fish community assemblage and habitat metrics
from the study system to predict the distribution of lionfish across coral reef habitats. As lionfish continue to spread and
increase in abundance across the region, these data may be used to target areas of management and control priority.
LITERATURE CITED
Table 1. Density and total length (TL) of Pterois volitans
on coral reef sites off southwest New Providence, Bahamas. Means are shown ± 1 SD.
Site
Density
TL (cm)
1
345
±
165
22
±
6
2
233
±
82
24
±
7
3
213
±
177
24
±
7
4
141
±
122
23
±
5
5
99
±
45
25
±
7
6
65
±
30
21
±
7
7
65
±
55
20
±
7
8
40
±
10
28
±
3
9
28
±
30
18
±
5
10
28
±
27
20
±
6
11
27
±
10
20
±
6
12
27
±
35
24
±
5
13
10
±
12
30
±
3
Ablins, MA and MA Hixon. 2008. Invasive Indo-Pacific lionfish
Pterois volitans reduce recruitment of Atlantic coral-reef fishes.
Marine Ecology Progress Series 367:233-238.
Fishelson, L. 1997. Experiments and observations on food
consumption, growth and starvation in Dendrochirus
brachypterus and Pterois volitans (Pteroinae, Scorpaenidae).
Environmental Biology of Fishes 50(4):391-403.
Freshwater, D.W. et al. 2009). Mitochondrial control region
sequence analyses indicate dispersal from the US East Coast as
the source of the invasive Indo-Pacific lionfish Pterois volitans
in the Bahamas. Marine Biology 156(6):1213-1221.
REEF. (2009). Reef Environmental Education Foundation Volunteer
Survey Project Database. www.reef.org. (01/11/2008).
Whitfield, P.E., J.A. Hare, A.W. David, S.L. Harter, R.C. Muñoz, and
C.M. Addison. 2007. Abundance estimates of the Indo-Pacific
lionfish Pterois volitans/miles complex in the Western North
Atlantic. Biological Invasions 9(1):53-64.
Biology and Ecology of the Invasive Lionfishes, Pterois miles and Pterois volitans
JAMES A. MORRIS, JR.1, J.L. AKINS2, A. BARSE3, D. CERINO1, D.W. FRESHWATER4,
S.J. GREEN5, R.C. MUÑOZ1, C. PARIS6, and P.E. WHITFIELD1
1
National Oceanic and Atmospheric Administration, 101 Pivers Island Road, Beaufort, North Carolina 28516 USA ,2Reef
Environmental Education Foundation, 98300 Overseas Hwy, Key Largo, Florida 33037 USA, 3Department of Biology,
Salisbury University, 1101 Camden Avenue, Salisbury, Maryland 21801 USA, 4University of North Carolina Wilmington, Center for Marine Science, 5600 Marvin Moss Lane, Wilmington, North Carolina 28409 USA, 5Department of
Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6, Canada,
6
Applied Marine Physics, Rosenstiel School of Marine and Atmospheric Science, 4600 Rickenbacker Causeway, Miami,
Florida 33149 USA
ABSTRACT
The Indo-Pacific lionfishes, Pterois miles and P. volitans, are now established along the U.S. southeast coast, Bermuda,
Bahamas, and are becoming established in the Caribbean. While these lionfish are popular in the aquarium trade, their biology and
ecology are poorly understood in their native range. Given the rapid establishment and potential adverse impacts of these invaders,
comprehensive studies of their biology and ecology are warranted. Here we provide a synopsis of lionfish biology and ecology
including invasion chronology, taxonomy, local abundance, reproduction, early life history and dispersal, venomology, feeding
ecology, parasitology, potential impacts, and control and management. This information was collected through review of the
primary literature and published reports and by summarizing current observations. Suggestions for future research on invasive
lionfish in their invaded regions are provided.
KEY WORDS: Lionfish, invasive species, Pterois
Biologia y Ecologia del Pez Leon Invasor, Pteoris miles y Pterois volitans
Los peces león del Indo-Pacifico, Pterois volitans and Pterois miles, se están estableciendo a lo largo de la costa sur oriental
de los Estados Unidos, Bermuda, Bahamas y han comenzado a invadir el Caribe. Aunque los peces leones son populares en el
comercio de acuarios, poco es conocido de su biología y ecología. Dado el rápido establecimiento de los peces león y su impacto
potencial de estos invasores, los estudios comprensivos acerca de su biología y ecología son necesarios. Aquí proporcionamos una
sinopsis de la biología y ecología del pez león incluyendo la cronología de la invasión, taxonomía, abundancia local, reproducción,
historia temprana de la vida y dispersión, venomology, ecología de alimentación, parasitología, los impactos potenciales, y control y
gerencia. Esta información fue recogida por medio de la revisión de la literatura primaria y informes públicos y resumiendo
observaciones actuales. Las sugerencias para la investigación futura sobre el invasor pez león en las regiones invadidas son
proporcionadas.
PALABRAS CLAVES: Peces león, especie invasora, Pterois
Biologie et Ecologie de Rascasses volantes Invasives, Pteoris miles et Pterois volitans
Les rascasses volantes, Pterois volitans et Pterois miles, originaires de la zone tropicale indo-Pacifique sont aussi retrouvées le
long des côtes sud-est américaines, aux Bermudes, aux Bahamas, et sont actuellement entrain d’envahir les Caraïbes. Alors que ces
poissons sont très demandés dans le commerce des poissons d’aquarium, peu de données relatives à leur biologie et à leur écologie
sont connues. Dans ce contexte, d’invasion rapide de ce poisson et des impacts potentiels sur les communautés de poissons récifaux
endogènes, nous essayons de mettre en évidence la biologie de reproduction, les habitudes alimentaires et les caractères venimeux en
utilisant les observations menées en laboratoire et sur le terrain. Concernant la reproduction de ce poisson, les observations menées
montrent que c’est un animal itéropare, asynchrone avec de multiples pontes par saison (le nombre de ponte étant indéterminé). Les
mesures visant à déterminer la périodicité de ponte montrent que ces poissons pondent mensuellement, avec des périodes de ponte
au cours de la plupart des mois du calendrier d’où le caractère invasif de cette espèce. Les expérimentations conduites au laboratoire
sur la prédation des rascasses juvéniles ont montré que ces derniers ne constituent pas de véritables proies pour les poissons récifaux
endémiques à cause de leurs défenses venimeuses. L’analyse du contenu stomacal révèle essentiellement des crustacées et des
poissons des espèces fourragères incluant les poissons dévolus à la pêche commerciale ou de loisirs, comme le vivaneau et le mérou.
Ces travaux fournissent un nouvel éclairage en ce qui concerne la biologie intégrée et l’écologie des rascasses volantes non
endémiques, et démontrent le besoin d’une détection précoce et systématique de cette espèce et la mise en œuvre de solution rapide
pour faire face à cette invasion dans l’écosystème marin.
MOTS CLÉS: Rascasses volantes ,espèce invasion, Pterois
INTRODUCTION
The lionfish invasion in the Northwestern Atlantic and
the Caribbean represents one of the most rapid marine
finfish invasions in history. Despite being a popular
member of the marine ornamental aquarium trade, little
was known regarding the biology and ecology of these
lionfishes prior to this invasion. Information on lionfish
abundance, dietary habits, predators, and seasonality of
reproduction are scarce. Most of what has been published
on lionfish relates largely to lionfish envenomations, which
Page 410
commonly occur during aquarium husbandry or as a result
of poor handling by home aquarists.
Invasive lionfish are a concern to coastal managers due
to their potential threat on fisheries resources, native fish
communities, and human health. Since 2000, National
Oceanic and Atmospheric Administration (NOAA)
researchers have partnered with non-governmental
organizations, academics, and other federal and state
agencies to develop a programmatic response to the
lionfish invasion. The following provides a synopsis of
information on the biology and ecology of the invasive
lionfishes that have invaded the Northwestern Atlantic and
Caribbean, and a discussion of future research needs and
management options.
Invasion Chronology
Many non-native marine ornamental fishes have
been reported along the U.S. East Coast, with a “hotspot”
of introductions occurring in South Florida (Semmens et
al. 2004). Lionfish have been documented off Palm Beach,
Boca Raton, and Miami, Florida beginning in 1992; and
Bermuda, North Carolina, South Carolina and Georgia
beginning in 2000 (Hare and Whitfield 2003, REEF 2008,
USGS 2008, Whitfield et al. 2002). Since 2004, lionfish
have become widespread in the Bahamas (REEF 2008,
USGS 2008, Whitfield et al. 2007). More recently,
lionfish were reported in the Turks and Caicos and Cuba
(Chevalier et al. 2008) in 2007, and in the Cayman Islands,
Jamaica, Dominican Republic (Guerrero and Franco 2008),
U S. Virgin Islands, Belize, and Barbados in 2008 (REEF
2008, USGS 2008). Juvenile lionfishes have also been
reported along the U.S. northeast coast including Virginia,
New York, Rhode Island, and Massachusetts since 2001.
These northeastern specimens are incapable, however, of
overwintering due to thermal intolerance (Kimball et al.
2004), and they are not considered established.
It is nearly impossible to determine which introduction
event(s) allowed lionfish to become established. Research
on the genetic variation of the lionfish populations is
providing insight into the minimum number of lionfish and
the geographic origin of founder population(s) (Hamner et
al. 2007). Interestingly, this is not the first documented
invasion of Pterois sp. as Golani and Sonin (1992) reported
a Mediterranean invasion of P. miles from the Indian
Ocean via the Suez Canal.
Taxonomy
Pterois miles and P. volitans are morphologically
similar and distinguishable in their native range by
meristics, with P. volitans exhibiting one higher count of
dorsal and anal fin rays when compared to P. miles. This
difference was documented by Schultz (1986) who
reported that P. miles is found in the Red Sea, Persian Gulf,
and Indian Ocean (excluding Western Australia) and P.
volitans is found in the Western and Central Pacific and
Western Australia. Kochzius et al. (2003) used mitochon-
drial DNA analyses to show that specimens identified as P.
miles and P. volitans were genetically distinct. Their
geographic sampling did not allow the determination of
whether this distinction was at the species or population
level. Hamner et al. (2007) analyzed specimens identified
as P. miles and P. volitans from additional areas of their
native range, including Indonesia, where they are sympatric. They found that the two taxa are clearly distinct
supporting the designation of two species. Analyses with
different molecular markers and additional geographic
samples of species in Pterois and the out-group comparison
with the closely related genus Dendrochirus, support the
classification of P. miles and P. volitans as separate
species. Recent efforts by Hamner et al. (2007) have
confirmed that:
i) Both P. miles and P. volitans were introduced
along the U.S. East Coast,
ii) P. volitans comprises approximately 93% of the
population, and
iii) A strong founder effect (i.e. low genetic diversity)
is evident among Atlantic specimens.
The genetic structure of invasive lionfish in the
Caribbean is presently unknown. Only one species (P.
volitans) has been confirmed along the Bahamian archipelago. Documentation of genetic change and adaptation of
lionfish populations in their invaded range is warranted
(e.g., Morris and Freshwater 2008). Greater understanding
of lionfish genetics could assist with validation of reef fish
dispersal and connectivity models in the Northwestern
Atlantic, Caribbean, and Gulf of Mexico.
Local Abundance
Whitfield et al. (2007) provided the first assessment of lionfish densities off North Carolina and reported
an average of 21 lionfish per hectare across 17 locations in
2004. Lionfish densities off North Carolina have continued to increase. Recent assessments off New Providence,
Bahamas indicate lionfish densities are more than 18 times
higher than the 2004 North Carolina estimates (Green and
Côté 2008). The cryptic nature of lionfish make them
difficult to census. It is likely that estimates of lionfish on
complex coral reef habitats under-represent local abundance of juveniles. Thus, these density estimates should be
considered conservative. Further, lionfish densities in the
Bahamas are more than eight times higher than estimates
from their native range (Green and Côté 2008). Few
published data are available, however, from the IndoPacific region providing high uncertainty for this comparison. In their invaded Atlantic and Caribbean ranges, it is
unclear when lionfish densities will reach carrying
capacity. Given that many reef fishes along the east coast
of the U.S. and Caribbean are overfished (Hare and
Whitfield 2003), lionfish might be utilizing vacated niche
attributes such as increased availability of forage fishes and
reef space.
Morris, J. et al. GCFI:61 (2009)
Monitoring of lionfish densities across habitat types
using standardized indices of abundance is needed to
determine when lionfish abundances reach carrying
capacity. Lionfish densities are expected to vary depending on such factors as seasonality, local recruitment, local
niche availability, and fishing pressure. Studies assessing
the drivers controlling lionfish densities in specific habitats
are needed to support lionfish control measures and to
identify potential pathways for new invaders.
Reproduction
The Pteroines, including P. miles and P. volitans,
are gonochoristic; males and females exhibit minor sexual
dimorphism only during reproduction (see Fishelson 1975).
Lionfish courtship has been well described by Fishelson
(1975) who provided a detailed description for the pigmy
lionfish, Dendrochirus brachypterus, and reported similar
courtship behaviors for Pterois sp. According to Fishelson,
lionfish courtship, which includes circling, side winding,
following, and leading, begins shortly before dark and
extends well into nighttime hours. Following the courtship
phase, the female releases two buoyant egg masses that are
fertilized by the male and ascend to the surface. The eggs
and later embryos are bound in adhesive mucus that
disintegrates within a few days, after which the embryos
and/or larvae become free floating.
P. miles and P. volitans ovarian morphology is similar
to that reported for D. brachypterus (Fishelson 1978) in
that these fishes exhibit cystovarian type ovaries (Hoar
1957) with ooctyes developing on stalks or peduncles. The
oocytes are terminally positioned near the ovary wall,
which secretes the encompassing mucus shortly before
spawning.
The seasonality of lionfish reproduction
throughout their native range is unknown. Invasive
lionfish collected off North Carolina and in the Bahamas
suggests that lionfish are reproducing during all seasons of
the year.
Early Life History and Dispersal
Larval stage descriptions for P. miles and P.
volitans are incomplete with only one report by Imamura
and Yabe (1996) describing five P. volitans larvae
collected off northwestern Australia. Scorpaenid larvae
exhibit two morphologically distinct groups characterized
as “morph A” and “morph B” by Leis and Rennis (2000).
Pteroine larvae are grouped among the “morph B”
morphotypes, whose traits include: large head, relatively
long and triangular snout, long and serrated head spines,
robust pelvic spine, and pigment confined to the pectoral
fins (Leis and Rennis 2000) and postanal ventral and dorsal
midlines (Washington et al. 1984). Pterois sp. meristic
characters are reported as 12 - 13 dorsal spines, 9 - 12
dorsal rays, three anal spines, 5 - 8 anal rays, 12 - 18
pectoral rays, one pelvic spine, five pelvic rays, and 24
vertebrae (Imamura and Yabe 1996; Leis and Rennis
2000).
Page 411
The size of P. miles or P. volitans larvae at hatching is
unmeasured, but is likely to be approximately 1.5 mm
based on reports for P. lunulata (Mito and Uchida 1958;
Mito 1963). The specific planktonic larval duration of
lionfish is also unknown, although Hare and Whitfield
(2003) estimated it to be between 25 to 40 days based
estimates for Scorpaena (Laidig and Sakuma 1998).
Dispersal of lionfish presumably occurs during the
pelagic larval phase during which larvae can be dispersed
across great distances. For example, lionfish eggs released
in the Bahamas are capable of dispersing to New England
via the Gulf Stream. Larval connectivity models for reef
fishes (e.g., Cowen et al. 2006) provide insight into
lionfish larval dispersal and are valuable for predicting the
spread of lionfish as evidenced by the recent establishment
of lionfish in the Caribbean. Further lionfish dispersal into
the lower Caribbean and the Gulf of Mexico seems
imminent. Assuming a planktonic larval duration of 25 to
40 days (Hare and Whitfield 2003), the Caribbean and
Yucatan currents are capable of dispersing lionfish larvae
into the Gulf of Mexico from locations in the Caribbean
where lionfish are already resident (i.e., Cuba, Jamaica,
Cayman Islands) (Cowen et al. 2006). Based on the
rapidity of lionfish establishment along the U.S. East Coast
and the Bahamas, lionfish establishment along the southern
edges of Central America (Nicaragua, Costa Rica, and
Panama), the Yucatan peninsula, and the western Gulf of
Mexico is likely within a few years or less. Establishment
would also be facilitated by gyres such as the ColumbiaPanama Gyre and the Gulf of Mexico loop current, which
could provide a mechanism for lionfish to become
established in the Florida Keys.
Venomology
Lionfish are venomous with their spines containing
apocrine-type venom glands. Each spine of the lionfish
(except caudal spines) is venomous including 13 dorsal
spines, three anal spines, and two pelvic spines. The spines
are encased in an integumentary sheath or skin and contain
two grooves of glandular epithelium that comprises the
venom producing tissue. Spine glandular tissue extends
approximately three quarters the distance from the base of
the spine towards the tip (Halstead et al. 1955).
Lionfish envenomation occurs when the spine’s
integumentary sheath is depressed as it enters the victim.
This process tears the glandular tissue allowing the venom
to diffuse into the puncture wound (Saunders and Taylor
1959). The toxin in lionfish venom contains acetylcholine
and a neurotoxin that affects neuromuscular transmission
(Cohen and Olek 1989). Lionfish venom has been found to
cause cardiovascular, neuromuscular, and cytolytic effects
ranging from mild reactions such as swelling to extreme
pain and paralysis in upper and lower extremities (Kizer et
al. 1985). Antivenom of the related stonefish (Synanceia
ssp.) is highly effective in neutralizing lionfish venom
activity (Shiomi et al. 1989, Church and Hodgson 2002).
Page 412
The severity of sting reactions in humans is dependent
upon such factors as the amount of venom delivered, the
immune system of the victim, and the location of the sting.
Records of home aquarists stung by lionfish provide a
comprehensive assessment of how lionfish stings affect
humans (Kizer et al. 1985, Vetrano et al. 2002). The
probability of lionfish envenomation is higher when
handling smaller-sized lionfish because the venom
glandular tissue is closer to the tip of the spine (Halstead et
al. 1955).
The effectiveness of lionfish venom defense in their
invaded range is in question. Maljković et al. (2008)
reported that lionfish were found in the stomachs of
groupers; however, this observation provides no assessment of the frequency of lionfish consumption by grouper.
Furthermore, laboratory behavioral experiments suggest
that groupers actively avoid lionfish, even during periods
of extreme starvation. Additional research is needed
towards understanding predatory interactions between
lionfish and native predators.
Work by Sri Balasubashini et al. (2006a, 2006b)
indicated that lionfish (P. volitans) venom contains
antitumor, hepatoprotective, and antimetastatic effects in
mice suggesting a promising application for cancer
research. Depending on the outcome of this research and
the subsequent demand for lionfish venom, bioprospecting
of venom from invasive lionfish could assist with fishery
development.
Feeding Ecology
In the Red Sea, lionfish (P. miles) have been
reported to feed on assorted taxa of benthic fishes including damselfish, cardinal fish, and anthias (Fishelson 1975,
Fishelson 1997). However, in the Pacific Ocean, P.
lunulata were observed to feed primarily on invertebrates
including penaeid and mysid shrimps (Matsumiya et al.
1980, Williams and Williams 1986). Assessments of
invasive lionfish feeding suggests that lionfish are largely
piscivorous, but also feed on a number of crustaceans. The
particular taxa of highest importance in invasive lionfish
diet will likely vary by habitat type and prey availability.
Feeding, growth, and starvation of P. volitans from the
Red Sea was investigated by Fishelson (1997) who
reported that lionfish stomachs can expand over 30 times in
volume after consuming a large meal. This capability
supported Fishelson’s hypothesis that lionfish were capable
of longterm fasting, and demonstrated their ability to
withstand starvation for periods of over 12 weeks without
mortality. Fishelson (1997) also measured daily consumption rates in the laboratory for six size classes of lionfish
ranging from 30 - 300g and found that lionfish consumed
approximately 2.5 – 6.0% of their body weight per day at
25 - 26 °C. Preliminary observations suggest that lionfish
in their invaded range can consume piscine prey at rates
greater than reported earlier by Fishelson (1997). Quantification of the feeding ecology of lionfish including
consumption rates and prey selectivity will permit better
assessment of the impacts of their predation on local reef
fish communities.
Parasitology
Knowledge of the parasites infecting native and nonnative lionfish is scant. No comprehensive survey of
protozoan or metazoan parasites of either host (P. miles or
P. volitans) has been published. There are, however, a few
isolated records of single parasite species such as monogeneans from the Red Sea (Paperna 1972, Colorni and
Diamant 2005) and Japan (Ogawa et al. 1995), copepods
also from Japan (Dojiri and Ho 1988), and leeches from
Japan (Paperna 1976) and the Florida coast (Ruiz-Carus et
al. 2006). Most published records of lionfish parasites are
of ectoparasites; the only record of an endoparasite is of a
new myxosporean species, Sphaeromyxa zaharoni which
was found in a lionfish gall bladder from the Red Sea
(Diamant et al. 2004). Recent observations of invasive
lionfish collected off North Carolina and in the Bahamas
have found low prevalence of endo- and ectoparasites
when compared to parasites of native reef fishes. Future
research describing parasites of invasive lionfish will
provide a unique study of opportunistic parasitism by
common parasites of marine reef fishes.
Potential Impacts
Potential ecological impacts of lionfish on local
reef fish communities will vary depending on the abundance of top level predators, the forage fish community,
the density of lionfish, and the geographic location. Local
studies that provide observations of lionfish impacts on
community structure and the abundance of forage fishes
are needed. The first evidence of lionfish impacts in their
new range was provided by Albins and Hixon (2008) who
reported a 79% reduction in forage fish recruitment on
experimental patch reefs in the Bahamas during a five
week observation period. Analysis of the potential impact
of lionfish consumption on whole coral reef fish communities is also being documented in the Bahamas, where data
on stomach contents are being combined with abundance
estimates of the prey community across various habitat
types and seasons. Given the high levels of lionfish
biomass found at some locations (Whitfield et al. 2007),
the predatory removal of forage fishes is a growing
concern, because many other top level predators (i.e.,
potential food competitors with lionfish) are overfished or
in low abundance (Hare and Whitfield 2003).
It is unclear if lionfish predation on economically
important species such as juvenile serranids will harm
stock rebuilding efforts. Economically important species
were relatively low in importance in the lionfish diet of the
Bahamas, but this could be a direct result of their low
abundance in the forage fish community. Research that
monitors lionfish predation on economically important
species is needed.
Morris, J. et al. GCFI:61 (2009)
Lionfish impacts on tourist recreational activities have
been observed. Some locations have posted warning signs
advising of the potential for lionfish envenomation. As
lionfish densities increase, so too does the risk of envenomations. It is unknown whether increasing lionfish
densities will reduce recreational activities and cause
economic hardship. This will be dependent on factors such
as the prevalence of warning signs, the density of lionfish,
and the effectiveness of education and outreach.
Control and Management
Management of marine finfish invasions are confounded by highly diverse and wide-ranging habitats, swift
ocean currents, and jurisdictional constraints. Prevention is
the least expensive and most effective management option.
There are currently two lionfish management and control
efforts in Bermuda and the Bahamas. Bermuda initiated a
lionfish culling program in 2006 that included a training
program, collecting license, and a special dive flag
allowing commercial and recreational fishers to spear
lionfish along nearshore reefs. A video description of this
program can be seen at http://www.youtube.com/watch?
v=LNbKjiUCGRU . Bahamian fisheries officials instituted
a lionfish kill order to fishermen in 2005. They have also
actively engaged the public with educational seminars
devoted to promoting lionfish as a food fish with the hopes
that human consumption will support fishery development.
Grassroots, “adopt a reef programs”, are being developed
in Eleuthra (see www.lionfishhunter.com) that encourage
local citizens to take ownership of small reefs and to
protect them from lionfish impacts. Some tourist locations,
such as resorts, are physically removing lionfish by
spearfishing and handnets to reduce the risk of swimmer
interaction. It is unclear how effective these approaches
will be, because too little is known about the rate of
lionfish recruitment and movement among the various
habitat types. Recently, NOAA researchers have developed techniques to trap lionfish, thus providing a means of
removal from deeper waters and larger areas that are
impractical for diver removal.
CONCLUSIONS
The lionfish introduction provides a reminder of how
rapid a non-native species can become established and
potentially compete with native fishes for resources. Early
detection and rapid response efforts are of utmost importance in the marine environment due to the complexity and
ineffectiveness of eradication measures. An early detection
and rapid response program has been developed in south
Florida (a hotspot for marine introductions), which utilizes
and coordinates resources from over thirty state, federal,
and non-governmental organizations in the region.
Programs such as this represent the first line of defense for
marine introductions and should be endorsed and supported
by local managers. Future research on invasive lionfish
should focus on understanding and reducing their ecologi-
Page 413
cal impacts, the scale of which is yet to be determined.
ACKNOWLEDGEMENTS
We thank the NOAA Aquatic Invasive Species Program, NOAA
National Centers for Coastal Ocean Science, the NOAA Undersea
Research Program (Grant No. NA030AR4300088), the National Science
Foundation (Grants OCE0825625, OCE0550732, PEET No.0328491,
DBI-MRI No.0618453), the GEF Coral Reef Targeted Research Program,
Connectivity Working Group, and the Natural Sciences and Engineering
Research Council of Canada for funding support. We also thank J.
Langston and P. Schofield (USGS) for providing lionfish sightings
records and D. Ahrenholz and D. Evans for their helpful review of this
manuscript.
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The Invasion of Indo-Pacific Lionfish in the Bahamas: Challenges for a National Response Plan
1
KATHLEEN SULLIVAN SEALEY1, 4, LAKESHIA ANDERSON2, DEON STEWART3, and NICOLA SMITH4, 5
University of Miami Department of Biology, Coral Gables, Florida USA; 2 Department of Marine Resources, Nassau,
Bahamas; 3 Bahamas Environment, Science and Technology Commission, Nassau, Bahamas;
4
Marine and Environmental Studies Institute, College of The Bahamas, Nassau, Bahamas;
5
Department of Zoology, University of British Columbia, Vancouver, BC, Canada
ABSTRACT
The invasion of the Indo-Pacific lionfish to Bahamian waters raises considerable concern due to the uncertainty of its
ecological impacts and its potential threats to commercial fisheries and human safety. Lionfish have been reported throughout the
archipelago and are the focus of several research and monitoring initiatives. The Bahamas has a National Invasive Species Strategy,
but marine invasions require unique partnerships for small islands developing states to develop realistic management goals and
actions. The Government of The Bahamas has limited funds to address major resource management issues; hence, collaboration
with non-governmental agencies and tertiary education institutions is imperative. The establishment and spread of lionfish has
created a novel opportunity for the formation of innovative public-private partnerships to address the ecological, economic and
social impacts of biological invasions.
KEY WORDS: Lionfish, invasion, reefs
La Invasión en las Bahamas del Pez León del Indo-Pacifico:
Un Caso de Investigación, Planificación, y Manejo
La invasión del pez león del Indopacífico en las aguas de Las Bahamas ha generado una gran preocupación debido a la
incertidumbre sobre su efecto ecológico y la posible afectación a la pesca comercial, el turismo y la seguridad de la población. El
pez león se ha registrado en todo el archipiélago y ha sido objeto de investigación y monitoreo. Aunque Las Bahamas posee un
Plan Estratégico Nacional para las Especies Invasoras, las marinas requieren de esfuerzos institucionales conjuntos para recopilar y
analizar la información, lanzar iniciativas educativas, y elaborar medidas realistas de manejo. El gobierno de Las Bahamas posee
fondos limitados para enfrentar los problemas prioritarios de manejo; por eso es indispensable la colaboración con agencias no
gubernamentales e instituciones terciarias de educación. La invasión del pez león ha creado una oportunidad para la formación de
asociaciones innovadoras entre instituciones privadas y públicas con el fin de abordar las consecuencias ecológicas, sociales y
económicas de los invasores biológicos.
PALABRAS CLAVES: Pez león, invasión, arrecifes
L’invasion des Rascasses de L’Indo-Pacifique aux Bahamas:
Problèmes de Recherche, D’éducation et de Gestion
L’invasion de la rascasse de l’indo-pacifique dans les eaux des Bahamas génère une grande préoccupation liée à l’incertitude
de ses impacts écologiques et de ses répercussions possibles sur la pêche commerciale, le tourisme et la santé humaine. La rascasse
a été observée sur l’ensemble de l’archipel et fait l’objet de plusieurs programmes d’études et de surveillance. Les Bahamas
possèdent un plan stratégique national contre les espèces invasives mais les espèces marines nécessitent des collaborations
institutionnelles fortes pour compiler et analyser les informations, promouvoir des programmes éducatifs et développer des moyens
réalistes de gestion. Le gouvernement des Bahamas possède des fonds limités pour répondre à ces questions majeures de gestion des
ressources, c’est pourquoi il est indispensable d’avoir des collaborations avec des agences non gouvernementales et des institutions
éducatives. L’invasion des rascasses a créé une nouvelle opportunité pour la formation d’associations innovantes public-privé dans
le but d’analyser les conséquences écologiques, sociales et économiques des envahisseurs biologiques.
MOTS CLÉS: Rascasse, Bahamas
INTRODUCTION
The Commonwealth of The Bahamas faces significant
challenges in the management and protection of marine
resources over the 1,200 kilometer-long archipelago. Stressors on the marine environment include overfishing, marine and terrestrial sources of pollution, climate
change, and invasive species. Biological invasions involve
both human and non-human mediated forms of dispersal in
which non-native species successfully arrive, survive, and
reproduce in a novel locality and then spread throughout a
region (Carlton 1987, 1989). The establishment and spread
of exotics represent one of the most pressing threats to the
diversity and distinctiveness of ecological communities
(see Clavero and Garcia-Berthou 2005, Lowe et al. 2000,
Sax and Gaines 2008, McKinney and Lockwood 1999,
2005, Vitousek et al. 1987). Moreover, the rate and
magnitude of species introductions are increasing worldwide while the economic costs associated with the impacts
and control of invaders are estimated in billions of dollars
per year in the US alone (Lowe et al. 2000, Pimentel et al.
2000, 2005).
Sullivan-Sealey, K. et al.
Addressing the current state of invasions is therefore a
daunting task for even the most well-equipped management teams. Invasive species management underscores the
long-standing tension between decision-makers’ need for a
rapid, cost-effective and yet, scientifically-informed
response to environmental issues and the incremental
nature of advances in scientific knowledge. On the one
hand, emergent crises demand a timely and viable response
to both real and perceived public concerns. On the other
hand, the technical and scientific expertise required to
guide management options often involves a comparably
slow development process. This strain is further exacerbated in small island developing states in which the threat
from invasions is high and an immediate concern but
national capacity to address the issue in terms of monetary
resources and scientific expertise is limited. The recent
introduction of the venomous Indo-Pacific lionfish (Pterois
volitans and P. miles) to the Western Atlantic Ocean brings
to focus the substantial challenges faced by small developing nations such as The Bahamas to effectively manage
high priority invaders and develop realistic prevention and
early detection programs for other exotics.
Invasive species management is a long-term proposition that entails partnerships between both local and
regional governmental and non-governmental agencies.
Accurate and continually updated scientific knowledge in
addition to ongoing monitoring is critical to understanding
the risks associated with lionfish establishment and spread.
The National Invasive Species Strategy for The Bahamas
provides a good starting point for work in this area (BEST
2003), but the unique challenges presented by such a
widespread and venomous marine invader may require a
greater degree of innovation. This paper presents a brief
overview of:
i) The unique challenges in creating a NRLP,
especially in establishing a central focused goal;
ii) Management options for a NRLP, and
iii) Assessment of stakeholder grounds, and potential
partnerships for a NRLP.
Part of the introduction to the management issues is a
short review of lionfish ecology and the invasion history.
A summary of events related to the Lionfish invasion of
The Bahamas is presented in Appendix 1.
LIONFISH ECOLOGY AND INVASION HISTORY
Lionfish (Pteroinae) are a subfamily of the scorpionfishes (Scorpaenidae). There are 17 different species of
lionfish that occur within five different genera (Kochzius et
al. 2003). All Pterois species contain venomous dorsal,
anal and pelvic spines (Randall et al. 1997). Pterois
volitans (Linnaeus 1758) and a closely related form, P.
miles (Bennett 1828), share similar morphological
resemblance and are generally considered to be allopatric
sibling species native to the sub-tropical and tropical
waters of the Indo-Pacific, where they are carnivorous midlevel predators on coral, rocky and sandy substrates
GCFI:61 (2009)
Page 405
(Schultz 1986, Fishelson 1997). P. volitans has a pelagic
egg and larval stage, matures between 180 - 190mm and
can grow up to 350 mm or more in total length (Fishelson
1997, Imamura and Yabe 1996). Lionfish have few natural
predators most likely due to the venomous nature of the
species. In the Indo-Pacific, Bernadsky and Goulet (1991)
report an isolated case of a pacific cornetfish Fistularia
commersonii that consumed a juvenile P. miles, while in its
invaded range of The Bahamas, groupers may be a
potential predator. This is supported by the discovery of
juvenile P. volitans in the stomachs of adult Nassau
grouper, Epinephelus striatus, and tiger grouper, Mycteroperca tigris (Maljkovic et al. 2008).
Lionfish occur outside of their native range in the
Mediterranean Sea as well as the Atlantic Ocean. Pterois
miles entered the Mediterranean Sea via the Suez Canal as
a Lessepsian migrant (Golani and Sonin 1992) while both
P. volitans and P.miles were most likely introduced to the
Western Atlantic in the early 1990s off the coast of Florida
via aquarium releases (Courtney 1995, Hare and Whitfield
2003; Ruiz-Carus et al. 2006, Semmens et al. 2004,
Whitfield et al. 2002). P. volitans is now established and
abundant along the US southeast continental shelf,
Bermuda and several Caribbean countries including The
Bahamas, The Cayman Islands, Cuba, and The Turks and
Caicos (Albins and Hixon 2008, Hare and Whitfield 2003,
Meister et al. 2005, Ruiz-Carus et al. 2006, Snyder and
Burgess 2006, USGS NAS Database 2008, Whitfield et al.
2002, 2007). Since 2008, lionfish have also been reported
in the Dominican Republic, Jamaica and St. Croix (Guerro
and Franco 2008, USGS NAS Database 2008). Unlike P.
volitans, which is now widespread and abundant in the
Atlantic, P.miles has only been observed in its invaded
range in relatively small numbers along the US eastern
seaboard (Hamner et al. 2007). The overall impact of
lionfish in the Atlantic remains largely unknown, but many
suggest that the species will negatively affect marine
communities by decreasing the abundance of a wide range
of reef associated fish via direct predation and competitive
interactions in which lionfish monopolize food resources
(see Albins and Hixon 2008, Hare and Whitfield 2003,
Whitfield et al. 2007).
UNIQUE CHALLENGES TO INVASIVE SPECIES
MANAGEMENT IN THE BAHAMAS
The Bahamian archipelago is a unique ecological
system, with real limitations to how much humans can alter
the landscape without irreversible changes to the environment. Invasive alien species has been recognized an
important cause for the loss of national biological diversity
of both natural communities and species. The primary
environmental sensitivities are related to water use, nutrient
cycles and island hydrology, and link coastal land use to
the adjacent marine resources. Thus, the vulnerability of
The Bahamas to near shore marine invasive species is very
high. The Government agencies and non-government
Page 406
organizations have a high level of problem recognition and
awareness, yet the general population has a poor understanding of what an “invasive alien species” is. Many
invasive alien plant species such as the Australian pine
(Casuarina equestifolia) are long established in the
country, and considered part of the natural landscape. The
challenge of just being able to monitor the ecology of an
archipelago with 30 different island population centers is
substantial. The management of invasive species requires
a coordinated response between many different islands
with varying capacities. Like many environmental issues,
the threat invasive species is the consequence of inadequate
infrastructure, regulation and planning both for the fastgrowing population on New Providence, and the emerging
Family Island communities.
MANAGEMENT OPTIONS
The consensus at the fall 2008 Lionfish Stakeholders’
Meeting for The Bahamas was that there is an urgency to
develop and implement a National Lionfish Response Plan
(NLRP) in lieu of the precautionary principle. The mission
of a NRLP should be fourfold:
i) To maintain the distinctiveness and diversity of
Bahamian marine communities,
ii) To protect commercially important fisheries,
iii) To safeguard public health, and
iv) To reduce the growth and spread of lionfish
populations.
Ideally, the ultimate goal of a NLRP would be the
eradication of lionfish and prevention of a possible reintroduction of the species. Realistically, the goal would
be to reduce lionfish population growth and spread in
targeted areas such as Marine Protected Areas and beaches
frequently used by the general public. For any invasive
species management plan to be effective it must be
informed by sound science, objective data and the financial
reality of those charged with its implementation. Table 1
presents a summary of potential lionfish management
options suggested at the fall 2008 Lionfish Stakeholders’
Meeting. Each option was subjectively ranked by meeting
attendees. However, we emphasize that none of these
managements options can be implemented or seriously
considered without prior rigorous scientific research and
economic analyses of their viability.
STAKEHOLDER GROUPS AND
POTENTIAL PARTNERSHIPS
One of the most remarkable aspects of the Indo-Pacific
lionfish invasion is the wide range of stakeholders and
resource user groups that are potentially impacted by its
establishment and spread.
The lionfish invasion is
unprecedented for The Bahamas in terms of the scope of its
potential impacts across many different marine and coastal
resource user groups. Because of the large number of
private businesses and individuals included in the stake-
holder groups (Table 2), there are likely to be a number of
privately funded initiatives independent of any government
action. The geographic extent and rapid rate of spread of
the invasion coupled with the large and diverse body of
stakeholders demands an equally rapid and coordinated
planning and implementation process on the part of the
government.
One challenge in bringing together all the stakeholder
groups is building a common body of information or
knowledge, the “baseline” information on the lionfish and
its impacts. Science-based management requires a close
coordination between research and management decisionmaking. Emerging information from researchers emphasizes the challenges of management, and a NLRP requires
engagement of all stakeholder groups for years to
come. There are four “functional” components of the
plan:
i) Accessing and the management of adequate
funding,
ii) Lionfish research, information management and
monitoring,
iii) Outreach and education on lionfish control and
prevention efforts, and
iv) Revision and further development of invasion
policy and regulations.
No component is more important than another, all
components work in concert to implement a management
plan.
The NLRP will require new and innovative approaches
to resource management and regulation. In addition, new
international partnerships should to be forged to meet the
scientific research and information needs. The level of
information exchange and sharing of resources would
likely change dramatically when foreign researchers are
engaged to help address a national research agenda. Table
3 outlines some basic research areas, with an assessment as
to whether sufficient progress could be made with capacity
within the country. Research done collaboratively with
more student exchanges and formal training will serve to
build the long-term resource management capacity.
Perhaps the biggest challenge is developing a national data
management system for natural resource and biological
diversity management. The NLRP can be an initial step
towards larger environmental management goals by
providing the necessary public focus, funding and international collaborations.
CONCLUSION
The writing and implementation of a National Lionfish
Response Plan for The Bahamas is a culmination of many
long term efforts to implement broader “ecosystem
management” of natural resources and build an integrated
coastal zone management plan. The invasion of the IndoPacific lionfish is in many ways symptomatic of the overall
challenges for Small Island Developing States (SIDS) to
Sullivan-Sealey, K. et al.
address environmental threats that cut across government
agencies and require additional scientific and technical
capacity beyond the borders of the country. If The
Bahamas can successfully plan and implement actions to
control lionfish populations, it will be an important step
forward in inter-departmental cooperation and privatepublic partnerships. The management actions require both
regulatory and non-regulatory approaches to the invasion
control, with a critical role assigned to public education
and outreach programmes.
From the initial stakeholders meetings, it is clear that
the overall level of “science literacy” in the country will
come into play in building public support for management
goals and actions. Fishers, diver operators and Family
Island fisheries officers all have opinions on the movement, behaviour and ecology of lionfish based on observations and conversations, yet there is very little information
about the science of fish population dynamics and invasion
ecology. Often, this lack of “science literacy” leads to
unrealistic expectations for management goals, and
underestimates of the cost for management actions.
Workshops and seminars are critical to bring accurate,
factual and up-to-date information to a broad audience of
stakeholders, and focus an educational outreach campaign
on the broader science of invasive species. Marine alien
species invasions are a threat to The Bahamian environment not only from the aquarium trade, but also through
ballast water introductions associated with port operations.
The focus of any outreach and public education campaign
should be on both the specific stewardship actions required
for individuals as well as the importance of a broader
knowledge of the local environment and its ecology.
Clearly, the areas in the above tables where there was a
lack of stakeholder consensus were due largely to a lack of
information or knowledge by some stakeholder groups.
Managing expectations and getting the best lionfish
population control results for the resources invested will
depend on an aggressive and innovative outreach campaign.
This need for accurate and current information on the
biology and ecology of lionfish in Bahamian waters may
present some new challenges in partnerships between the
Government of The Bahamas and foreign research permit
holders. The control of lionfish populations will be a
management issue for likely decades to come. The
Bahamas would wish to both increase its internal capacity
to study, analyze and archive information on invasive
species, as well as build stronger and more collaborative
relationships with foreign researchers working in the
country. Special conditions or arrangements may be
necessary on research permit holders to insure that results
are presented back to the NLRP management team in a
timely manner, and that datasets are available for compiling national archives and databases. A clear focus in
management planning can help refine the requests for
funding and research priorities. The Bahamas will play an
GCFI:61 (2009)
Page 407
important role as a regional leader in marine invasive
species research and management as the NLRP continues
to develop.
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Golani, D. and O. Sonin. 1992. New records of the Red Sea fishes,
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Guerrero, K.A. and A.L. Franco. 2008. First record of the Indo-Pacific
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Hamner, R.M, D.W. Freshwater, and P.E. Whitfield. 2007. Mitochondrial cytochrome b analysis reveals two invasive lionfish species
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Kochzius, M., Solier, R., Khalaf, M.A., and D. Bloom. 2003. Molecular
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(Scorpaenidae, Pteroinae) based on mitochondrial DNA sequence.
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world's worst invasive alien species: a selection from the global
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Maljkovic, A., T.E. Van Leeuwen, and S.N. Cove. 2008. Predation on
the invasive red lionfish, Pterois volitans (Pisces: Scorpaenidae), by
native groupers in the Bahamas. Coral Reefs. Online:28 March
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McKinney, M. and J. Lockwood. 1999. Biotic homogenization: a few
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Barrier Reef and Coral Sea. University of Hawaii Press, Honolulu,
Hawaii USA.
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Sax, D.V. and S.D. Gaines. 2008. Species invasions and extinction: the
future of native biodiversity on islands. Proceedings of the National
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Copeia 1986:686-690.
Semmens, B.X., Buhle, E.R., Salomon, A.K. and C.V. PattengillSemmens. 2004. A hotspot of non-native marine fishes: evidence
of the aquarium trade as an invasion pathway. Marine Ecology
Progress Series 266:239-244.
Snyder, D. and G. Burgess. 2006. The Indo-Pacific red lionfish, Pterois
volitans (Pisces: Scorpaenidae), new to Bahamian ichthyofauna.
Coral Reefs. Online:25 October 2006.
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Hawaii: biological effects and opportunities for ecological research.
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Whitfield, P.E., T. Gardner, S.P. Vives, M.R. Gilligan, W.R. Courtenay,
G.C. Ray, and J.A. Hare. 2002. Biological invasion of the IndoPacific lionfish Pterois volitans along the Atlantic coast of North
America. Marine Ecology Progress Series 235:289-297.
Whitfield, P.E., J.A. Hare, A.W. David, S.L. Harter, R.C. Munoz, and
Atlantic. Biological Invasions 9:53-64
Consumption Potential of Invasive Lionfish (Pterois volitans) On Caribbean Coral Reefs
STEPHANIE GREEN and ISABELLE CÔTE
Department of Biological Sciences Simon Fraser University 8888 University Drive Burnaby, BC V5A1S6 Canada
EXTENDED ABSTRACT
First reported from Atlantic coral reefs in 2004, Indo-Pacific lionfish (Pterois volitans) have spread rapidly around the
Caribbean basin (Freshwater et al. 2009, Morris et al. 2009) and are now one of the most abundant predators of their size on
invaded Bahamian coral reefs (REEF 2009). Following this rapid range expansion, temporal data from the Reef Environmental Education Foundation reveal exponential increases in the local abundance of lionfish in several regions (REEF
2009). There is great concern about the ecological and economic impacts of dense lionfish populations on Caribbean coral
reef systems (Green and Cote 2009). Analyses of lionfish stomach contents from the Bahamian archipelago reveal that
lionfish prey on over 50 species of native Caribbean reef fish (Green Unpublished data, Morris and Akins 2009) and studies
on experimental reefs reveal that a single lionfish can reduce the recruitment of native Caribbean fishes by 80% through
predation (Albins and Hixon 2008). However, to understand the cumulative impact of lionfish on Caribbean coral reef fish
communities, and set targets for lionfish population management, we must link the magnitude and rate of lionfish prey
consumption to the production of their prey on invaded reefs.
To meet these objectives, we synthesized data on invasive lionfish population parameters, bioenergetics and diet to
created a probabilistic model (Vose 2008) of lionfish prey consumption on invaded reefs and then compared these consumption rates with estimates of production of their fish prey. To parameterize our model, we collected data on prey-sized
fish density, diversity and size distribution, and lionfish density, size distribution and diet collected from 13 invaded coral
reef sites off the southwest coast of New Providence, Bahamas, from May through to September 2008.
Our analyses reveal that invasive lionfish populations far exceed sustainable levels on the majority of the coral reefs we
studied. Using our model to quantify the density of lionfish that could be sustained by their prey populations at each site
revealed that most reefs require significant reduction in lionfish abundance and that ‘sustainable’ lionfish densities depend
not only prey production rates, but on the size distribution of lionfish at each site.
The main tool being considered by managers to control invasive lionfish population is manual removal by divers.
However, the cryptic nature of lionfish may introduce challenges in effectively detecting their presence across the reefscape.
In particular, there may be a bias towards detecting and removing only the largest size classes of lionfish inhabiting a reef
area. We used our model to evaluate the effect of this potential size bias on the ability of removal efforts to keep lionfish
abundances at sustainable levels. Using data from a study site harbouring a high density of large lionfish (density: 344 ±
164 lionfish/ha (mean ± SD); body mass: 296 ± 211 g/lionfish), we simulated our model for three removal scenarios. We
varied the smallest size class of lionfish detected and removed from 15cm, to 20cm, and 25 cm total length, respectively.
To determine how the minimum size of lionfish removed affects the sustainability of the consumption by the remaining
lionfish, we simulated the difference between annual prey consumption by lionfish and the annual production of their prey
base for each scenario. Results were only considered sustainable if the removal effort reduced annual prey consumption to
a level equal to or less than annual prey production at the site.
We found that targeting all individuals greater than 15cm, 20cm and 25cm total length would remove 89%, 87%, and
79% of the population, respectively. However, there were substantial differences in the sustainability of each removal
scenario (Figure 1). Our analyses revealed in two of the three scenarios, both targeting lionfish greater than 20cm and 25cm
total length for removal, lionfish prey consumption was not reduced to sustainable levels (i.e. net annual biomass production of their prey was less than zero). However, our model revealed that removing all individuals greater than 15cm total
length from the site reduced the consumption capacity of the remaining population to levels that were far surpassed by the
production of their prey, and was therefore sustainable. These results indicate that potential size selectivity in removal
efforts can significantly affect the success of lionfish population management efforts. Our model provides a method by
which regional resource managers can assess the risk of local reef fish populations to lionfish predation, and set clear targets
for control and management efforts on invaded Caribbean reefs.
KEYWORDS: Lionfish, invasive species, predator, fish production
Tasas de Consumo Potenciales del Pez León Invasor (Pterois volitans)
en los Arrecifes de Coral del Caribe
Desde el año 2004 cuando se registro la primera aparición del pez león (Pterois volitans) especie invasiva procedente del IndoPacifico en los arrecifes de coral del Atlantico occidental, el pez león se ha extendido rápidamente en la cuenca del Caribe. Para entender
el impacto de la depredación del pez león sobre la comunidad de peces en las Bahamas y para predecir los impactos potenciales sobre el
mar del Caribe, creamos un modelo probabilístico de la tasa de consumo de presas por el pez león en 13 arrecifes invadidos en las
Proceedings of the 62nd Gulf and Caribbean Fisheries Institute
November 2 - 6, 2009 Cumana, Venezuela
Green, S. and I. Côte GCFI:62 (2010)
Page 359
Bahamas y comparamos estos valores con las estimaciones de las tasas de producción sus peces presa. Nuestro análisis revela que el
pez león tiene la capacidad de consumir presas en los arrecifes con una tasa mayor a la que las que las poblaciones de peces de arrecife
pueden recuperarse. Para fijar objetivos para la gestión, usamos el modelo para cuantificar las densidades de peces león que podrían
mantener las tasas de producción de peces presa disponible en cada zona. Las densidades sostenibles estimadas no dependen solo de la
tasa de producción de las presas sino también del tamaño de las distribuciones de los peces león que habitan en cada zona. Nuestro
modelo proporciona una herramienta a los gestores de los recursos regionales para evaluar el riesgo de los peces león sobre las
poblaciones de peces de arrecife locales y para establecer objetivos de control y esfuerzos de gestión.
PALABRAS CLAVES: Especie invasiva, Pterois volitans, producción sus peces
Consommation Potentielle du Rascasse Envahissant (Pterois volitans)
Sur les Récifs De Corail Antillais
D'abord annoncé des récifs de corail de l'Atlantique en 2004, le rascasse Indo-Pacifique (Pterois volitans) s'est étendu rapidement
autour de la cuvette antillaise et est maintenant un des prédateurs les plus abondants de leur taille sur les récifs de corail envahis des
Bahamas. Pour comprendre l'impact de la prédation par le rascasse sur les communautés de poisson natifs aux Bahamas et prédire leurs
impacts potentiels au large des Caraïbes, nous avons synthétisé des données sur les paramètres démographiques de rascasse envahissant, la bio-énergétique et l'alimentation pour créer un modèle probabilistique de consommation de proie de rascasse sur les récifs
envahis et avons comparé ces taux de consommation aux estimations de production de leur poisson de proie. Pour paramétrer notre
modèle, nous avons recueilli des données sur la densité du poisson de la taille de poisson proie, la diversité et la distribution de taille et
la densité rascasse, la distribution de taille et l'alimentation recueilli de 13 sites de récif de corail envahis de la côte du sud-ouest de la
Nouvelle Providence, Bahamas, de mai à septembre 2008. Nos analyses révèlent que le rascasse a le potentiel d'enlever la proie de
nombreux récifs à un taux bien plus élevé que les populations de poissons de récif peuvent se reconstituer. Nous avons utilisé aussi
notre modèle pour quantifier la densité du rascasse qui pourrait être soutenue par la production de poissons de proie disponibles sur
chaque site. Les densités de rascasse 'durable' dépend non seulement des taux de production de poisson, mais de la distribution de taille
de rascasse sur chaque site. Notre modèle fournit une méthode par laquelle les directeurs de ressource régionaux peuvent évaluer le
risque de la population locale de poissons de récif à la prédation du rascasse et fixent des objectifs clairs du point de vue de la densité
du rascasse pour le contrôle et les efforts de gestion sur les récifs antillais envahis.
MOTS CLÉS: Rascasse, espèces envahissantes, prédateur, production de poissons
LITERATURE CITED
Figure 1. Predicted net prey production (mean ± non-parametric bootstrapped 95% confidence intervals) under
three removal scenarios: Removal of all lionfish greater than 15cm TL (a), 20cm TL (b), and 25cm TL (c).
Ablins, M.A. and M.A. Hixon. 2008. Invasive Indo-Pacific lionfish Pterois volitans reduce recruitment of Atlantic coral-reef fishes. Marine Ecology
Progress Series 367:233-238.
Fishelson, L. 1997. Experiments and observations on food consumption, growth and starvation in Dendrochirus brachypterus and Pterois volitans
(Pteroinae, Scorpaenidae). Environmental Biology of Fishes 50(4):391-403.
Freshwater, D.W. et al. 2009. Mitochondrial control region sequence analyses indicate dispersal from the US East Coast as the source of the invasive
Indo-Pacific lionfish Pterois volitans in the Bahamas. Marine Biology 156(6):1213-1221.
Morris, J.A., J.L. Akins, A. Barse, D. Cerino, D.W. Freshwater, S.J. Green, R.C. Muñoz, C. Paris, and P.E. Whitfield. 2009. The biology and ecology of
invasive lionfish. Proceedings of the Gulf and Caribbean Fisheries Institute 61:409-414.
REEF. 2009. Reef Environmental Education Foundation Volunteer Survey Project Database. www.reef.org. (01/11/2008).
PRESENTATIONS
Reconstructing the Western Atlantic
lionfish Invasion
R. Betancur-R.
A. Acero P.
G. Ortí
A. Hines
A. Wilbur
W. Freshwater
Pterois volitans
Pterois miles
Pterois volitans
Pterois miles
Introduced into Florida!
Popular ornamental fishes
~80K lionfish specimens
imported into Florida in 40
years
First non-native marine fishes
established across the W. Atlantic
Non-native lionfish reports as of June 2010
Lionfish progression
Schofield (2009): Aquat. Invas.
Lionfish progression
Schofield (2009): Aquat. Invas.
Lionfish as model system
Research aims/questions
1- Establish the geographic extent of invasive P.
volitans and P. miles
Lionfish reports: P. volitans? P. miles?
Difficult morphological differentiation!
2- Reduction or increment in genetic diversity?
Reduced or increased levels of genetic diversity?
•Founder event
Native population
Invasive population
bottleneck
•Founder event
Native population
Invasive population
bottleneck
REDUCED DIVERSITY!
•Multiple introductions from different populations
Invasive population
Native pop. A
Native pop. C
Native pop. B
•Multiple introductions from different populations
Invasive population
Native pop. A
Native pop. B
INCREASED DIVERSITY!
Native pop. C
2/3 of aquatic invasions
(Roman & Darling, 2007)
3- Single or multiple independent lionfish
introductions across WA?
Single introduction?
Multiple independent introductions?
3- Single or multiple independent lionfish
introductions across WA?
4- Population structure and insights into
Caribbean connectivity patterns
Lionfish as model system
•Marine biology: insights into Caribbean connectivity
patterns
Cowen et al. (2006): Science
Data
•Mitochondrial d-loop sequences
Native lionfish
1- Pterois volitans vs. P. miles
n
h
P. volitans 734
9
P. miles
1
21
P. miles is restricted to the north; P. volitans is ubiquitous
2- Genetic diversity in P. volitans
n= 734
h= 9
HD= 0.60
n= 70
h= 36
HD= 0.97
Founder eff.
2- Genetic diversity in P. miles
n= 21
h= 1
n= 10
h= 8
Founder eff.
Increased genetic diversity in 2/3 of aquatic invasions
Invasive population
Native pop. A
Native pop. C
Native pop. B
Increased genetic diversity in 2/3 of aquatic invasions
GENETIC PARADOX!
Invasive population
Native pop. A
Native pop. C
Native pop. B
3- Single or multiple introductions of P. volitans?
D-loop haplotype network
No evidence of multiple introductions
Mitochondrial data are
consistent with the
lionfish progression
from sightings
Rapid range expansion
•30K eggs per spawning event
•Spawn as often as every four days
•Release buoyant egg masses with pelagic
larval duration estimated at 25+ days
4- Population structure in P. volitans?
h= 9
h= 4
Secondary founder effect in the Caribbean
4- Population structure in P. volitans?
Secondary founder effect in the Caribbean
Conclusions
1- P. miles is restricted to the US east coast and
Bermuda; P. volitans is ubiquitous and much more
abundant
2- Reduction in genetic diversity in the WA as a
result of a founder effect: genetic paradox
3- Mitochondrial data show no evidence of multiple
independent introductions. Consistent with a
range expansion out of the epicenter (Florida)
4- Two population groups (northern vs. Caribbean)
Directions
•Obtain samples from the eastern Caribbean and
collect microsatellite data:
•
Fine-scale resolution into population
structure and Caribbean connectivity
•
Species boundaries
Acknowledgements
•Lab members at the Ortí, Universidad Nacional
de Colombia sede Caribe, and UNCW labs
Effects of invasive Pacific red lionfish on
Bahamian coral-reef fish communities:
preliminary results from a large-scale,
long-term experiment
Mark A. Albins
Department of Zoology
Hixon Lab Lionfish Research
1. Effects on native coralreef communities
– Experimental
– Observational
2. Ecological release and
biotic resistance
– Recruitment, movement,
growth and survival
– Time budgets, predation
rates etc.
– Community interactions
•
•
•
Competition
Predation
Parasitism
Effects on Atlantic
Coral-Reef Communities
?
Experimental Patch Reefs
1m
200 m
Predation Experiment
Coney effect
35% reduction
p=0.105
Lionfish effect
Combined effect
85% reduction
88% reduction
p=0.001
p=0.001
Albins (in review) Biological Invasions
Effects on Atlantic
?
Small scale
Large
Short term
Long
term
?
?
Large-scale, Long-term Experiment
•
•
•
•
10 large reefs (2500-5000 m2)
Paired based on habitat similarity
Baseline surveys
Low and high lionfish density
– Low ~0-100/hectare
– High ~500-600/hectare
• Quarterly surveys
• Quarterly treatment maintenance
Large-scale, Long-term Experiment
Each Site:
2 Plots – 100 m2 / each
4 Strips – 50 m2 / each
Change in Density (fish/m2 ± SEM)
All Species
Combined
Low Lionfish Density
High Lionfish Density
Change in Density (fish/m2 ± SEM)
All Species
Combined
Low Lionfish Density
High Lionfish Density
82%
reduction
52%
reduction
Time (quarters)
Effects on Atlantic
?
?
?
Large scale
Long term
?
Funding and Support
Provided by:
?
Special Thanks
Academic Committee: Mark
Hixon, Bruce Menge, Lisa
Madsen, Selina Heppel,
George Bohlert
Colleagues: Chris Stallings,
Darren Johnson, Mark
Christie, Tim Pusack, Tye
Kindinger, Kurt Ingeman,
Kimberly Page-Albins
Post Graduate Assistants:
Gabe Scheer, Julia Lawson
Undergraduate Assistants:
Robbie Lamb, Megan Cook,
Emily Pickering, Wendel
Raymond, Alyssa Alder
Perry Institute Staff: Brenda
Gadd, Meredith Newman,
Eric Lamarre
The Impacts of the Indo-Pacific Lionfish (P.
volitans and P. miles) on Fish Assemblages in
Near Shore Benthic Reefal Habitats of the
Central and Southern Bahamas
Nicholas Bernal 1, Alexio Brown2
and Kathleen Sullivan Sealey1, 2
1Department
of Biology, University of Miami, Coral
Gables, Florida 33124
2School of Natural Sciences, College of The
Bahamas, Nassau, The Bahamas.
Coastal Ecology of The Bahamas
•
•
•
•
Completing 10 years of surveys of major islands in
the Bahamian archipelago
Look at near shore marine environments, including
patch reefs, seagrass beds, hard-bottom and
mangrove creeks
Examine patterns in fish species assemblages with
season, degree of coastal alteration and water
quality
Lionfish have been encountered since 2006 surveys
2006
2007
2003
2002
2010
2005
2004
2008
2009
RESEARCH QUESTION:
How have lionfish impacted fish community
level diversity in near shore habitats over
time?
Great Exuma
Great Inagua
Habitat Classifications:
Near Shore Fringing Reef
Near Shore Patch Reef
Channel Reef
Hardbar
Some sites can be 1m> at low tide!
Roving Diver Surveys




Average survey is 45 minutes per snorkeler.
All sites used in the study have aggregate
bottom time of over 180 minutes.
Sites restricted within 200m of shore (usually
less than a hectare)
Allows for rapid assessment of fish
assemblages in habitats around an island with
multiple observers.
Diversity Indices Summary Table
Average Shannon (H') Diveristy by Habitat and Year
Year
Hardbar
Channel
Patch
Fringing
1996
N/A
4.371
4.093
N/A
2002
N/A
4.292
3.977
N/A
2010
3.942
4.237
3.962
3.978
Average Berger Parker (d) Diversity by Habitat and Year
Year
Hardbar
Channel
Patch
Fringing
1996
N/A
16.854
13.64725
N/A
2002
N/A
15.739
12.51575
N/A
2010
11.917
15.132
12.249
11.865
11/1/10
Mean Shannon
Diversity
Mean Shannon Diversity From 1996-2010
5
4.8
4.6
4.4
4.2
4
3.8
3.6
3.4
3.2
3
1996
2002
2010
Year
Mean Berger Parker
Diversity
Mean Berger Parker Diversity From 1996-2010
18
16
14
12
10
8
6
4
2
0
1996
11/1/10
2002
Year
2010
Results:
Bray Curtis Similarity Dendogram (Years)
EX-TS
EX-HB EX-RC
EX-SR
EX-FC
Results:
PCA Diagram
Inagua Sites
EX-FC
EX-SR
Conclusions:
•
•
•
•
•
Observable shifts in fish assemblages dependant on
habitat type and island.
Shift in five sites on Great Inagua attributable to
differences in parrotfish abundance.
Fowl Cay (EX-FC) changes related to temporal decline
in Nassau grouper abundances.
Great Exuma patch reefs shifting due to change in
abundance of: Silversides, Grunts, Damselfish, and
Chromis species
Hardbar habitats on both islands show shift due to
increase in lionfish abundance.
Lionfish are not successful
everywhere – what limits their
success and abundance?
What are the preferential habitats for lionfish?
Thank You!
University of Miami
Department of Biology
Invasive red lionfish in shallow habitats
of the Turks & Caicos Islands
JAB Claydon,1 J Batchasingh,2 MC Calosso,1 SE Jacob,1 K Lockhart 2
1
The School for Field Studies, Center for Marine Resource Studies
South Caicos, Turks & Caicos Islands, B.W.I.
2
Department of Environment & Coastal Resources
Turks & Caicos Islands, B.W.I
Introduction
Regionally, emphasis on deep habitats (SCUBA)
Shallow habitats overlooked
Mangrove & seagrass nursery
Critical habitats for fisheries species
TURKS & CAICOS ISLANDS
Data collection
Opportunistic sampling
2007 to 2008
Systematic sampling
2009 to 2010
Tournament specimens
2009 to 2010
Opportunistic sampling 2007 - 2008
Shallow
Deep
15
Sightings month-1
n
28
1
TL (cm)
10.2
27.7
N
10
10 km at 16 m
6 km at 24 m
5
0
A
S
O
2007
N
D
J
F
M
A
2008
M
J
J
Systematic sampling 2009 - 2010
Mangrove = 4
Seagrass = 60
Total lionfish = 107
Reef = 43
3m
Systematic sampling 2009 - 2010
CPUE
0.4
0.3
Chi2 test
p < 0.05
0.2
0.1
0
Seagrass
Reef
3m
Seagrass
> 90% in blowouts
Deep
n = 126
n = 809
15
Shallow
5
10
T-test
p < 0.05
0
Density (%)
20
Size vs. depth
0
10
SL (cm)
20
30
3m
Invasion timeline
2007
2008
2009
2010
N
1 km
Invasion timeline
2007
2008
2009
2010
N
1 km
Invasion timeline
2007
2008
2009
2010
N
1 km
Invasion timeline
2007
2008
2009
2010
N
1 km
Invasion timeline
2007
2008
2009
2010
N
1 km
Invasion timeline
2007
2008
2009
2010
N
Ontogenetic migration ?
1 km
Conclusions
Highest densities of lionfish in seagrass
Larger lionfish deeper
Delay in invasion of deeper habitats ?
Ontogenetic shift in habitat use ?
Future directions
Ontogenetic habitat shifts
Tagging & telemetry
Deep & shallow sampling
Effect on Nassau grouper
Competition & predation
Population density
Acknowledgments
The School for Field Studies for financial and logistical support
Staff and students of SFS Center for Marine Resource Studies
In particular: J Clauson, C Cormier, F Hoogakker, B Hooper,
K Ketch, C Lamendola, D Myers, D Short, L Sperry
Turks & Caicos Islands fishermen and dive operators
jclaydon@fieldstudies.org
mcalosso@fieldstudies.org
Why Are Lionfishes (Pterois, Scorpaenidae) So Rare
In Their Native Ranges?
Terry J. Donaldson, David Benavente and Roxie Diaz
University of Guam Marine Laboratory
UOG Station
Mangilao, Guam 96923 USA
email: donaldsn@uguam.uog.edu
What is Rarity?
• Species with low abundance
• Species with small geographic ranges
• Species that may be found only in a few
specialized habitats
• These components are not mutually exclusive
A Fourth Definition
• Species with wide geographical ranges but low
adult abundances at any given locality
Cheilinus undulatus
(Labridae)
Photos: J.E. Randall
Bolbometopon muricatum
Labridae: Scarinae
What Causes Species to be Rare?
• Relationship between body size and range size
• Relationship between body size and abundance
• Environmental adaptations resulting from range
size variability at higher latitudes
• Habitat requirements and resource availability
• Poor ability to exploit marginal habitats or other
resources
• Poor mobility
• Low reproductive effort
• Poor recruitment
After Gaston (1994); Jones, Munday and Caley (2002)
The Unusual Suspects:
Lionfishes (Pterois, Scoraenidae)
Pterois antennata
Photographs: J.E. Randall
Pterois radiata
Pterois volitans
Why Are Scorpionfishes So Rare?
Photographs: J.E. Randall and R.E. Myers
Tropical and Subtropical Scorpionfishes
• Some have a high degree of habitat specificity
• Strong site fidelity because of poor mobility
• Behavior
• Ambush or hunting predators
• Long larval life but poor recruitment?
• Prey of larger predators limits abundance of adults and
juveniles?
• Sampling artifacts: cryptic and/or nocturnal
Visual surveys at selected Pacific localities (n = 632 transects)
Transect Type and Number
Locality
Guam
Northern Mariana Islands (limestone)
Northern Mariana Islands (volcanic)
Palau
Southwest Palau Islands
Yap-Ulithi, Micronesia
Chuuk, Micronesia
Kosrae, Micronesia
Kwajalein, Marshall Is.
Majuro, Marshall Is.
American Samoa
Sta. Ysabella, Solomon Is.
Visayas, Philippines
Total
Timed Swim GPS Timed
63
43
5
0
65
21
0
11
6
3
0
22
12
251
7
0
0
0
0
0
0
0
0
0
0
0
3
10
Belt
157
4
0
11
0
8
55
0
7
7
122
0
0
371
Number of transects out of a total of 632 transects where lionfishes
were observed. Pant = Pterois antennata; Prad = P. radiata;
Pvol = P. volitans.
Transect type
Pant
Prad
Timed GPS
0
0
0
Timed swim
3
5
23
Belt
3
0
3
Total
6
5
26
0.8%
4.1%
Percent
1.9%
Pvol
What Causes Species to be Rare?
• Relationship between body size and range size
• Relationship between body size and abundance
• Environmental adaptations resulting from range
size variability at higher latitudes
• Habitat requirements and resource availability
• Poor ability to exploit marginal habitats or other
resources
• Poor mobility
• Low reproductive effort
• Poor recruitment
Body Size, Range Size and Abundance
• Species with wide geographical ranges but low
adult abundances at any given locality
Cheilinus undulatus
Photos: J.E. Randall
Bolbometopon muricatum
Habitat
Pterois antennata
Pterios volitans
Pterois radiata
Pterois volitans
Resource Use and Behavior
• Lionfishes are social
• Shelter together
• Hunt cooperatively
Resource Use: Being a Predator
• Adults are piscivorous but also feed upon crustaceans
• Nocturnal predators but also feeding during daylight
and crepuscular periods (dusk and dawn “switchover”)
• Prey limited?
Doubtful.
Long Larval Life and Poor Recruitment
• Morphological and life history specializations of
larvae facilitate long larval periods and greater
dispersal
• Wide geographic ranges
• Why poor recruitment?
Many larvae lost in “vastness” of the IndoWest Pacific?
Predation upon larvae, post-larvae and small
juveniles promotes low abundance locally?
Prey of Larger Predators?
• Largely anecdotal information or small sample
sizes
• Scorpaenids found in stomachs of groupers
(some groupers not much larger than their prey)
• Moray eels will prey opportunistically upon adults
• But, how much larger? What preys upon postlarval juveniles?
Poor Mobility and High Site Fidelity
Scorpaenopsis macrochir
Pterois volitans
Pterois antennata
Rarity, “crypticity”, nocturnal behavior and
sampling error
Turning the Question Around:
Why Aren’t Lionfishes Rare in the
Western Atlantic?
Some Possible Explanations (But Not All)
• Lack of effective predators upon adults
• A surfeit of suitable habitat (= disturbed habitat)
• The geography and physics of larval distribution
• Lack of effective predators upon post-larval juveniles
• Competitive release as invasive species
• Altered states: shifts in life history strategies that promote
rapid growth, early maturity and reproduction
Acknowledgments
• Jack Randall- B.P. Bishop Museum (photographs)
• Robert Myers- Coral Graphics (photographs)
• Surveys funded or supported directly or indirectly by:
NOAA, U.S. Fish and Wildlife Service, U.S. National
Park Service, U.S. Navy, The Nature Conservancy, and
the University of Guam Marine Laboratory
Red Lionfish control strategies in the
Caribbean UK Overseas Territories
(Cayman Islands, British Virgin
Islands and Turks and Caicos Island).
Bradley Johnson1, Shannon Gore2, Kathy Lockhart3
1
Department of Environment, Cayman Islands, bradley.johnson@gov.ky
2 Conservation & Fisheries Department, British Virgin Islands, sgore@gov.vg
3 Department of Environment and Coastal Resources, Turks and Caicos
Islands, kglockhart@hotmail.com
Nov ‘07
Mar ‘10
Feb ‘08
Nov ‘07
Mar ‘10
Feb ‘08
Joint Nature Conservation Committee (JNCC)
Funding Nov ‘09
British Virgin Islands (B.V.I.)
Marker System, #’s low
B.V.I.
Marker System, #’s low
Wine Cork Request
B.V.I.
Public Awareness
– Get initial sighting reported
– Only trained responders
– No licensing needed
• TCI MPA and CI required
B.V.I.
Training
– REEF
• TCI & CI
– CORE - Caribbean Oceanic
Restoration and Education
B.V.I.
School/Business Visits – TCI & CI also
B.V.I.
Capture Gear
– Donated 40 sets
– Purchased with
JNCC funds
TCI & CI additional
Image: reef.org
Turks and Caicos Islands (T.C.I.)
Tasting Events
Supermarkets, public events etc.
T.C.I.
Restaurant Initiative
Promote lionfish as a food fish
T.C.I.
Convert Commercial Fishermen
Safe to eat, care when handling
T.C.I.
Year-long Tournament
– Grand Prize $3,000 for
3,000 fish
Cayman Islands
C.I.
Licensing courses held regularly
C.I.
Licensing courses held regularly
Over 600 culler licensed since March 2009
C.I.
License conditions:
• Only for taking lionfish
• Now allowed to:
– Take while on SCUBA
– Take from MPA
– Wear gloves
– Take fish under 8”
• All fish killed
• Data sent to DoE
• No spearing devices or
noxious substances
Photo: Jason Washington
C.I.
Resident participation
• Enjoyment
• Recreational activity
• Environmental awareness
C.I.
Dive Operators/Companies
• Involved from the start, even without proper gear
C.I.
C.I.
C.I.
Problems encountered by dive companies
• More & larger lionfish
– time/depth constraints
– Smaller gear not practical anymore
• Nets bulky and cumbersome
• Responsibility to guests first
• Requested pole spears
C.I.
Conditions for Dive Companies to spear:
• Only licensed cullers
• Only DoE issued spears
• Only while leading dives
• Spears tagged & licensed to company &
boat
• Companies provide list of employed
licensed cullers
Can’t import or manufacture spear guns or parts
C.I.
Expand spearing program
• DoE supervised culls
• Spears remain with DoE
• Import/manufacture
more spears
• Assign Fisheries Officers
to lead culls
C.I.
Tournaments
• Private companies organize
• Must follow DoE
Tournament Guidelines
• DoE acts in advisory and
support role
– Measuring at weight station
– Training chefs on handling
fish
• Less time required
• Enforcement presence
Facebook
• Turks and Caicos Lion Fish Public
Awareness Campaign
• BVI lionfish
• DoE Lionfish Culling Group
Thank You
Photo: Jason Washington
Implementation of the
Management Plan
For the Control of the Lionfish In Puerto Rico
• 4 Years
Carmen Rosa Valentín
• 15 minutes
Luis Raúl Saez
Joel Meléndez
Geovanny Negrón
Everything Start
• July 25, 2007
– Caribbean
Photo: Jaime Matos
• Vieques
– November 14, 2008
Everything Start
• Fuete y Verguilla en
Octubre 2008 (Volumen
2, Número 4)
– Sr. Daniel Matos
– “Los beneficios de las
Vedas para los Pescadores
comerciales”/ “The benefits
of the fishing seasons for
commercial fishermen”
Las especies marinas pueden ser sobre
explotadas y pueden desaparecer /
Marine species can be overfished and can
disappear
• Fishing gear and
methods / regulations
Everything Start
• GIS – Overlap
– Ecosystems
– The People (Divers,
Fisherman's, Water
Sport, Public)
– Lionfish
• Not in my Backyard
cartoonstock.com
Público
Marinas
Buzos
Arrecife
Deportes Acu
Manglar
Protegemos Ambiente
Everything Start
• March 5, 2010
– William C. Coles,
Ph.D.
USVI, Chief Environmental
Education
Aquatic Education
Coordinator
Division of Fish and
Wildlife
45 Mars Hill
Frederiksted, VI 00840
– 340-772-1955
• Train the Trainers
Puerto Rico
ACTIONS / STRATEGIES
Some Areas Low Visibility –
Season
Habitats
Seagrass (Dra. Roberson)
Mangrove (Pedro Padilla)
Reef (Pedro Padilla)
Natural & Artificial Structures
Jaime Matos
People Concentration & Natural
Reserves
Organize the Program:
Cooperation Levels
Operaciones
Supervisor de Operaciones
Responsable de Apoyo
Administración
Gerencia
Apoyo
Logística de Campo
Seguridad y Protección
Emergencias
Pescadores
Técnicos de Respuesta Rápida
Estudiantes
Médicos
Restaurantes
Operaciones
Pescadores
Gobierno
Comunidades
Técnicos de Respuesta Rápida
Estudiantes
Médicos
Restaurantes
Logística de Campo
Seguridad y Protección
Emergencias
•
Work
–
Público
Fallow up
•
•
How are you?
What you need?
–
–
Provide what they need
when things are not working
–
Keepers
•
Act
Rosa Valentin Vision / Luis Saez
Marinas
Buzos
Arrecife
Deportes Acu
Manglar
Protegemos Ambiente
• Part 1 - Seminar - General Information
• Part 2 - Workshop – How to make
Ornamental Fishing tools
• Part 3 – Water Training – How to Used
• DATABASE
• Lionfish Credit
Program
• Laws & Regulation
• Tools for report
– UMET
• Support
– Pure Adventure
– Fideicomiso
Conservación
– DNAR – Managers NR
• Aguadilla Lionfish
Derby
– January 2011
• $4.00 Credit per
Lionfish +
– $1.00 report
Terry Chevako Bava
www.caribbeanlionfish.org
Restaurants
• Añasco
–
–
•
Dorado
–
–
•
El Fogón de la Curva
$1.00 x Pound
ChoPHouse
9 Lionfish minimum
Luquillo
–
–
La Parrilla
$1.50 x Pound
• DNA Results
– February 30, 2011
• Habitat Description &
Fish Population
– Reefs
• January 1, 2011
– Seagrasses
• January 1, 2011
– Mangrove
• April 30, 2011
• Report - Lion Fish No
take zones
– Universities & NGO,s
– Natural Reserves
– Low People
Concentration
• www.caribbeanlionfish.org
• Mayor Update
– Directory
• Who is doing What?
– Education Materials (Free)
• Power Points / Videos
Publishing Date: January 1, 2011
Important Point 2010 & 2011
• Support to the Communities
– Seminars & Workshops
• New Videos for 2011
– Eat the Lionfish & Protect our
Fish (Kids)
• Fishing Regulations
Amendments
• Lionfish Markets
– Fajardo, Aguadilla, Ponce
• Continue Telling the People
– Never Give UP!!!
www.mascurioso.com
Thanks
Reinaldo Rios
Johnathon Fulop
Caribbean & Eastern Canada Sales Representative
Anna Backe
National Sales Manager, TUSA
Tamara Trinidad
Dr. Robert Mayer
Joseph “Joe” Gulli
Susan Soltero
Noemi Peña
Craig Lilyestrom
Dr. William Coles
Carlos Mendez
Fisherman Associations of Aguadilla
Fisherman Association of Cataño
Marcos Hanke
Capt. Jose Carlos Gonzalez
Brenda Bell Cerezo
Fred Lentz
Questions or Comments?
© Carlos Jiménez
Lionfish in Costa Rica:
threats, actions, and opportunities
Helena Molina Ureña, Ph.D.
Universidad de Costa Rica
63th GCFI, Nov 4, 2010
1
© Carlos Jiménez
Lionfish in Costa Rica:
a love – hate relationship
Helena Molina Ureña, Ph.D.
Universidad de Costa Rica
63th GCFI, Nov 4, 2010
2
Lionfish in Costa Rica
2009
2002
1992
3
Lionfish in Costa Rica: threats
Feeding habits
• Voracious carnivores
üAmbush predators
üPrey unusually large
üCompetitor à predator
• Nighttime? feeders
üCaribbean C.R. à full
guts during the day
© Helena Molina-Ureña
Juvenile lionfish, 2.5 cm TL
Puerto Viejo, October 17, 2009
4
Native habitat
• Coral & rocky reefs
© Carlos Jiménez
Young lionfish
Costa Rica, May 2009
üWarm waters (≥ 15ºC)
üCalm waters à bays, coves
ü0 – 50 m depth
Caribbean habitat
• Seagrass beds
• Mangrove
• Shipwrecks, artificial reefs
Nemo, Vero Beach
FL, Aug 2010
ü0 – ~333 m depth
5
Natural predators
• Large groupers
• Sharks
• Overfished across the
Caribbean
• Other lionfish
Maximum size
• ~ 38 cm TL
• 47.5 cm TL
Grouper caught off Uvita Island
Diario Extra, June 14, 2010
6
Governance issues
Captured lionfish
Puerto Viejo,
February 2010
© Oscar Gutiérrez
• Slow institutional response
• National Park legislation
• No decree declaring
lionfish an invasive
species
• Lack of funding
7
Tropical Eastern
Pacific
• Potential invasion
üUnconfirmed reports
üNorth and Central
Pacific coast of CR
üNot through Panama
Canal
8
Lionfish in Costa Rica: actions
Lionfish
Interinstitutional
Task Force (LITF):
4 axes
1. Prevention à
monitoring
2. Mitigation / Control /
Impact reversal
Time scale Spatial scale
3. Scientific research
• Short term • National
4. Outreach
• Mid run
• Regional
9
LITF: Costa Rica
• C.R. Fisheries Institute
decree
• Permanent monitoring
à early detection
• Outreach & training
workshops à South
Caribbean communities
• Collector licences
• Health practioner training
• Data collection
10
Gear design
1.Costa Rican
version of the
Hawaiian sling
2.Containers
11
Lionfish: monitoring data sheet
10/05/09 Pollito
Danny Mora
9º43'36.3" 82º48'36.2"
2759-xxxx
6
15, 21
R (P / A)
2 peces comiendo juntos / buceo
noche
12
Lionfish: removal data sheet
CR0907-0001
09/05/09
Pollito
9º43'36.3" 82º48'36.2"
12
15
Bichero
13
Lionfish: sampling data sheet
CR0907-0001
02/07/09
Pollito
9º43'36.3"
82º48'36.2"
10
23
14
Lionfish: sampling data sheet
P
Sí
Hígado
4D, 2A, 1P
Sí
Sí
I, D
15
Lionfish in Costa Rica: opportunities
New native predators
• Large groupers
• Large snappers
• Moray eels
ü “Teach” predators to
feed on lionfish
Give reefs a chance
• Learn to live w/ lionfish
• Rearrange food webs
Nassau Grouper,
~90 cm LT
Cave Reef, Bahamas.
Photo: Lyle Gremillion
13 October, 2009
16
Lionfish in Costa Rica: opportunities
GCFI Net
• Dispersion monitoring &
history
• Scientific & technical forum
ü Biology
ü Ecology
ü Genetics
ü Management
• Arrival warnings
• Contacts
Search “Lionfish”
GCFINet
Nov 03, 2010
(230 matches)
17
¡GRACIAS!
© Oscar Gutiérrez
18
INSTITUTO DE ESTUDIOS CARIBEÑOS
CENTRO DE INVESTIGACIONES MARINAS
TROPHIC AND REPRODUCTIVE ASPECTS OF THE LION FISH PTEROIS VOLITANS, IN
SAN SANDRÉS ISLAND, BIOSPHERE RESERVE- SEAFLOWER, COLOMBIAN CARIBBEAN
ASPECTOS TRÓFICOS Y REPRODUCTIVOS DEL PEZ LEÓN PTEROIS VOLITANS, EN SAN
ANDRÉS ISLA, RESERVA DE BIOSFERA - SEAFLOWER, CARIBE COLOMBIANO
Por
Santos-Martínez, Adriana; Acero P., Arturo; Sierra-Rozo, Omar
San Juan de Puerto Rico, noviembre de 2010

El pez león Pterois volitans (Linnaeus, 1758) es una especie
originaria del Indo-Pacífico que ha invadido y colonizado los arrecifes del
Gran Caribe.
 Representa
un alto riesgo ambiental por ser depredadora,
ponzoñosa, con pocos depredadores y euritolerante.
 En Colombia se ha registrado en el Caribe insular y continental; en el
Archipiélago de San Andrés - RB Seaflower, se observó por primera vez
en Providencia en noviembre de 2008.

En 2009 se inició la investigación de ésta especie, con el estudio de
su población para estimar su impacto en Seaflower, con la meta de
contribuir a su manejo.


El archipiélago de San Andrés,
Providencia y Santa Catalina, reserva
de la Biosfera - Seaflower (Unesco,
2000), constituye el Caribe
occidental colombiano

La isla mayor es San Andrés, la
cual tiene un barrera arrecifal en el
costado este.

Se evidencia el deterioro en
los arrecifes de coral en San Andrés
de cerca del 50% y sobrepesca de
poblaciones de importancia
comercial.
 Se cuenta con una zonación de
AMP - Coralina
FASE DE CAMPO
Búsqueda en arrecifes – Buceo y censos visuales
Captura manual con bolsa y congelado
FASE DE LABORATORIO
Procesamiento y mediciones
(identificación, peso y longitud)
Identificación de sexo y análisis de contenido estomacal
SAN ANDRÉS
SITIOS DE
BUCEO
MAYORES
DENSIDADES
PTEROIS VOLITANS
(Guía de Buceo
Pérez et al., 2009)
 Se estima de acuerdo con los buzos y las observaciones que se han
detectado cerca de 500 individuos del pez león en ocho meses (noviembre
2009 a junio de 2010) en los lugares de buceo alrededor de la isla.
 En el primer semestre se observaron por lo general individuos solitarios
o en parejas y actualmente es más frecuente ver grupos entre tres y siete
individuos.
 El pez león es más abundante en los arrecifes coralinos en el presente
semestre que en el anterior; p. ej. en los sitios de buceo Parguera y Trampa
Tortuga, Faro y Cantil de la Piscinita (Com. Per. J. Veleño y D. Ladino).
 Es más frecuente ver ejemplares pequeños (juveniles) en el costado
occidental entre 2 y 25 m, pero se observan hasta 80 m (Com. Per. D.
Guggenheim y D. Ladino).
La población de pez león está aumentando en los arrecifes, pese a la pesca
de buzos y entidades (Coralina, gobernación, colegios, UNC).
ASPECTOS BIOLÓGICOS
Relación longitud peso
Histograma de frecuencia
de longitud
Peso total (g)
600
y = 1,0969e0,2292x
R² = 0,929
N=37
500
400
n
300
200
100
0
0
5
10
15
20
25
30
Longitud total (cm)
Crecimiento alométrico, con mayor aumento
en peso que en longitud.
Factor de Condición promedio K= 1.36028
(10x5). Alto comparado con otras especies ícticas
16
14
12
10
8
6
4
2
0
4.6
8.4
12.1
15.8
19.5
23.3
27.0
Ámbito de longitudes (cm) n=37
A partir de los 15 cm se empieza a ver el
sexo (en algunos casos)
ASPECTOS REPRODUCTIVOS
Sexo y estados gonadales P.
volitans (% n=40)
Proporción de sexos
P. volitans (% n=40)
Juvenil
52%
Hembra
20%
Inmaduro
Madurando
60
40
20
Macho
28%
Relación Hembra : Macho = 0.7
Mayor proporción de machos
0
Hembra Macho
Indifer.
Juvenil
Mayor % de machos madurando
ASPECTOS TRÓFICOS
Proporción de Grasa en Vísceras
Media
52%
Mucha
24%
Poca
24%
Sin
0%
Los peces analizados tenía grasa en un alto porcentaje, se usó la
escala propuesta por Lorenzo-Cobo (1993)
ASPECTOS TRÓFICOS
Proporción del Grado de Repleción
Proporción del Grado de
Digestión
Lleno
0%
Medio
lleno
39%
Fresco
7%
Vacio
16%
Casi
vacio
45%
Parcial.
Dig.
45%
Digerido
48%
ASPECTOS TRÓFICOS
Porcentaje de frecuencia por ítem
Porcentaje numérico por ítem
1.4
Resto moluscos
2.7
Resto Vegetal
42 %
Crustaceos
13.7
Peces
19.2
Otolitos
Resto peces
20.5
Crustaceos
20.5
58 %
Peces
21.9
M. Organica
0.0
5.0
10.0
15.0
20.0
25.0
0
20
40
60
ASPECTOS TRÓFICOS
Especies del contenido estomacal P.
volitans
 La población del pez león ha aumentado en los arrecifes coralinos de
isla de San Andrés y se empiezan a observar individuos en zonas de pastos
marinos y muelles del costado oriental; lo que hace probable que lleguen a
los manglares e impacten más la comunidad íctica y de crustáceos por la
abundancia de juveniles en ese ecosistema.
 Entre los peces hallados por nosotros en los contenidos estomacales de
Pterois se pueden mencionar serránidos y lábridos
 Entre los crustáceos identificados se destacan portúnidos, ermitaños,
galatheidos y stomatópodos
 Se requiere profundizar la investigación y emprender acciones
interinstitucionales e internacionales, para poder contribuir mejor al manejo
de AMP y RB Seaflower y por lo tanto del resto del Caribe.
 Trabajo de investigación interinstitucional
(Acero et al., 2010).
 Campaña de educación ambiental escrita
(Folleto UN Gavio, 2010), TV.
 Cursos para buzos, estudiantes,
pescadores y comunidad.
 Trabajo internacional – región (acciones
y protocolos conjuntos).
INSTITUTO DE ESTUDIOS CARIBEÑOS
CENTRO DE INVESTIGACIONES MARINAS
 Universidad Nacional de Colombia - Sede
Caribe, IEC y CECIMAR
Directivos
Estudiante Michel Orellano Chia
 Buzos del Caribe, San Andrés
David Guggenheim,
Edon Rafael Fraile Montego
Jaime Veleño
Manuel Angarita
Diego Ladino
Lanchero Marcos Urrego
Secretarias Mirna Zambrano y Yanet Freire
 Buzos independientes
Paolo Usseguio
Moises Gómez
 Fotos
A. Santos-Martínez, UNC - Sede
Caribe, IEC
David Guggenheim y colaboradores,
Buzos del Caribe, San Andrés
Amilcar Cupul, Universidad de
Guadalajara – Centro Universitario
de la Costa.
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