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| Poster presented at 2015 Florida Fish and Wildlife Commission, Florida Fish and Wildlife Research Institute, Fisheries-Independent Monitoring Annual Meeting |
Range of the Zebratail Blenny, Hypleurochilus caudovittatus Bath, 1994, with comments on identification of the Hypleurochilus complex in the Gulf of Mexico
¹Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, Charlotte Harbor Field Laboratory, 585 Prineville Street, Port Charlotte, FL, 33954, USA
²Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, 100 8th Avenue SE, St. Petersburg, FL, 33701, USA
3Univeristy of Florida, Florida Museum of Natural History, Ichthyology Department, Dickinson Hall, Newell Dr. and Museum Rd., P.O. Box 117800, Gainesville, FL 32611, USA
*Corresponding author: Email: Chrystal.Murray ttarpon@gmail;com
Running head: range of Hypleurochilus caudovittatus
Abstract
Background
Range-restricted marine species present compelling case studies for testing
hypothesized latitudinal shifts due to climate change. We examined the
distribution of the recently described Zebratail Blenny, Hypleurochilus
caudovittatus, historically thought to
occur only in the northeastern Gulf of Mexico. Because of its recent
description, it was necessary to search collections for three additional Hypleurochilus
species to identify all available
specimens.
Results
Cephalic pore branching complexity was the most reliable way to distinguish
between H. caudovittatus
and its congeners (Fig. 1). Based on multi-decadal
fisheries-independent and museum data, H. caudovittatus ranges from Perdido Key, Florida in the northeastern Gulf of Mexico to
at least off Marco Island, Florida in the southeastern Gulf of Mexico. This
species reaches at least 64 mm standard length and occurs in estuaries and continental
shelf waters from 0.5 m to at least 30 m deep.
Conclusions
We propose H. caudovittatus as a candidate species to study the future effects of climate change
because of its restricted range and existing barriers that may prevent range
adjustments.
Keywords
Blenniidae
– climate change – range-restricted fishes – larval dispersal – semi-enclosed
seas
Background
We
examined the distribution, habitat, and hydrographic
requirements of the Zebratail Blenny, Hypleurochilus caudovittatus Bath 1994, which was thought to occur
only within a narrow range in the northeastern Gulf of Mexico, to establish a
historically accurate range with which to gauge future potential change.
Efforts to identify a species’ range entail searching fishery-independent and
museum collection databases to best determine a historical range that can act
as a baseline for future comparisons (Booth et al. 2011).
With
the recent description, we suspected that specimens were present in museum
collections that had been identified as closely-related blenniids. Thus, to
accurately establish the range of H.
caudovittatus, it was also necessary to examine records of three sympatric Hypleurochilus species[PG1] :
Crested Blenny, Hypleurochilus geminatus
(Wood 1825); Featherduster Blenny, H. multifilis (Girard 1858); and Barred
Blenny, H. bermudensis Beebe and Tee-Van 1933.
Models for future climate change
predict gross changes in the distribution of fishes, even over slight
temperature changes (Roessig et al. 2004). Coastal and estuarine fishes with restricted
ranges are vulnerable to local extinction if the effects of climate change
limit access to key habitats or generate barriers that prevent adequate larval
dispersal. Range-restricted marine species present
compelling opportunities for studies of latitudinal shifts due to ongoing
climate change and ocean warming (Parmesan 2006),
and recent studies have begun to link these factors to range shifts in fishes (Sorte et al.
2010; Last et al. 2011; Feary et al. 2014). Species currently residing near
their thermal tolerance limits are most at risk (Booth et al. 2011). For some
species, such as Alewives, Alosa
pseudoharengus (Wilson 1811), and American shad, Alosa sapidissima (Wilson 1811), along the east coast of the U.S.,
a northerly shift in ranges has been documented (Nye et al. 2009). As water
temperatures increase, species trapped by the northern boundary will not be
able to escape (Biasutti et al. 2012), and they could be extirpated from the
south (Cheung et al. 2009). Perry et al. (2005) documented a similar range
retraction from the south for 13 taxa in the semi-enclosed North Sea, including
the Snakeblenny, Lumpenus lampretaeformis
(Walbaum 1792); Blue Whiting, Micromesistius
poutassou (Risso 1827); and Redfishes, Sebastes
spp.
Using
long-term fisheries-independent monitoring data and museum specimens, our goals
were to revise the range and maximum size of H. caudovittatus, discuss its environmental preferences, and review
species descriptions to explore novel characteristics that separate H. caudovittatus from possible sympatric
congeners.
Methods and Materials
Field Sampling
Twenty-four years (1989–2013) of the
Fisheries-Independent Monitoring (FIM) program of the Florida Fish and Wildlife
and Conservation Commission’s Fish and Wildlife Research Institute data were queried for H.
caudovittatus. These data were collected in eight major estuaries on both
coasts of Florida, including Florida Bay at the southern tip of the peninsula.
Early FIM monitoring included fixed-station and seasonal stratified-random
sampling. Beginning in the late 1990’s, sampling has been monthly using gear
that targets small fishes: center-bag seines (21.3 × 1.8-m, 3.2-mm stretch
mesh) and otter trawls (6.1-m, 38-mm stretch mesh, 3.2-mm stretch mesh liner).
Associated data include locality, depth, habitat, physiochemical conditions,
and standard length (SL). Detailed data collection and sample processing
procedures are outlined in Poulakis et al.
(2004).
Offshore fisheries-independent monitoring data from the 2008–2013
Southeast Area Monitoring and Assessment Program (SEAMAP)
groundfish trawl surveys were also queried for H. caudovittatus. These data were collected in 4–104 m depths along
the West Florida Shelf from the Dry Tortugas north to the Florida-Alabama
border in the northern Gulf of Mexico (Switzer et al. 2015). Briefly, the
survey uses a standard 12.8-m SEAMAP shrimp trawl towed at 3 knots for 30
minutes. Additional survey details are outlined in Rester (2014).
Museum specimens
Specimens
examined for this study came from eight museum collections (abbreviations
follow Sabaj Pérez
2014): AMNH = American Museum of Natural History, New York, NY; NMNH = National
Museum of Natural History, Smithsonian Institution, Washington, DC; UF = Florida Museum of Natural History, University of Florida,
Gainesville, FL; CAS = California Academy of Sciences, San Francisco, CA; SU
= Stanford University, ichthyology
collection housed at CAS; FSBC = Florida Fish and Wildlife Research Institute,
St. Petersburg, FL; TCWC = Texas A&M University Texas Cooperative Wildlife
Collection, College Station, TX; and UMMZ = University of Michigan Museum of
Zoology, Ann Arbor, MI. Notes on the geographic distribution of Hypleurochilus were queried from online
collections data. We examined H.
bermudensis, H. geminatus, and H. multifilis from the western Atlantic
and the Gulf of Mexico to compare cephalic pore branching complexity, cirri
counts, anal fin ray counts, and pigment as separating characteristics.
Taxonomy follows Page et al. (2013).
Diagnostic
characteristics selected for this study were based on previous studies of Hypleurochilus in the Caribbean,
Florida, Gulf of Mexico, and Atlantic Ocean (Randall 1966; Bath 1994; Williams
2002; Hopfensperger 2003; McEachran and Fechhelm 2005).
Hypleurochilus spp. from museum collections that were
re-identified for this study are indicated below by an asterisk. Official museum
records have been updated to reflect the corrected identifications. Materials
examined are listed alphabetically by institution and then by collection date
(Table 1).
Hypleurochilus
caudovittatus—Museum collections were examined from Perdido Key, Florida to offshore of Marco Island,
Florida in the northeastern Gulf of Mexico. AMNH
16927* two specimens, near Tarpon Springs, 13 November 1941; 16926* one
specimen, Tarpon Springs, 23 November 1941, condition: stiff, specimen
rehydrated following Singer (2014); FSBC 1090 one specimen, 14.74 nautical
miles west of Long Key, 31 January 1959; 7469 one specimen, 11.3 nautical miles
west of Siesta Key, 17 July 1970; 18213 one specimen, Holmes Beach, 24 July
1994; 19411 Charlotte Harbor, 21 July 1997; 19063 one specimen, Tampa Bay, 08
September 1998; 19412 one specimen, Pine Island Sound, 12 October 1998; 19122
one specimen, Tampa Bay, 08 February 1999; 25279 three specimens, Charlotte
Harbor, 20 April 2011; 25281 one specimen,
Bokeelia, 17 May 2011; 28643 Caloosahatchee River,
23 January 2015; SU 60103* one specimen, Pine Island Sound, Cabbage (formerly
Palmetto) Key, 07 August 1938; TCWC 659.02 two specimens, Cedar Key, Gomez
Island, 11 June 1976; UF204038* four specimens, Boca Grande Pass, 06 July 1958;
185898* one specimen, east jetty at St. Andrew Bay, 22 July 1959; 76253 three
specimens, St. Marks Channel, 30 January 1977; 42090* two specimens, off Cedar
Keys between Seahorse and Snake Keys, 11 October 1985; UMMZ 108026* one
specimen, Lemon Bay, 07 March 1935; USNM 131928* one specimen, Cortez, 20 March
1906; 34724* one specimen, Cedar Key, December 1883.
Hypleurochilus geminatus—Museum collections of this
species were examined from Galveston Island, Texas to Panama City, Florida in
the northern Gulf of Mexico (GOM) and Fort Matanzas to the Indian River Lagoon along
the east coast of Florida. AMNH 8986, two specimens,
Indian River Lagoon, Florida, 07 November 1920; 77323, one specimen, Florida,
Santa Rosa, Fort Pickens jetties, 05 July 1958; 85353 one specimen, Florida,
Santa Rosa, Fort Pickens jetties, 01 October 1977; 52059 three specimens, off
Mobile, Dauphin Island and vicinity, Alabama, July 1982; 249303 one specimen,
GOM, Mississippi, Ocean Springs, 19 July 2006; 249336 one specimen,
Mississippi, Ocean Springs, 19 July 2006; CAS 213389 two specimens, GOM, Florida, Pensacola, Santa Rosa, no collection
date;[ME2] [MC3] FSBC12016 two specimens,
Florida, St. Lucie Inlet, 10 August 1967; TCWC 2615.04* one specimen, GOM,
Florida, near Panama City, St. Andrews Park, 24 August 1978; 11311 four
specimens, GOM, Texas, Galveston Island, 26 April 2001; 11315 one specimen,
GOM, Texas, Galveston Island, 26 April 2001; 11327 four specimens, GOM, Texas,
Galveston Island, 26 April 2001;
11316; 16 specimens, GOM,
Texas, Galveston Island, 25 May 2001: UF 2734, one specimen, GOM, Florida,
Escambia County, 15 August 1953; 67439* three specimens, GOM, Florida,
east jetty at St. Andrew Bay, 22 July 1959; 153504 one specimen, GOM, Florida,
Santa Rosa Sound, 10 October 1976; UMMZ 139406 one specimen, Florida, Matanzas
River south of Matanzas Inlet, 19 August 1936; USNM 49711 one specimen, GOM,
Florida, Big Marco Passage, 1901.
Hypleurochilus bermudensis—Museum collections of this species were examined from the
northeastern Gulf of Mexico off Mobile, Alabama to the Dry Tortugas off south
Florida. AMNH 86897* one specimen, GOM, Alabama,
Mobile Ship Channel, 22 April 1981; USNM 116805 seven specimens, GOM, Florida
Keys, Dry Tortugas, no collection date; UF 5699 one specimen, GOM, Florida,
jetties at Panama City, 30 September 1956; 11844 three specimens, GOM, Florida,
Dry Tortugas, east side of Loggerhead Key, 23 May 1965; 153074, one specimen,
eastern GOM, Florida, west of Sanibel Island, 23 October 1977; 28370 one
specimen, eastern GOM, Florida, twelve-foot ledge off St. Petersburg, 08 June
1979.
Hypleurochilus multifilis—Museum collections of this
species were examined from the northern Gulf of Mexico off Louisiana to the
panhandle of Florida. AMNH 260605* two specimens,
GOM, Florida, Santa Rosa, Fort Pickens, 05 July 1958; 87262* three specimens,
GOM, Florida, Panama City rock jetty, Saint Andrews State Park, 26 June 1973; 83995* two
specimens, GOM, Florida Middle Ground, 21 May 1975[PG4] ; 35753* one specimen,
GOM, Alabama, Mobile, Dauphin Island, 117 October 1975; 260604* two specimens,
GOM, Alabama, Mobile, Dauphin Island and vicinity, July 1982; TCWC 11315* one
specimen, GOM, Texas, Galveston Island, 26 April 2001; 11316 one specimen, GOM,
Texas, Galveston Island, 25 May 2001; UF 134634 five specimens, GOM, Florida,
Choctawhatchee Bay, 05 July 1958; 69206* one specimen, GOM, Florida, Destin, west jetties at
East Pass, mouth of Choctawhatchee Bay, 27 June 1968; 70188* one specimen, GOM,
Florida, St. Andrews State Park, 14 July 1970; USNM 217324* three specimens,
GOM, Louisiana, 07 September 1977.
Analysis
Multiple techniques were used to identify and analyze fishes used
in the study. Photographs of cephalic pore branching complexity were included
to aid identification. To highlight cephalic pore branching complexity, Hypleurochilus
spp. were stained with Alcian Blue 8 GX dissolved in 90% ethanol. The stain was
applied to the head of the fish with a number 0 fine paint brush to highlight
the pores and the cluster connections under the skin. Small amounts of Alizarin
Red S and Rose Bengal Disodium Salt were used to enhance the skin surface. A length-frequency was plotted to document overall size
distribution, including freshly caught and museum specimens.
Results
Diagnosis
Combtooth
blennies (family Blenniidae) of the genus Hypleurochilus
(Gill 1861) inhabit mostly subtropical and tropical
seas and are small, cryptic fish with scaleless tube-shaped bodies. They are demersal
spawners with modified ventral fins for clinging to structure in strong
currents and a single row of comb-like teeth. Hypleurochilus are further characterized by the following: gill
openings restricted to the side of the head extending ventrally to level of
pectoral-fin base; elongated supraorbital cirri and nasal cirri (Fig. 1); smooth ventral edge on upper lip; no
teeth on the vomer; large recurved caniniform teeth generally at rear of both
jaws; dorsal fin XII, 13–16; anal fin II, 14–17; pectoral fin 14; pelvic fin I,
3-4; truncated or rounded caudal fin; segmented caudal fin rays 13–15 with some
principle rays branched; continuous dorsal fin with slight notch between
spinous and soft-rayed sections (Randall 1966; Bath 1994; Williams 2002; Hopfensperger
2003; McEachran and Fechhelm 2005).
Previous
studies used the branching pattern of supraorbital cirri to distinguish Hypleurochilus species (Bath 1994;
Hopfensperger 2003), but we found this characteristic was highly variable by
species and size. However, the cephalic pore system is an arrangement of raised
sensory canals transiting though the neurocranium, terminating in dermal pores.
The complexity of canal branching and grouping of the associated dermal pores were
considered when examining this characteristic. We found cephalic pore branching
to be useful in distinguishing Hypleurochilus
spp. in all but the smallest (<20 mm SL) specimens. We looked at the
following cephalic canals and associated dermal pores to determine cephalic
pore branching complexity: infraorbital (IOP) (posterior to eye), preopercular
(POP), and supratemporal (STP) (above and behind eye) (Fig. 1). In H. caudovittatus and H. geminatus there
are (simple condition) single, unbranched canals leading to individual dermal
pores of IOP and POP series. Pores in the
STP group in H. caudovittatus were mostly
branched, whereas those in H. geminatus
were mostly single and unbranched. In H. bermudensis (intermediate
condition) half or less of the pores in IOP and POP lead to branched dermal
pores with the STP group forming a floret shape (Hopfensperger 2003). In H.
multifilis (complex condition) there are groups of three or more dermal
pores in IOP, POP, and STP series (Hopfensperger 2003), with an STP pore count of
ten or greater and the STP group forming a multibranching floret[ME5] [MC6] .
Focal species description
Hypleurochilus
caudovittatus Bath, 1994
Cephalic
pore branching complexity simple, rarely intermediate; supraorbital cirri 1–8;
nasal cirri 1–4; caudal fin with three or four dark bands on a translucent
background; anal-fin II, 16; a dark spot may occur on membrane between first
and second dorsal-fin spines (Bath 1994; Williams
2002; Hopfensperger 2003; McEachran and Fechhelm 2005).
Observed
in aquaria, colors are variable; head and body dark purple or gray with brown
blotches and mottling or dark olive or blackish purple, perhaps with white
outer margin on anal fin (Fig. 2). Dorsal and anal fins may be mottled. Caudal
fin with 4–6 distinct dark bars over white background. In formalin-preserved
and ethanol-stored condition, caudal fin retains bands and head and body are dark
gray to yellowish-brown with blotches. In older preserved specimens, the body
is orange-yellow and the blotches fade, but the caudal fin bands are apparent.
Based
on the holotype, UF 100283, Williams (2002) reported the maximum length of H. caudovittatus as 48.7 mm standard length (SL) (= 59 mm total length (TL)). However,
H. caudovittatus
captured in fisheries-independent samples collected between 1989 and 2013 (see
Methods and Materials) ranged from 11 to 66 mm SL (mean = 31.0 mm; Fig. 3[PG7] ). The three largest H. caudovittatus (FSBC 25281 and FSBC
28643), were 64 mm, 64 mm, and 66 mm SL (maximum TL = 76.0 mm).
Hopfensperger
(2003) reported a maximum depth of occurrence of 20.1 m. Our fisheries-independent
samples have increased the maximum depth of occurrence to 33.1 m.
Range extension of Hypleurochilus caudovittatus
A comprehensive investigation of Hypleurochilus collected during fisheries-independent sampling
(1989–2013) and housed in museum collections expanded the known range of H. caudovittatus (Table 2; Fig. 4). The
range of H. caudovittatus was
originally established by Bath (1994) using capture localities of types and comparative
material—west and northwest coasts of Florida, Sarasota (New Pass) to St.
Andrew’s Bay, northwest end of Shell Island. This range has been repeated in
subsequent publications (Williams 2002; Hopfensperger
2003; McEachran and Fechhelm 2005; Kells and Carpenter 2011; Williams et al.
2014).
In
inshore habitats, H. caudovittatus have
been collected outside the previous range from within the Charlotte Harbor
estuarine system. Specimens have been collected in fisheries-independent
samples in Lemon Bay (south of Sarasota), southern Pine Island Sound (near
Captiva Island), and in the mouth of the Caloosahatchee River (salinity range: 17.1–39.0; mean: 31.4 Practical Salinity
Units (PSU)). The Caloosahatchee River specimen is the
southernmost H. caudovittatus from
inshore collections.
In
offshore habitats, H. caudovittatus have
been collected outside the previous range from localities further west and
south. Specimens
have been collected in fisheries-independent samples west-southwest of Marco
Island and Boca Grande Pass, south-southwest of Cedar Key, west-southwest of
Horseshoe Beach (northwest of Cedar Key), and south of the western end of
Perdido Key. The Marco Island specimen extends the southern range by 189.2 km,
and the Perdido Key specimen extends the western range by 171.8 km.
Extensive
faunal studies have reported no H.
caudovittatus from Florida Bay (Flaherty et al. 2013), the Dry Tortugas (Switzer et al. 2015), the Florida Keys
reef tract (Smith et al. 2011), or along the east
coast of Florida (Kupschus and Tremain 2001; Solomons
and Tremain 2009). We reidentified 13 H. caudovittatus in museum collections
from the northern Gulf of Mexico, the West Florida Shelf, and contiguous passes
and bay systems. One reidentified H.
caudovittatus (SU 60103) was collected in Pine Island Sound during August
1938 and is the southernmost specimen from museum collections.
Based on these new data, we amend the traditional range and revise
it to 97.2 km west southwest of Marco Island, Florida to 4.73 nm south of the western
end of Perdido Key, Florida (Fig. 4).
Outtaxa descriptions
Hypleurochilus geminatus[PG8] :
Cephalic pore pattern simple (Fig. 1); supraorbital cirri 5–11; nasal cirri 2–4;
anal-rays II, 17; dorsal-fin spot present; no caudal-fin pigment. Specimens examined
ranged from 13-73 mm SL (16-80 mm TL). Maximum known length is 89 mm TL (Williams
2002). Museum specimens examined (see Methods and Materials) occurred within
the previously published range: New Jersey to east-central coast of Florida
(Williams 2002) and the entire U.S. Gulf of Mexico coast south to at least Veracruz, Mexico (Smith-Vaniz et al. 2014).[MC9] [PG10]
Hypleurochilus bermudensis:
Cephalic pore pattern intermediate (Fig. 1); supraorbital cirri 1–6; nasal
cirri 1–6; anal-rays II, 15; dorsal-fin spot present; in life and preserved—body
with six broad saddles above mid-line. Specimens examined ranged from 19 to 48 mm
SL (23–56 mm TL), and maximum known length is 100 mm TL (Williams 2002). Museum
specimens examined (see Methods and Materials) occurred within the previously published
range: Bermuda, Bahamas, Florida Keys, Dry Tortugas, and Gulf of Mexico (Williams 2002; Williams et
al. 2014).
Hypleurochilus multifilis:
Cephalic pore pattern complex (Fig. 1); supraorbital cirri 21 or fewer
branches; nasal cirri 1–15; anal-rays II, 17; dorsal-fin spot present; may have
dark spot at caudal-fin base. Specimens examined ranged from 49 to 97 mm SL,
and maximum known length is 102 mm TL (Williams 2002). Museum specimens
examined (see Methods and Materials) mostly occurred within the published range: Gulf of
Mexico (Williams 2002; McEachran and Fechhelm 2005; Smith-Vaniz et al. 2014) [MC11] two specimens were re-identified from
the Florida Middle Grounds during this study (AMNH 83995)
and suggests H. multifilis
occurs in depth greater than 18 meters.
Discussion
One factor that defines the range of H. caudovittatus, and its congeners, is habitat. Mature
Hypleurochilus spp. are typically restricted
to structured, hard-bottom habitats including estuarine reefs, inlet and pass
jetties, and coastal marine reefs (Hastings and
Springer 2009). Hopfensperger (2003) and McEachran and Fechhelm (2005)
reported that H. caudovittatus is
generally found on sandy substrates; however, according to Macpherson (1994)[PG12] [MC13] the majority of blenniid species
show a negative association with muddy-sandy bottoms. Bath (1994) stated that H. caudovittatus is common inside inlets
along the Gulf of Mexico and associated with hard bottoms along the west
Florida coast. Extensive areas of marsh, bayou
systems, and terrigenous sediment dominate the region west of our revised
northern range (Gore 1992), which may act as a barrier to western range
expansion for H. caudovittatus.
Similarly, these habitats may influence populations of other Hypleurochilus spp., especially those
that occur east and west of the Mississippi River.
Hypleurochilus
caudovittatus examined in our study were
captured inshore and offshore over the limestone dominated West Florida Shelf. Small
(<48 mm SL) H. caudovittatus collected during inshore sampling were
associated with sponge and bryozoan bycatch,
suggesting these are suitable inshore habitats for juveniles. The low
number (n=10) of H. caudovittatus >47.8
mm SL from our sampling suggests that H. caudovittatus moves from juvenile
habitats to more complex and under-sampled habitats in rocky passes and
offshore. The greatest number of H. caudovittatus
documented for this study (96.2%) were
captured within the Central Barrier (Cape Romano (25°50'35.91"N, 81°40'50.15"W[MC14] ) –Tarpon Springs, Florida) and Big
Bend Limestone (Tarpon Springs–Alligator Point, Florida (29°53'35.84"N,
84°22'53.81"W) coasts described by Gore (1992) (Fig. 5).
Hydrography
is another range-limiting factor. Weisberg et al. (2014)
determined that near-bottom currents transport larval, pre-settlement Gag, Mycteroperca microlepis (Goode and Bean 1879) across the West Florida Shelf
(WFS) toward shore. Presently, there is no published information on the larval
duration of Hypleurochilus; however, Raventós and Macpherson[PG15] [MC16] (2001) documented a short
pelagic larval duration of <50 days for the Blenniidae. In this study, no H.
caudovittatus occurred outside of the WFS. This raises the
hypothesis that shoreward transport of pre-settlement H. caudovittatus is similarly
directed by the deep-water circulation across the WFS. Examination of SEAMAP ichthyoplankton sample data collected in the Gulf of
Mexico may corroborate onshore-directed transport as a factor that limits Hypleurochilus
spp. dispersal.
Salinity is a third major factor in the distribution of fishes in
all life stages. At the northwest boundary of the H. caudovittatus range, seasonal freshwater outflow from the
Mississippi River and neighboring rivers produces salinity fields <28 PSU
entrained in eddies proximate to the Mississippi–Alabama–Florida (MAFLA) Shelf
and Desoto Canyon (Morey et al. 2003). At the
southern edge of the H. caudovittatus range,
managed freshwater discharge from the Florida Everglades generates dramatic
changes in salinity (Montague and Ley 1993). Hypleurochilus caudovittatus
captured during inshore and offshore sampling showed an affinity for high salinities (32.2–35.7 PSU). Periodic episodes of lower
salinity on either side of the revised range is likely a major barrier for
settlement of H. caudovittatus. Other Hypleurochilus
spp., especially those with more extensive ranges such as H. geminatus and H. multifilis,
may be able to tolerate lower salinities.
Hypleurochilus
caudovittatus captured in intensive
sampling inshore and offshore over the West Florida Shelf are likely
under-reported due to under-sampling in high-relief areas anywhere within its
range (i.e., gear selection). The relatively low number of H. caudovittatus collected over
25 years of intensive fisheries-independent sampling indicates either that gear
selectivity is marked or that this species is not abundant anywhere in its
range.
During the course of this study, as we confirmed and re-identified
museum specimens using the cephalic pore patterns, several avenues of possible
future research emerged. For example, H.
geminatus has only recently been
confirmed in the Gulf of Mexico and its extent remains largely undocumented.
Similarly, H. multifilis is poorly known and our research suggests that it is
more widely distributed that previously thought. In addition to the need for
more taxonomic studies, future research on larval dispersal and salinity
tolerance may help clarify the ranges of these interesting species.
Conclusions
In
this study, we have extended the known geographic range of H. caudovittatus and provided data on habitat use by this species. This information may prove useful
should future conservation strategies be required to manage the species. In
addition, H. caudovittatus, with its distinct range in the eastern Gulf of
Mexico, presents an interesting case for monitoring range shifts due to climate
change. Its restricted distribution is ideal for periodic vulnerability
assessment to determine if the range shrinks from the south.
Further
studies of H. caudovittatus should focus on serial collection of larval stages
and adults and include age-and-growth studies, genetic analysis (at this time
there is one DNA barcode sequence available in public records), and
histological examination to determine breeding season and population stability.
Continued monitoring will be necessary to best define trigger points that
affect the range of H. caudovittatus and other range-restricted
fishes in semi-enclosed seas.
Declarations
Acknowledgements
We thank Sabine Jessel at the Senckenberg
Research Institute and Natural History Museum[PG17] [MC18] ;
Barbara
Brown and James Van Tassell (AMNH); Victor Springer, Jeffery Williams, and
Sandra Raredon (NMNH); Rob Robins Jr. and Randal Singer (UF); David Catania (CAS
and SU); Joan Herrera, Alfred Thomson, and Molly Phillips (FSBC); Charles
Idelberger, Alejandro Acosta, Richard Paperno, and Robin Grunwald (FWRI); Heather Prestridge (TCWC); Douglas
Nelson (UMMZ); James Maclaine at the
British Museum of Natural History (NHM); Luke Tornabene (TAMUCC); and Ramon
Ruiz-Carus, Benjamin Victor, and Mitchell Roffer who were not associated with
any institution, but contributed curatorial assistance, specimens, data, and
expertise. Alfred
Thomson and Bland Crowder[PG19] improved earlier versions of the
manuscript.
Availability of data and materials
Data
available in a public (institutional, general, or subject specific) repository
that does not issue datasets with DOIs (non-mandated deposition).
Competing interests
The
authors declare that they have no competing interests.
Author’s contributions
CM conceived the study, obtained and identified specimens, and wrote the manuscript. JH identified specimens, conducted length-frequency analysis, and drafted sections of the manuscript. RM edited the manuscript and aided species identification. JD provided data analysis and created the range map. GT cleared, stained, and photographed the specimens for Fig. 1. GP developed the study, aided procurement of specimens, wrote and edited the manuscript. All authors read and approved the final manuscript. This manuscript remains unpublished as of 2018.
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Correspondence should be addressed to: C. H. Murray, Fish and Wildlife Research
Institute, Florida Fish and Wildlife Conservation Commission, Charlotte Harbor
Field Laboratory, 585 Prineville Street, Port Charlotte, FL, 33954 USA, Chrystal.Murray@MyFWC.com.
Summary of Zebratail Blenny,
Hypleurochilus caudovittatus material examined. See Figs.1 and 3 for pictures of characteristics.
|
Material examined |
Cephalic pore |
Supra-orbital cirri |
Nasal cirri |
Anal fin rays |
Caudal fin pigment |
|
AMNH
16927-1 |
Simple |
5 |
2 |
16 |
Bands
present |
|
AMNH
16927-2 |
Simple |
4 |
1 |
16 |
Bands
present |
|
FLMNH
42090-1 |
Simple |
4 |
1 |
16 |
Bands
present |
|
FLMNH42090-2
|
Simple |
5 |
2 |
16 |
Bands
present |
|
FLMNH
67439-4 |
Simple |
4 |
2 |
16 |
Bands
present |
|
FSBC
07469 |
Simple |
6 |
2 |
16 |
Bands
present |
|
FSBC
18213 |
Intermediate |
8 |
3 |
16 |
Bands
present |
|
FSBC
19063 |
Simple |
4 |
2 |
16 |
Little
pigment |
|
FSBC
19122 |
Simple |
5 |
2 |
16 |
Bands
present |
|
FSBC
19411 |
Simple |
4 |
1 |
16 |
Bands
present |
|
FSBC
19412 |
Simple |
3 |
2 |
16 |
Bands
present |
|
FSBC
25279-1 |
Simple |
3 |
2 |
16 |
Bands
present |
|
FSBC 25279-2 |
Simple |
5 |
2 |
16 |
Bands present |
|
FSBC 25279-3 |
Simple |
5 |
3 |
16 |
Bands present |
|
FSBC
28643 |
Simple |
7 |
2 |
16 |
Bands
present |
|
SU
60103 |
Simple |
5 |
2 |
16 |
Bands
present |
|
TCWC
659.02-2 |
Simple |
1 |
4 |
16 |
Bands
present |
|
TCWC
659.02-3 |
Simple |
6 |
2 |
16 |
Bands
present |
|
UMMZ108026
|
Simple |
6 |
2 |
16 |
Bands
present |
|
USNM
34724 |
Simple |
3 |
1 |
16 |
Bands
present |
|
USNM
131928 |
Simple |
5 |
3 |
17 |
Bands
present |
|
USNM
034724 |
Simple |
3 |
1 |
16 |
Bands
present |
Source
data for the Zebratail Blenny, Hypleurochilus
caudovittatus range extension. See Methods and Materials for museum codes.
|
Source |
Specimen |
Latitude |
Longitude |
Location off
Florida |
|
Type |
UF 68927 |
30°07'20.82"N |
85°43'47.22"W |
St. Andrew’s Bay |
|
Type |
FSBC 7469 |
27°19’41.90”N |
82°35’20.00”W |
Sarasota |
|
Museum |
SU 60103 |
26°38' 60.00"N |
82°13'00.00"W |
Pine Island Sound |
|
Inshore |
FSBC 28643 |
26°31'31.26" N |
82°08'27.48"W |
Caloosahatchee River |
|
Offshore |
SEAMAP 2009 |
30°11' 39.00"N |
87°31'16.80"W |
Perdido Key |
|
Offshore |
SEAMAP 2013 |
25°37’17.30”N |
82°38’35.00”W |
Southwest of Marco Island |
Fig. 1
Cephalic sensory canal pore complexity for (A) Zebratail Blenny, Hypleurochilus caudovittatus, (B) Crested Blenny H. geminatus, (C) Barred Blenny, H. bermudensis, and (D) Featherduster Blenny, H. multifilis. Infraorbital pores (IOP), preopercular pores (POP), and supratemporal pore (STP) complexity distinguish the species. BCC = branched cephalic canal. Note: branching and counts of supraorbital and nasal cirri (C) are highly variable and should not be used to distinguish the species. Photo credits: G.E. Ramos-Tafur.
Fig. 2
Length
frequency of Zebratail Blenny, Hypleurochilus
caudovittatus. Black bars refer to collections by Florida’s Fisheries-Independent
Monitoring program; white bars refer to museum collections. Numbers indicate numbers
of specimens from each source. The shape of the curve suggests that the
complete size range of the species is represented.
Fig. 3
Life coloration of the Zebratail Blenny, Hypleurochilus caudovittatus FSBC 25281 ♂ standard length 64.0 mm, total length 74.0 mm. Collected from 26°43'13.98"N 82° 09'09.18"W Charlotte Harbor, southwest coast of Florida, eastern Gulf of Mexico, 17 May 2011.
Revised[PG24] range for the Zebratail Blenny, Hypleurochilus caudovittatus. SEAMAP = Southeast
Area Monitoring and Assessment Program trawl survey; SRS = Stratified-Random
Sampling.
Fig.
5
Number
of Zebratail Blenny, Hypleurochilus
caudovittatus by latitude in the Gulf of Mexico, Florida, 1989–2013. The low
number of fish caught at 28°N is likely due to limited fisheries-independent
sampling effort.