Management of Biological Invasions (2020) Volume 11, Issue 3: 372–398
CORRECTED PROOF
Research Article
A low number of introduced marine species at low latitudes:
a case study from southern Florida with a special focus on Mollusca
Fred E. Wells1,2,* and Rüdiger Bieler2
1 School
of Molecular and Life Sciences, Curtin University, GPO Box U1987, Perth, Western Australia 6845, Australia
Integrative Research Center, Field Museum of Natural History, Chicago, Illinois 60605, USA
2 Negaunee
Author e-mails: fred.wells@curtin.edu.au (FEW), rbieler@fieldmuseum.org (RB)
*Corresponding author
Citation: Wells FE, Bieler R (2020) A low
number of introduced marine species at
low latitudes: a case study from southern
Florida with a special focus on Mollusca.
Management of Biological Invasions 11(3):
372–398, https://doi.org/10.3391/mbi.2020.11.3.02
Received: 24 May 2020
Accepted: 22 July 2020
Published: 12 August 2020
Thematic editor: Katherine Dafforn
Copyright: © Wells and Bieler
This is an open access article distributed under terms
of the Creative Commons Attribution License
(Attribution 4.0 International - CC BY 4.0).
OPEN ACCESS.
Abstract
The anthropogenic transfer of non-indigenous marine species (NIMS) into new
areas of the oceans is a key issue. Despite increasing research effort in recent years
many fundamental questions remain to be answered before we can effectively manage
the issue. One question is whether the greater number of NIMS thus far documented
in temperate waters is real or an artefact of fewer surveys being undertaken in tropical
environments. Another one is whether poor taxonomic knowledge of the biodiverse
tropics hides NIMS that actually occur there. Extensive taxonomic work in three
Pacific localities (Guam, northern Western Australia and Singapore) has been
collated in previous papers showing that there are relatively few NIMS in these
biodiverse environments. The present paper replicates investigations for a low
latitude environment in southern Florida in the Atlantic Ocean. The focus area
includes the extensive Florida Keys coral reef environment, the southern margin of
the Everglades on Florida Bay and the major PortMiami. Only 48 NIMS were
identified in a literature-based compilation of 4,615 species; 15 species were
represented by isolated records and have not established populations, leaving only
33 NIMS that are established or whose status is unknown. Records for Mollusca,
the group with the most species (1,153) in the compilation, were individually
researched and taxonomically verified. It is argued that the relative paucity of
NIMS is not a straightforward temperature-driven tropical/temperate issue, but
instead there are biological factor(s) restricting the ability of NIMS to colonise
biodiverse environments compared to less diverse areas.
Key words: introduced marine pests, biotic resistance, artificial reefs, Florida Keys,
biodiversity databases, molluscs
Introduction
The anthropogenic transfer of non-indigenous marine species (NIMS)
from one part of the world’s oceans to another is one of the key issues in
protecting marine environmental diversity (Johnson and Chapman 2007;
Molnar et al. 2008; Katsanevakis et al. 2014b; Crowe and Frid 2015). There
has been growing concern about the increasing number of marine
invasions reported and their perceived effects. For example, the recent
introduction of the Indo-Pacific lionfish Pterois volitans and P. miles into
Florida and the Caribbean Sea (Albins and Hixon 2008; Hackerott et al.
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Introduced marine species in southern Florida
2013; Côté et al. 2013) has received extensive publicity. There are numerous
anthropogenic mechanisms for species introductions. Shipping, either as
biofouling (Hewitt 2002; Hewitt et al. 2004; Yeo and Chia 2010; Yeo et al.
2011; Jaafar et al. 2012) or in ballast water (Carlton 1985), is a dominant
component in most areas. Construction of canals, particularly the Suez Canal,
is another important cause. Deliberate introductions include aquaculture
species and the release of unwanted aquarium species. Inadvertent
introductions can include species attached to deliberate introductions, such
as organisms adhering to introduced oysters (e.g., Lavesque et al. 2020).
The numbers of NIMS are truly staggering. Eight hundred twenty-one
species are known to have been introduced to the Mediterranean Sea,
largely a result of Lessepsian migration through the Suez Canal (Zenetos et
al. 2017). A smaller number of species have migrated through the canal
from the Mediterranean to the Red Sea; the numbers of NIMS in both
areas continue to increase as new discoveries are made. A total of 343
NIMS has been recorded in Hawaii (Eldredge and Smith 2001). Fofonoff et
al. (2018) provide data on 276 marine and estuarine NIMS in California,
190 of which are in San Francisco Bay alone (Foss 2008). A study published
15 years ago recorded 99 NIMS in Port Philip Bay, Melbourne, Australia
(Hewitt et al. 2004) and undoubtedly there have been more introductions
since then. In addition, there are many cryptogenic species whose native
ranges cannot yet be determined, some of which may have been introduced
through anthropogenic mechanisms.
NIMS can have various deleterious effects by disrupting native ecosystems,
outcompeting local species, threatening commercial fisheries, introducing
diseases and fouling industrial structures (Hayes et al. 2005; Wells et al. 2009).
Fortunately, most of the NIMS have no apparent adverse effects; only a
small proportion become marine pests (Hayes et al. 2005; Wells et al. 2009).
Most studies have reported fewer NIMS in tropical waters than in
temperate environments (e.g., Coles and Eldredge 2002; Hewitt 2002;
Hutchings et al. 2002; Huisman et al. 2008; Hewitt and Campbell 2010;
Freestone et al. 2011, 2013). Several potential causes have been proposed
for this, including an increase in biotic interactions such as predation and
competition as a result of the higher tropical diversity making it more
difficult for species to become established (Hewitt 2002). Alternatively, it
has been suggested that the lower number of tropical NIMS is simply a
result of fewer studies resulting in fewer detections, or our lack of taxonomic
knowledge of the biodiverse tropics may result in NIMS remaining
undetected (Hewitt 2002). The relative paucity of NIMS in tropical
environments was specifically addressed by Hewitt (2002), who compared
the results of four tropical and four temperate surveys of Australian ports
conducted with the same techniques. Fifty-eight NIMS were detected; 48 in
the temperate ports and only 28 in the tropical ports.
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Introduced marine species in southern Florida
Wells (2018) investigated whether the apparent low number of tropical
NIMS is real or an artefact of a lack of taxonomic knowledge along the 800
km long coast of the Pilbara region in northwestern Australia. The shallow
water marine biota of the Pilbara has been extensively investigated,
primarily by studies led by the Western Australian Museum, and
identifications have been made of 5,532 species across a wide range of taxa.
Prior to the development of an iron ore mining industry in the 1960s the
Pilbara had been visited by relatively few vessels from overseas or
interstate, limiting the opportunities for NIMS introductions. This
changed in the early 2000s with the commencement of a ten-year boom in
iron ore mining and liquefied natural gas construction projects. Strict
marine quarantine procedures were instituted in the Pilbara to minimise
the introduction of NIMS and extensive monitoring programs were
undertaken to detect any species that had penetrated the quarantine
barriers. Only 17 NIMS have been detected in the Pilbara, compared to 54
in southern Western Australia. Only one species (the ascidian Didemnum
perlucidum) on the Australian national marine pest list of 55 species
(NIMPCG 2009a, b) occurs in the Pilbara; it also occurs on the west and
temperate south coast of Western Australia. In contrast 12 species on the
Australian national marine pest list occur in southern Australia (DAWE
2020).
The Pilbara study was repeated in Singapore (Wells et al. 2019). In contrast
to the relatively undisturbed Pilbara marine environment, international
trade in Singapore goes back at least to the 1300s. European vessels first
arrived in the 1500s, and vessel numbers increased rapidly in the early
1800s when Singapore became a British colony (Yeo et al. 2011). Singapore
is now one of the busiest ports in the world and is connected to over 600
ports in 120 countries (MPA 2017). In 2016 there were 138,998 vessel
arrivals involving a total of 593 million tonnes of cargo and a million
passengers. Arrivals included a large number of high-risk vessels, such as
barges, tugboats, dredges, oil rigs and similar vessels that remain in port
areas for long periods (MPA 2017). Following the downturn in
international shipping in 2008 associated with the global financial crisis
many vessels remained in port for months, increasing the risk of biofouling
and NIMS introductions (Floerl and Coutts 2009). Seebens et al. (2013)
ranked Singapore as the number one port in the world for the risk of
marine bioinvasions. The shallow water marine biota of Singapore has
been extensively studied by the National University of Singapore, with
3,650 species recorded, but Wells et al. (2019) found only 22 NIMS in
Singapore waters. Only three of these (the mussels Brachidontes striatulus,
Mytella strigata and Mytilopsis sallei) were potential marine pests.
The present study replicates the Pilbara and Singapore studies in low
latitude southern Florida. The location was chosen for several reasons. It is
in a different ocean, the western North Atlantic. The coral cay archipelago
Wells and Bieler (2020), Management of Biological Invasions 11(3): 372–398, https://doi.org/10.3391/mbi.2020.11.3.02
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Introduced marine species in southern Florida
of the Florida Keys is a large, coral reef marine environment that is
biodiverse and well documented. Although the Keys are outside the
tropics, the biota is tropical. The Florida Current originates in the South
Atlantic and Caribbean Sea and carries warm, marine water from the
Caribbean to the Keys (FKNMS 2020). Florida Bay, to the northeast of the
Keys, is shallow water and abuts the ecologically important Florida
Everglades, where the variety of invasive terrestrial and freshwater species
is a considerable problem (Ferriter et al. 2006). Salinities in Florida Bay are
highly variable. They reached a maximum of up to 70 PSU in the late
1980s. Monthly monitoring from 1998 to 2004 showed a range of 24 PSU
in October 1999 just after Hurricane Irene to 42 in July 2001 after a
drought (Kelble et al. 2007). The Miami area, which includes PortMiami, is
situated just north of the Keys, on the east coast of Florida’s peninsula. The
port advertises itself as the cruise capital of the world, with 55 cruise ships
operating from the port and visiting the Bahamas, Caribbean and Mexico.
In 2019, 958 cargo ships entered the port, importing 5.7 million tons of
cargo and exporting 4.4 million tons (PortMiami 2020). There were over
950,000 registered boats in Florida in 2018 (FHSMV 2020), many of which
are trailered between locations and thus provide an additional mechanism
for introductions.
Materials and methods
Separate literature and internet searches were undertaken of the shallow
water benthic marine biota of southern Florida, including macroscopic
invertebrates, fishes and marine plants. Marine birds, mammals, reptiles,
parasites and microscopic species were excluded.
The literature search commenced by examining a major three volume
work Gulf of Mexico–Origins, Waters, and Biota on the biota of the Gulf of
Mexico edited by Felder and Camp (2009). Numerous chapters in the
biodiversity volume provide information on specific taxonomic groups
authored by specialists in the respective taxa. While there is a consistent
format, there are some differences in the treatment of the different taxa.
The Gulf for the purposes of that study was divided into four quadrants,
with our study area falling into the border area of north-east and southeast quadrants. Few distributional point data are provided in that study, so
the taxonomic chapters were used to develop an Excel database of Gulf of
Mexico species in each taxon. Specific mentions of species occurring in the
Florida Keys and data on NIMS in the Gulf of Mexico (provided in some
chapters) were noted.
The literature search then built on the information sourced from Felder
and Camp (2009). Relevant references were examined for marine species
reported as occurring in the Florida Keys, Florida Bay and Biscayne Bay,
where PortMiami (25.77°N; 80.17°W) is located. Miami Beach is on the
north-eastern side of Biscayne Bay. Both the bay and seaward side of Miami
Wells and Bieler (2020), Management of Biological Invasions 11(3): 372–398, https://doi.org/10.3391/mbi.2020.11.3.02
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Introduced marine species in southern Florida
Figure 1. Project region at the southern tip of the State of Florida, encompassing several large
federally protected areas, including the Florida Keys National Marine Sanctuary (FKNMS, dark
blue) and three U.S. National Parks (Biscayne, Everglades, and Dry Tortugas). Two major
cities, Miami in Miami-Dade County (with PortMiami indicated by a box) and Key West in
Monroe County, are marked. 50, 100, and 200 m isobaths indicated.
Beach were included. The Florida Keys was broadly defined to include Dry
Tortugas (24.63°N; 82.92°W) and Florida Strait to a depth of 300 m (Figure 1).
The University of Miami commenced publication of the Bulletin of Marine
Science of the Gulf and Caribbean in 1951. Many early papers concentrated
on the biota near the university laboratory at Virginia Key in Biscayne Bay.
Over the years the geographical coverage of the journal broadened and it
was renamed in 1965 as the Bulletin of Marine Science. Because of the
regional relevance, all issues of the journal through 2019 were examined
for additional occurrence data and mention of NIMS.
The internet searches used Google Scholar to search for taxonomic
papers for the localities of Florida Keys, Florida Bay, Biscayne Bay, Miami,
Dry Tortugas and Florida Strait. Each locality was searched individually for
all of the taxa listed in Table 1 and also for general biotic surveys. We also
analysed records in the study area for all of the taxa in Table 1 from GBIF
(2019).
An Excel spreadsheet was constructed for each of the taxa in Table 1
showing species recorded for Florida Keys, Florida Bay and Biscayne Bay.
Whereas the spreadsheets provided a wealth of distributional data, we are
aware of quality control issues in such data compilations (e.g., as discussed by
Ball-Damerow et al. 2019) and have, were possible, checked outliers against
additional sources. In addition, it must be noted that the data capture and
georeferencing of North American marine invertebrate collections lags
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Introduced marine species in southern Florida
Table 1. Numbers of species recorded in various taxonomic groups in southern Florida and the primary sources of information.
Taxonomic group
Multiple taxa
Marine algae
Number of species
544
Seagrasses
Mangroves
Sponges
7
4
112
Corals
83
Other cnidarians
111
Nemerteans
Polychaetes
22
573
Sipunculans
Bryozoans
20
122
Crustaceans
813
Echinoderms
184
Brachiopods
Molluscs
10
1153
Ascidians
Fishes
23
834
Total
4615
Primary sources of information
Voss and Voss (1955); Smith et al. (2007); NOAA (1995); FLDEP (2018), GBIF (2019)
Tabb and Manning (1961); Humm (1963, 1964); Ballantine (1996); Croley and
Dawes (1970); Hine and Humm (1971); Mathieson and Dawes (1975); Zieman et
al. (1989); Frankovich and Fourqureane (1997); Dawes et al. (1999); Cho and
Fredericq (2006); Glardon et al. (2008); Fredericq et al. (2009); Burke et al. (2011)
Zieman et al. (1989); Ley et al. (1994)
Ley et al. (1994); UF/IFAS (2019)
Tabb and Manning (1961); Callahan (2005); Alvarez et al. (1998); Rützler et al. (2009);
Stevely et al. (2010)
Wheaton and Jaap (1988); Jaap et al. (1989); Cairns (2000); Callahan (2005);
Cairns et al. (2009)
Wallace (1909); Bayer (1961); den Hartog (1980); Wheaton and Jaap (1988); Jaap
et al. (1989); Fautin and Daly (2009); Opresko (2009); Cairns and Bayer (2009);
Calder and Cairns (2009)
Correa (1961); Norenburg (2009)
Treadwell (1911); Monro (1933); Hartman (1951, 1959); Taylor (1966); Ebbs
(1966); Perkins (1979, 1980, 1981, 1984, 1985); San Martín (1991, 1992);
Fauchald (1992); Rouse (1994); Fauchald et al. (2009); Bastida-Zavala et al. (2017)
Rice (2009)
McCann et al. (2007); Winston and Maturo (2009), Osburn (1914), Bastida-Zavala
et al. (2017)
Voss and Voss (1955); Provenzano (1959); Moore and McPherson (1963); Biffar
(1971); Moore et al. (1974); Abele and Kim (1986); Griffith (1987); Holmquist et
al. (1989); Thomas and Barnard (1992); Criales et al. (2000); Smith et al. (2007);
Felder et al. (2009); Gittings (2009); Reaka et al. (2009); Schotte et al. (2009);
Carlton et al. (2011), Tavares (2011).
Thomas (1962, 1964); Kier and Grant (1965); Singletary (1971); Pawson et al.
(2009)
Santagata and Tunnell (2009)
Clark (1994); Mikkelsen and Bieler (2000, 2004, 2007); Bieler and Mikkelsen
(2003, 2004a, 2004b, unpublished data); Bieler et al. (2004); Lyons and
Moretzsohn (2009); Rosenberg et al. (2009); Turgeon et al. (2009); Collins et al.
(2019); NPS (2020)
Plough and Jones (1939); Cole and Lambert (2009); Rocha et al. (2012)
Roessler (1970); Emery (1973); Jones and Thompson (1978); Sogard et al. (1989);
Thayer and Chester (1989); Ley et al. (1994); Serafy et al. (2003); Wiley and
Simpfendorfer (2007); McEachran (2009); Starck et al. (2017); Hepner (2017);
Bannerot and Schmale (2002)
behind those of vertebrates and flowering plants and the GBIF data
holdings do not yet reflect the majority of actual collections-based records
for many groups (e.g. Sierwald et al. 2018).
One of the authors of this paper (RB) has worked extensively over the
last two decades on the molluscan diversity of the Florida Keys and Florida
Bay (Bieler and Mikkelsen 2003, 2004a, b; Collins et al. 2019; Mikkelsen
and Bieler 2000, 2004, 2007), with focus on non-native taxa (Bieler et al.
2004, 2017). This provided the opportunity to verify individual published
records of NIMS for this phylum and to evaluate potential additions to that
list. To better ascertain actual distribution data for molluscs, we analysed
aggregated listings of museum collection records. For the region, there are
several data aggregators providing such services, e.g. (in order of increasing
taxonomic and geographic inclusiveness), InvertEBase (2020), IDigBio (2020),
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Introduced marine species in southern Florida
and GBIF. This information was used to update the molluscan species list
for the present study. We disregarded unique records of shells that were
likely discarded as food or decorative items; or were introduced to the
region as part of beach nourishment projects.
To avoid double counting, only taxa identified to species were included.
Those cited as tentatively identified (e.g., referred to family or genus sp. A,
sp. 1, etc.) were not included as there was no way of determining specieslevel identity across studies and there was no mechanism for assessing
whether or not the taxon was introduced to southern Florida.
Information was derived on NIMS during all of the above literature and
database surveys. In addition, a specific internet search was undertaken
using terms such as marine invasive species, introduced marine species,
etc. coupled with the specific localities. Further, broader databases on
introduced species in Florida and the United States were examined. In
particular, the National Exotic Marine and Estuarine Species Information
System (NEMESIS) (Fofonoff et al. 2018) was used to identify NIMS.
Species recorded during this search were checked against the World
Register of Marine Species (WoRMS 2019) and the names updated where
appropriate. The WoRMS category “marine” was used to determine the
habitat occupied by a species for inclusion on the NIMS species list. The
World Register of Introduced Marine Species (WRiMS 2019) was also
consulted. While WRiMS requires considerable work to verify the
information contained, it is the most comprehensive resource available.
One of the problems is that the native ranges of many widespread
species are not known; these species are referred to as cryptogenic. For
example, Fauchald et al. (2009) list 854 polychaete species from the Gulf of
Mexico, 181 of which are considered to be potential invaders (but these are
not specified in the publication). In a study such as the present paper, with
thousands of species across a wide range of taxa, it is not possible to
accurately determine the published ranges of all species. We have adopted
a very conservative approach of restricting the term cryptogenic to species
listed as such by Fofonoff et al. (2018). Two molluscs reported as “potentially
introduced” by Bieler et al. (2017) are also listed here as cryptogenic as they
essentially used that phrase as equivalent to cryptogenic.
Results
A total of 4,615 taxa were identified in the study area (Table 1). The most
diverse groups were molluscs (1,153 species), fishes (834), crustaceans
(813), polychaetes (573) and marine algae (544). Apart from polychaetes,
these are well known groups that tend to have large-bodied species.
Ascidians, a group known to include marine invasive species, were not well
represented in the study with only 23 species.
Forty-eight species are non-indigenous to the study area (Table 2);
an additional 19 species are cryptogenic (Supplementary material Table S1).
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Table 2. Non-indigenous marine species (NIMS) recorded in southern Florida. Abbreviations: Keys, Florida Keys; FB, Florida
Bay; BB, Biscayne Bay; WoRMS, World Register of Marine Species; WRiMS, World Register of Introduced Marine Species.
Group/Species
Location
Presumed source
WRiMS
Established
Notes
BB
Botanical garden
escape
No
Yes
Spread from cultivation at Fairchild Tropical
Botanic Garden where trees had been planted in
the late 1960s. Spread was discovered in 2008 and
eradication is ongoing (UF/IFAS 2019). Recorded
by GBIF (2019) from BB. The species closely
resemble the native Laguncularia racemosa (L.)
C.F. Gaertn.
Carijoa riisei (Duchassaing &
Michelotti, 1860)
Keys, BB
Shipping
Yes
Diadumene lineata (Verrill,
1869)
Keys, BB
Shipping
Yes
Tubastraea coccinea Lesson,
1829
Keys, BB
Shipping?
Yes
BB
Shipping
Yes
Yes
Keys, BB
Shipping
Yes
Yes
Keys, BB
Shipping
No
Yes
BB
Shipping
No
Rare
Amphibalanus amphitrite
(Darwin, 1854)
Keys, BB
Shipping
Yes
Yes
Amphibalanus reticulatus
(Utinomi, 1967)
Keys, BB
Shipping
Yes
Yes
Balanus trigonus Darwin,
1854
Keys, BB
Shipping
Yes
Yes
Caprella scaura Templeton,
1836
BB
Shipping
Yes
Unknown
Charybdis hellerii (A. MilneEdwards, 1867)
Keys
Shipping
Yes
Yes
Cyclograpsus integer H. Milne
Edwards, 1837
Ligia exotica Roux, 1828
Keys
Shipping
No
Yes
Keys, BB
Shipping
No
Yes
MANGROVE
Lumnitzera racemosa Willd.
CNIDARIANS
Presumed to be Although described from the Virgin Islands,
established.
Carijoa riisei is an Indo-West Pacific species. It
was first recorded from Dry Tortugas in 1869 and
BB in 1947 (Fofonoff et al. 2018).
Presumed to be Native range from Hong Kong to Japan. Widely
established.
introduced to east and gulf coasts (Fofonoff et al.
2018). Recorded from the Keys and BB by GBIF
(2019).
Yes
Indo-West Pacific species. Cairns (2000), Fenner
& Banks (2004); Ferry (2009), Precht et al. (2014),
Bieler et al. (2017), Fofonoff et al. (2018), GBIF
(2019).
POLYCHAETES
Ficopomatus uschakovi (Pillai,
1960)
Hydroides elegans (Haswell,
1883)
Poecilochaetus johnsoni
Hartman, 1939
Protula balboensis Monro,
1933
An Indo-West Pacific species reported by BastidaZavala et al. (2017) and Fofonoff et al. (2018).
Çinar (2013), Bastida-Zavala et al. (2017). The
map provided by Fofonoff et al. (2018) shows
Florida populations as cryptogenic, but the text
states the species is thought to have originated in
the Indo-Pacific and introduced in the western
Atlantic.
Described from southern California. Taylor (1966)
recorded the species in BB and other localities.
Recorded from the Keys and BB by GBIF (2019).
Thought to be an eastern Pacific species
introduced to the western Atlantic, but possibly
originated in the western Atlantic (Bastida-Zavala
et al. 2017; Fofonoff et al. 2018).
CRUSTACEANS
Possibly native to the Indo-West Pacific. Carlton
et al. (2011). Fofonoff et al. (2018) report that a
single specimen from Dry Tortugas may have been
from a ship's bottom. Moore and Frue (1974) from
BB. GBIF (2019) records from both the Keys and
BB.
Indo-West Pacific species. Carlton et al. (2011),
Fofonoff et al. (2018).
Broad Indo-West and eastern Pacific distribution
(Fofonoff et al. 2018). Recorded in southern
Florida by Carlton et al. (2011), Fofonoff et al.
(2018), GBIF (2019).
Described from Mauritius but origin is uncertain
(Fofonoff et al. 2018). Recorded from BB
(Fofonoff et al. 2018; GBIF 2019).
Indo-West Pacific. Fofonoff et al. (2018) reported
from Long Key, USGS (2019) from Monroe
County, and two GBIF (2019) two records in the
Keys.
Eastern Atlantic. Rathbun (1918) listed from Key
West and GBIF (2019) has several records.
Fofonoff et al. (2018) recorded as an Indo-Pacific
species found in Key West in 1883. Widely
distributed on US east coast and Gulf of Mexico.
GBIF (2019) records from both the Keys and BB.
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Table 2. (continued).
Group/Species
Location
Presumed source
WRiMS
Established
Notes
Limnoria pfefferi Stebbing,
1904
BB
Shipping
No
Unknown
Paradella dianae (Menzies,
1962)
Keys, BB
Shipping
Yes
Unknown
Penaeus monodon Fabricius,
1798
Penaeus vannamei (Boone,
1931)
Keys, BB
Aquaculture
release
Aquaculture
release
Yes
Yes
Yes
No
Pullosquilla litoralis (Michel
& Manning, 1971)
Keys
Yes
No
Sphaeroma terebrans Bate,
1866
Sphaeroma walkeri Stebbing,
1905
BB
Shipping,
aquarium release
or possibly
scientific
research.
Shipping
Indo-Pacific species recorded from Miami Beach.
Assumed to be a viable population (Fofonoff et al.
2018).
Native to west coast of North America, California
to Mexico. Recorded from BB (Fofonoff et al.
(2018). GBIF (2019) records from the Keys.
Tavares (2011), Fofonoff et al. (2018), USGS (2019)
and GBIF (2019) records from the Keys and BB.
Fofonoff et al. (2018) discuss the species in detail.
Wild catches are thought to have been the result of
escapes from aquaculture farms. No evidence of
living populations. GBIF (2019) records from both
the Keys and BB.
Indo-Pacific species. Single individual collected at
Key Largo in 1998 (Fofonoff et al. 2018).
Yes
Unknown
BB
Shipping
Yes
Unknown
Eualetes tulipa (Rosseau in
Chenu, 1843)
Keys, BB
Shipping
No
Yes
Hyotissa hyotis (Linnaeus,
1758)
Keys, BB
Shipping
Yes
Yes
Littorina littorea Linnaeus,
1758
Keys
Shipping
Yes
No
BB
Shipping
Yes
No
Keys
Discard?
Yes
No
Keys
Shipping
No
Yes
(single site)
BB
Shipping
Yes
Yes
Keys, BB
Shipping
Yes
Keys
Shipping
Yes
Yes
Didemnum psammatodes
(Sluiter, 1895)
Keys, BB
Shipping
Yes
Yes
Diplosoma listerianum (MilneEdwards, 1841)
Keys, BB
Shipping
Yes
Yes
Styela canopus (Savigny,
1816)
Keys, BB
Shipping
Yes
Yes
BB
Shipping
Yes
Yes
Keys, BB
Thought to be native to Indo-West Pacific but
widespread in tropical waters (Fofonoff et al. 2018).
Native to Indian Ocean, recorded from BB
(Fofonoff et al. 2018, GBIF 2019).
MOLLUSCS
Pinctada margaritifera
(Linnaeus, 1758)
Rapana venosa (Valenciennes,
1846)
Thylacodes vandyensis Bieler,
Rawlings & Collins, 2017
Eastern Pacific. Reported from southern Florida
for the first time in this paper. Earlier record by
Miller (1970) was misidentified as Petaloconchus
mcgintyi, from Miami. Widely introduced
elsewhere (e.g., Miloslavich 1995).
Indo-West Pacific. Bieler et al. (2004), Rosenberg
et al. (2009), Bieler et al. (2017). Apparently still
limited to artificial reef structures. Recorded by
GBIF (2019) in the Keys and BB.
Eastern North Atlantic. A single specimen from
Monroe County collected in 1952 (EDDSMapS
2020).
Indo-West Pacific species. Fofonoff et al. (2018)
provide several Florida records, including an
undated specimen from off Key West.
Western Pacific. Near Fiesta Key 1973 (single
live-collected specimen), USGS (2019).
Originally described from Keys shipwreck but
Bieler et al. (2017) provided molecular evidence
for a likely Pacific origin. Discussed as
“potentially invasive” by the authors. South
Florida (South Florida PBS 2018). Recorded by
GBIF (2019) in the Keys and BB.
ASCIDIANS
Ascidia sydneiensis Stimpson,
1855
Botrylloides violaceus Oka,
1927
Didemnum perlucidum
Monniot, 1983
Styela plicata (Lesueur, 1823)
Indo-West Pacific species reported by Fofonoff et
al. (2018).
Northwestern Pacific species (Fofonoff et al. 2018).
GBIF (2019) records from the Keys and BB.
Cryptogenic, but Florida populations are
introduced (Fofonoff et al. 2018). Simkanin et al.
(2016). GBIF (2019) records from the Keys.
Indo-West Pacific species recorded by Simkanin et
al. (2016), Fofonoff et al. (2018). GBIF (2019)
records from the Keys.
A complex of species, but introduced in the
northwest Atlantic (Fofonoff et al. 2018).
Recorded from the Keys and BB by GBIF (2019).
Indo-West Pacific species reported by Simkanin et
al. (2016), Fofonoff et al. (2018), GBIF (2019)
records from the Keys.
Possibly native to Northwest Pacific (Fofonoff et
al. 2018). Weiss (1948), Simkanin et al. (2016).
GBIF (2019) records from BB.
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Table 2. (continued).
Group/Species
Location
Presumed source
WRiMS
Established
Notes
BB
Aquarium release
No
No
Keys
Aquarium release
Yes
No
Keys
Aquarium release
Yes
No
Dascyllus aruanus (Linnaeus,
1758)
Keys, BB
Aquarium release
No
No
Hypsoblennius invemar SmithVaniz & Acero, 1980
Keys, BB
Shipping
No
Yes
Naso lituratus (Forster in
Bloch and Schneider, 1801)
Keys
Aquarium release
No
No
Platax orbicularis (Forsskål,
1775)
Keys
Aquarium release
Yes
No
BB
Aquarium release
Yes
No
Keys
Aquarium release
Yes
Yes
Keys, BB
Aquarium release
Yes
Yes
BB
Aquarium release
No
No
Zebrasoma flavescens
(Bennett, 1828)
Keys
Aquarium release
Yes
No
Zebrasoma veliferum (Bloch,
1795)
Keys
Aquarium release
Yes
No
Two individuals were removed from the Miami
Beach Marina in 2017 (USGS 2019). Considered
as of unknown status by Schofield & Akins
(2019).
Observed off Key West in 2006 (Schofield et al.
2009).
One specimen taken by spearfisherman off North
Key Largo (Huffpost 2013). Previously recorded at
several other Florida localities but considered to be
extirpated (Mundy 2005, Schofield et al. 2009).
One individual removed from Palm Beach County
in 2009 and two were sighted in the Miami Beach
Marina in 2017 (USGS 2019). GBIF (2019)
records from the Keys.
Native to Lesser Antilles and South America.
Schofield et al. (2009), Schofield & Akins (2019).
GBIF (2019) records from the Keys.
The species was sighted off Boca Raton in 2000
and 2001 and a single individual was removed
from Molasses Reef, Florida Keys in 2018 (USGS
2019).
Isolated specimens found off Keys and removed.
Schofield et al. (2009), Florida Museum (2019),
USGS (2019). GBIF (2019) records from Keys.
Considered as of unknown status by Schofield &
Akins (2019).
Three sightings have been made in Florida,
including one off North Miami Beach in 2002
(USGS 2019).
Widespread in southern Florida. Invasive
(Schofield 2009).
Widespread in southern Florida. Invasive
(Schofield 2009). GBIF (2019) records from the
Keys and BB.
Three individuals have been found in Florida,
including one in Biscayne National Park in 2018
(USGS 2019).
Isolated individuals recorded from several Florida
localities, including Marathon and Dry Rocks in
the Keys (USGS 2019).
Seen in three Florida localities from 2001 to 2003,
including Key Largo (Schofield et al. 2009). GBIF
(2019) records from the Keys.
FISHES
Acanthochromis polyacanthus
(Bleeker, 1855)
Cephalopholis argus
Schneider, 1801
Cromileptes altivelis
(Valenciennes, 1828)
Pomacanthus imperator
(Bloch, 1787)
Pterois miles (Bennett, 1828)
Pterois volitans (Linnaeus,
1758)
Zanclus cornutus (Linnaeus,
1758)
Six records of potential NIMS are rejected (Table S2). The 48 nonindigenous species are dominated by crustaceans (14 species), fishes (13)
and ascidians (7) (Table 2). Ten species of fishes, two crustaceans and three
molluscs have not established populations, leaving 33 NIMS that are
thought to have become established or whose status is uncertain. Twentyfour NIMS have been recorded from the Florida Keys and 28 from
Biscayne Bay.
Twelve of the non-indigenous fish species are thought to have been
introduced as aquarium releases. Specimens of the non-indigenous
mangrove were planted in a botanical garden. The species was later
discovered growing in a nearby stand of mangroves and has not yet been
eliminated. Two crustacean species were aquaculture releases and one
mollusc may have been discarded at the site where it was recorded. The
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likely introduction method of one species cannot be determined.
Interestingly, 29 of the 33 species of NIMS are thought to have been
introduced through shipping, but only one of the 13 fish species was
introduced through shipping.
Discussion
Overview of NIMS in Southern Florida
Ferriter et al. (2006) list 129 priority introduced terrestrial species and 83
priority introduced freshwater species the Florida Everglades, Florida Bay
and Florida Keys (non-priority taxa not listed). In contrast to the terrestrial
and freshwater environments, only 48 of the 4,615 marine species we
recorded in the southern Florida study area have been introduced, and
only 33 are believed to have established populations or whose population
status is uncertain. Twenty-four potentially established NIMS have been
recorded from the Florida Keys and 28 from Biscayne Bay.
The question arises: How complete is the analysis of NIMS in southern
Florida if there has been no specific survey for NIMS? However, even
targeted NIMS surveys such as the eight reported by Hewitt (2002) are
incomplete due to the limited extent of the surveys, absence of taxonomists
to identify key groups and the large number of cryptogenic species. Bishop
and Hutchings (2011) assessed the results of 46 NIMS surveys of
Australian ports and concluded that surveys for targeted species may
provide information on those species, but the surveys are not effective in a
broader context. Most NIMS are molluscs and crustaceans (Ruiz et al.
2000). Data for these taxa in southern Florida are extensive, with 1153
species of molluscs and 813 crustaceans recorded, but only 3 molluscs and
12 crustaceans were NIMS with established populations. Two of the
molluscs are apparently still limited to artificial reefs (the foam oyster
Hyotissa hyotis and the worm-snail Thylacodes vandyensis). Only one
species, the worm-snail Eualetes tulipa, is widely established. Likely of
eastern Panamic origin, it was first reported in Miami by Miller (1970) as
Petaloconchus mcgintyi, a misidentification. This species became tagged as
a potential invasive in Hawaiian waters (Bieler in Carlton 1999: 449; as
Vermetus alii). It is now known from many localities, including Venezuela
(Miloslavich 2009), Brazil (Spotorno-Oliveira et al. 2018) and India
(Jebakumar et al. 2015). Two other molluscs established in the Florida Keys
(the pyramidelloidean snail Cyclothyca pacei and the foam oyster Hyotissa
mcgintyi) are viewed as cryptogenic and need further study as to their
geographic origin. Fishes, with 834 species are well known in southern
Florida and NIMS have been closely monitored by Schofield and Akins
(2019). Of the 13 species of fish recorded as possible NIMS, ten are not
believed to have established populations (Table 2); only three are believed
to be anthropogenic introductions that are living in natural environments.
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Bryozoans can be a key component of NIMS (Wyatt et al. 2005). One
hundred twenty-two species were recorded in the present study, none of
which were NIMS. McCann et al. (2007) surveyed fouling communities of
four bays in Florida, the southernmost of which were Tampa Bay and
Indian River. Four bryozoan NIMS were detected; none of these were
recorded in the present study of southern Florida. Polychaetes are a major
gap. Although 573 species were recorded, their taxonomy and ranges are
poorly known. Çinar (2013) reported that 292 species of polychaetes have
been moved in world oceans by human transport and 180 have become
established; how many of these species are in southern Florida is not
known. Ascidians are another group with known NIMS. They are poorly
represented in the southern Florida data with only 23 species, but 7 of
these are NIMS. We conclude that, while incomplete, the data for southern
Florida are consistent with the completeness of similar studies.
Key invasive species in southern Florida
The two lionfishes, Pterois volitans and P. miles, established in southern
Florida (Schofield and Akins 2019), are having major impacts on native
reef systems (e.g., Green et al. 2012). Having been recorded in low
numbers along the Florida east coast since the 1980s, they were reported
from the Florida Keys in 2009 and have rapidly expanded in numbers
(Ruttenberg et al. 2012), especially along the reefs of the Florida Keys.
Various efforts are underway to remove lionfish from Florida waters,
including events for the general public (such as lionfish derbies and
removal days) to collect the species (FFWCC 2020).
Potential NIMS not found in southern Florida
There are a number of potential NIMS species that could be introduced to
southern Florida, that have not yet been recorded. It is interesting that
none of the 544 species of macroalgae recorded from southern Florida are
introduced despite the well-known invasives in the group and the presence
of records of introduced Caulerpa in other parts of Florida. Approximately
150 marine algal species have been introduced to the Mediterranean Sea
(Verlaque et al. 2004), with the genus Caulerpa attracting the most
attention. A small patch of an aquarium strain of C. taxifolia was first
detected off Monaco in 1984 (Meinesz et al. 1993). It became invasive and
spread rapidly in the northern Mediterranean (Glardon et al. 2008). The
strain was also reported from California (Jousson et al. 2000) and Australia
(Wiedenmann et al. 2001; Millar 2004). A second variety of C. taxifolia
discovered off southeastern Turkey (Cevik et al. 2007) spread westwards
and has been reported from Sicily (Picciotto et al. 2016). A third invasive
Caulerpa, C. cylindracea, has become widespread in the Mediterranean
(Verlaque et al. 2000; Boudouresque and Verlaque 2002) and the Canary
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Introduced marine species in southern Florida
Islands (Verlaque et al. 2004). Davidson et al. (2015) analyse the impacts of
these and other algal species worldwide. Jacoby et al. (2004) reported the
Indo-Pacific Caulerpa brachypus and native species becoming invasive in
Palm Beach County and Broward County, Florida. As aquaria are a
potential source of Caulerpa, Stam et al. (2006) examined the genetics of
256 individuals of Caulerpa being sold in aquarium shops and internet
sites and in field locations in Florida (including the Florida Keys), the
Bahamas, US Virgin Islands, and Honduras. Fourteen species were found.
Only a single individual of an invasive strain of C. racemosa was detected,
and this was in California.
The Asian green mussel Perna viridis is a highly successful invader
(Rajagopal et al. 2006) due to its short life span, rapid growth rate, rapid
sexual maturity, high fecundity, ability to colonise a wide range of habitats,
wide physiological tolerances, gregarious behaviour, suspension feeding
and ability to repopulate following a population crash (Morton 1997). It
was first detected in Trinidad, West Indies in about 1991, spread to
Venezuela by 1993 and now occurs widely in the Caribbean. The first
detection in Florida was in Tampa Bay in 1999 (Ingrao et al. 2001; Barber
et al. 2005). It has since been found at numerous localities on both the east
and west coasts of Florida (McGuire and Stevely 2018), but has not been
recorded in the Florida Keys. The absence of P. viridis in the Keys may
simply be an artefact, but it parallels the situation in northern Australia
where its absence may be due to oligotrophic water conditions (Huhn et al.
2017; Wells 2017).
The seagrass Halophila stipulacea was first detected in Grenada, West
Indies, in 2001 (Ruiz and Ballantine 2004). It is now widespread in the
Caribbean and is expected to continue its spread into the Gulf of Mexico
(Ruiz et al. 2017) but has not yet been recorded from southern Florida.
The Indo-West Pacific orange cup coral Tubastraea coccinea is
widespread in the Caribbean Sea and Gulf of Mexico, including the
southern Florida study area (Fenner and Banks 2004; Figueroa et al. 2019).
The congeneric T. tagusensis was recently detected at several sites,
including in northern Florida. Further research may well record the species
in southern Florida.
Methods of introductions
Apart from the apparent aquarium introductions of isolated individuals
and aquaculture introductions it is very difficult to determine the methods
of introduction of NIMS. As indicated above, biofouling (Hewitt 2002;
Hewitt et al. 2004; Yeo and Chia 2010; Yeo et al. 2011; Jaafar et al. 2012)
and ballast water (Carlton 1985) are the most important sources of NIMS
in most regions.
Fifty-five cruise ships operate from PortMiami, near the northern end of
Biscayne Bay. In 2018 the port also handled 1081 cargo vessels (PortMiami
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Introduced marine species in southern Florida
2020). There were 4,016 ship arrivals, including operations by 40 cruise ships
in 2019 in Port Everglades, 35 km north of PortMiami (Port Everglades
2020). Noting the difficulties in establishing introduction vectors discussed
above, shipping is the most likely source of 29 of the 33 NIMS recorded in
the study area (Table 2). Key West is only 200 km southwest of PortMiami
in a straight line, so it would be relatively easy for vessels to secondarily
disperse NIMS from either port to any location in the Keys.
Cruise ships have little requirement for ballast water. However, cargo
vessels delivered 5.7 million tons of cargo to Miami in 2019 and 4.4 million
tons were exported (PortMiami 2020). The ballast water associated with
these cargoes provides a mechanism for the importation of NIMS into
Miami, and also the export of species from the port. With the current
Covid-19 crisis many of the cruise ships have remained for months in
PortMiami with cruises suspended until 15 September (PortMiami 2020).
This increases the risk of biofouling accumulations with potential NIMS
species that could be exported to other ports once cruises resume (Floerl
and Coutts 2009).
Commercial trading vessels, including cruise ships and cargo vessels, are
regarded as low risk for the transfer of NIMS through biofouling as the
vessels have antifouling coatings, remain in ports for short periods and
move at relatively high speeds through the water. However, low risk does
not mean no risk. There are a number of areas on commercial trading
vessels where biofouling is likely to accumulate (Coutts and Taylor 2004;
DoF 2009). An antifouling coating (AFC) cannot be applied to some
structures, such as propellers and internal seawater systems. Vessels are
supported on blocks during drydocking when the AFC is applied; AFC
cannot be applied to the drydock support strips. All of these factors
increase the risk of biofouling accumulation, potentially including NIMS,
on the vessel.
Comparison of southern Florida with other geographical areas
The 33 NIMS in southern Florida is substantially fewer than 276 reported
for California (Fofonoff et al. 2018), 190 in San Francisco Bay (Foss 2008)
and 99 in Port Philip Bay, Melbourne, Australia (Hewitt et al. 2004).
Teixeira and Creed (2020) recently recorded 119 NIMS along the 8,000 km
coastline of Brazil. They demonstrated a generally increasing number of
NIMS with latitude, though the pattern was complicated by concentrations
of NIMS in areas where there were extensive maritime facilities. Even the
brief surveys conducted in four temperate Australian ports by Hewitt
(2002) reported more NIMS (58) than the present study. The low number
of NIMS in southern Florida is consistent with the paucity of NIMS
reported in other tropical studies: 85 in > 5,500 species recorded in Guam
(Paulay et al. 2002); 17 NIMS of 5,532 species in the Pilbara, northwestern
Australia (Huisman et al. 2008; Wells 2018); and 22 NIMS in 3,650 species
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Introduced marine species in southern Florida
in Singapore (Wells et al. 2019). These data support the contention that
NIMS are more common in temperate than tropical marine environments.
Hawaii, with 343 NIMS (Eldredge and Smith 2001) is an exception to
the low number of NIMS in the tropics. NIMS in Hawaii occur primarily
in disturbed areas and relatively few are in open coastal areas (DeFelice et
al. 2001). A rapid survey of 41 coral reef sites detected only 26 NIMS in 486
identified taxa; 17 were found at one or two sites and half of the sites had
three or less NIMS (Coles et al. 2006). However, Hawaii is biogeographically
isolated with a less diverse marine biota than other Indo-West Pacific
localities (Hutchings et al. 2002), suggesting the issue is not one of tropicaltemperate environments but instead an increased ability of NIMS to
colonise environments that are less biologically diverse.
Another example of increased NIMS in a less biologically diverse
environment is the Mediterranean Sea, which has a warm temperate biota.
Zenetos et al. (2017) reported that 821 invasive alien species have been
recorded in the Mediterranean. Katsanevakis et al. (2014a) divided the
introduction mechanisms of each species in the eastern Mediterranean into
four categories based on how well understood the introduction mechanism
was. Four hundred twenty species in the two best understood categories
were Lessepsian migrants through the Suez Canal, and additional Lessepsian
migrants have subsequently been reported (e.g. Steger et al. 2018). While
we could not find a species count, fewer anti Lessepsian migrations from
the Mediterranean into the Red Sea, which has a tropical biota, are known
(Rais Lasram et al. 2008). When the Suez Canal opened in 1869 two salinity
barriers restricted movement between the Mediterranean Sea and Red Sea.
The Bitter Lakes in the canal initially posed a high salinity barrier to
movement of species through the canal, but over time the salinity
decreased, removing the barrier. Also, the eastern Mediterranean had a
lower salinity than the Red Sea. Construction of the Aswan Dam in 1965
restricted flow from the Nile River, increasing salinity in the eastern
Mediterranean (Rais Lasram et al. 2008). Despite the salinity barriers, the
Red Sea bivalve Brachidontes pharaonis was first detected at Port Said at
the Mediterranean entrance to the canal in 1876 (Dogan et al. 2007). The
greater number of Lessepsian species is not due entirely to a greater
diversity of Red Sea biota as the dominant water flow in the Suez Canal is
from south to north (Rais Lasram et al. 2008).
Studies in Guam (Paulay et al. 2002), Pilbara, Western Australia (Wells
2018), Singapore (Wells et al. 2019) and southern Florida (this paper) are
all from biodiverse low latitude regions where the biota is relatively well
known, yet all four reported relatively few NIMS. Hewitt (2002) examined
four tropical and four temperate Australian ports using the same
methodology and found more NIMS in the temperate ports. This strongly
suggests that the relative paucity of NIMS in the studied environments is
not due to a lack of study or inability to detect NIMS caused by poor
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taxonomic knowledge, but rather by increased biological interactions in a
biodiverse environment (Hewitt 2002). One possible mechanism leading to
a larger number of NIMS in a less biodiverse setting is physical separation,
as was described by Zabin and Hadfield (2002), who found the Caribbean
barnacle Chthamalus proteus higher on the Hawaiian intertidal shoreline
than the native Nesochthamalus intertextus. Another approach was
undertaken by Freestone et al. (2011, 2013), who demonstrated that
increased predation could limit tropical invasions. Further study is
required to confirm these and possible other biological interactions to
explain the relative paucity of NIMS in diverse marine ecosystems.
Origin of NIMS in southern Florida
The presumed native ranges of 31 of the 33 NIMS recorded in southern
Florida are shown on Table 2: 21 species are from the Indo-West Pacific,
Northwestern Pacific or Indian Ocean; 1 (Balanus trigonus) has a broad
distribution in the Indo-West Pacific and Eastern Pacific; 6 are from the
Eastern Pacific; 2 are from the Eastern Atlantic; and 1 (the fish
Hypsoblennius invemar) is from the Lesser Antilles and South America. It
may have been introduced either naturally by currents or through shipping
(Schofield and Akins 2019).
PortMiami (2020) reports that cruises from the port travel to the
Bahamas, Caribbean and Mexico. Cargo trade is more widespread, with
46% with Latin America and the Caribbean, 37% to Asia and 16% to
Europe.
The generally north flowing oceanic circulation pattern in the region
provides a natural mechanism for the distribution of species from the Gulf
of Mexico and Caribbean to southern Florida. This is well illustrated by the
orange cup coral Tubastraea coccinea as discussed by Fenner and Banks
(2004) and Creed et al. (2016). The species was described from Bora Bora
in 1829 and was first recorded in the eastern Caribbean Sea in 1943. The
most likely source of the first introduction was by ship. The expanding
range of T. coccinea in the Caribbean and Gulf of Mexico follows the
pattern of the die-off of the urchin Diadema antillarum which began in
Panama in 1983–1984 and was spread by ocean currents. Tubastraea coccinea
was first seen on a shipwreck in Florida in 1999 and has subsequently been
found on a number of additional shipwrecks and other artificial habitats in
southern Florida (Fenner and Banks 2004; Creed et al. 2016; Bieler et al.
2017). As described above, Perna viridis was first detected in Trinidad in
about 1991 and since then has spread throughout the Caribbean and Gulf
of Mexico to both coasts of Florida (McGuire and Stevely 2018).
It is possible that some of the presumed naturally widespread ranges of
species known from the Caribbean, Gulf of Mexico and southern Florida
have in resulted from vessel traffic in the past. Such a possibility has been
discussed in detail in Singapore by Yeo et al. (2011).
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The fact that NIMS in southern Florida have been introduced from
other marine biogeographic regions rather than within the same region
parallels the situation in the previous studies in Western Australia (Wells
2018) and Singapore (Wells et al. 2019). The recent distribution of P. viridis
into eastern Indonesia (Huhn et al. 2017) is an exception.
It is noteworthy that many of the recognized NIMS and listed
cryptogenic species in southern Florida are larger-bodied and/or colourful
forms (foam oysters, orange cup corals, sea slugs, tropical fish) that are
more readily noticed than small-bodied and cryptic members of their
respective groups. However, in taxa where good data exist (e.g., molluscs),
there is no indication that smaller-bodied NIMS have escaped detection.
It is also noted that the current list for southern Florida will be modified
as additional information becomes available. Additional species are likely
to be recorded, either as new records or revised taxonomy of existing
known species. Alternatively, the number of NIMS in southern Florida may
be reduced as understanding of the origins of individual species improves.
For example, in the Mediterranean a group of experts in the taxonomy of
various phyla (Zenetos et al. 2017) made major changes to a NIMS list
published only a year earlier (Galil et al. 2016), excluding 72 species as
being not-established or native, but adding a similar number of new records.
In particular, application of recent advantages in genetic techniques will
enhance our understanding of NIMS, including those of southern Florida.
For example, Sun et al. (2017) used genetic barcoding to investigate the
status of the highly invasive serpulid polychaete Hydroides dianthus.
Although the species was described from New England and is considered
to be native to the east coast of North America, the genetic evidence
suggests it may have originated in the Mediterranean. In addition, a
distinct clade was detected in Texas that may represent separate species.
Similarly, Dias et al. (2018) used genetic techniques to detect three clusters in
the mytilid Perna viridis: (1) USA and Caribbean, (2) India and (3) Southeast
Asia. Figueroa et al. (2019) used morphological analyses to demonstrate
that there are three clades of the orange cup coral Tubastraea in the Gulf of
Mexico: T. coccinea, T. tagusensis and an intermediate clade. The
morphological results were confirmed by genetic analyses.
Thus, the present paper synthesizes our current understanding of NIMS
patterns in the southern Florida study area. The details will undoubtedly
change as more information is developed in future.
Acknowledgements
This paper draws on the work of many scientists who have documented the marine biota and
non-indigenous marine species in southern Florida and other parts of the world. We are grateful
to all of them for providing the information we have used. Dr John Huisman of the Western
Australian Herbarium, Perth and Dr Pamela Schofield of the United States Geological Survey,
Gainesville, Florida kindly provided literature and data that we could not otherwise access. We
appreciate the advice of three reviewers whose comments significantly improved the
manuscript.
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Funding Declaration
Research in the protected waters of the Florida Keys was conducted under Florida Keys
National Marine Sanctuary Research Permit FKNMS-2009-024 and Florida Keys National
Wildlife Refuge Special Use Permit 41580 to RB. The funders had no role in study design, data
collection and analysis, decision to publish, or preparation of the manuscript.
Ethics and Permits
Research on regional gastropods was funded under US National Science Foundation (NSF)
award DBI-0841760, on bivalves under award DEB-0732854 to RB. Coral reef-related studies
in the Florida Keys were supported by a grant from the Paul M. Angell Family Foundation to RB.
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Supplementary material
The following supplementary material is available for this article:
Table S1. Cryptogenic marine species recorded in southern Florida.
Table S2. Doubtful and rejected records of non-indigenous marine species (NIMS) recorded in southern Florida.
This material is available as part of online article from:
http://www.reabic.net/journals/mbi/2020/Supplements/MBI_2020_Wells_Bieler_SupplementaryMaterials.xlsx
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