NOAA Technical Memorandum NMFS NE 118
Tautog (Tautoga onitis) Life History
and Habitat
Requirements
by Frank W. Steimle1 and
Patricia A. Shaheen2
1National
Marine Fisheries Serv., Highlands, NJ 07732
2Rutgers Univ., New
Brunswick, NJ 07102
Print
publication date May 1999;
web version posted April 12, 2001
Citation: Steimle FW, Shaheen PA. 1999. Tautog (Tautoga onitis) Life History and Habitat
Requirements. US Dep Commer, NOAA Tech Memo NMFS NE 118; 23 p.
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Abstract
This report compiles and summarizes available information
on the tautog (Tautoga onitis), covering nomenclature and taxonomy,
distribution and habitat, reproduction, development, growth, feeding
and diet, behavior, population structure, natural and human-induced environmental
factors, and ecological roles. The report also identifies research needs
and includes an extensive bibliography.
Recent declines in this species' abundance and certain
known aspects of its life history and specific habitat requirements have
caused coastal fishery resource and habitat managers to believe the species
may need further conservation measures. Essential to developing an effective
conservation management strategy for this species is a thorough summary
of what is known of the species' life history and of the habitat requirements
for all life stages. This information will be important for developing
holistic approaches to managing a sustained population and fishery for
this species and closely associated members of its ecological community.
This review shows that although much is known about the
species, and studies on its life history and ecology are ongoing, there
are important gaps and conflicting or unconfirmed results in our understanding
of the species, and its needs, which should be addressed.
INTRODUCTION
The
tautog (Tautoga onitis) is a valuable recreational and commercial
fishery resource from Massachusetts to Virginia in the Northwest Atlantic.
It is commonly found on complexly structured, vegetated, or reef-like
habitats during post-larval stages. Fishery interest in the species has
increased in recent years, and additional management measures for this
fishery are being considered. To support the development of information
necessary to produce good public policy, an assemblage of existing information
on the species is needed (Atlantic States Marine Fisheries Commission
1995). This report compiles and summarizes available information on the
life history and habitat requirements of, and natural and human-induced
environmental threats to, tautog. It builds upon and substantially expands
the previous efforts of Auster (1989), Gray (1992), and others. This
review is needed because of: 1) recent increased fishing effort for,
and resource user conflicts over, tautog (DiLernia 1993); 2) sensitivity
of reef fish such as tautog to exploitation (Hostetter and Munroe 1993);
3) the species' particular habitat needs and the threats to this habitat;
and 4) the possible need to exercise additional management of at least
certain localized populations of this species (Hostetter and Munroe 1993).
Information on regional-level stock abundance, and detailed discussion
of state-level populations and harvests, are not included here, but have
been compiled by Lazar (1995).
As there are incidental references to this species in many documents
and papers, and as there are ongoing studies wholly or partially involving
this species, such a review cannot be definitive. It can, however, serve
as a stepping stone to adequate knowledge for fishery resource and/or
habitat management, or for fishery research planning, as have previous
reviews. Because of the scarcity or absence of certain information on
tautog, relevant information from studies of the its close labrid relative,
the cunner (Tautogolabrus adspersus) -- which has similar habitat
needs -- has sometimes been considered and discussed as probably an appropriate
estimate for tautog.
NOMENCLATURE
AND TAXONOMY
NOMENCLATURE
Valid Name
Tautoga onitis (Linnaeus, 1758) is the name recognized by the
American Fisheries Society (Robins et al. 1991). The generic term
is the original name applied to this fish species supposedly by Narragansett
Indians. The species term is derived from the Latin, onitis, which
means "a kind of plant"; but its application by Linnaeus is unclear (Smith
1907).
Synonymy
The species has a rich synonymic history which, following Jordan and
Evermann (1896-1900) and Jordan et al. (1930), includes: Labrus
onitis (Linnaeus 1758 and 1766); L. hiatula (Linnaeus 1766); L.
carolinus (Bonnaterre 1788); L. blackfish (Schopf 1788); L.
subfuscus (Walbaum 1792); L. tesselatus (Block 1792); Hiatula
gardeniana (Lacepede 1800); L. americanus (Block and Schneider
1801); L. tautoga rubens (Mitchill 1814, 1815); L. tautoga
alia (Mitchill 1814, 1815); Tautoga niger (Mitchill 1814,
1815); L. tautoga fusca (Mitchill 1815); T. tessellata (Cuvier
and Valenciennes 1839); T. americana (DeKay 1842); T. onitis (Gunther
1862; Uhler and Lugger 1876; Yarrow 1877; Jordan and Evermann 1898; Evermann
and Hildebrand 1910; Fowler 1912); H. onitis (Jordan 1886; Jenkins
1887; Bean 1891); and H. hiatula (Goode and Bean 1885).
TAXONOMY
Description
and Affinities
This species is a member of the Labridae, a family of lipped fishes
commonly called wrasses. This family takes its name from the presence
of conspicuous, thick, longitudinally folded lips which, along with other
characteristics such as the form of the jaws, give the mouth a peculiar
appearance (Smith 1907; Liem and Sanderson 1986). The mouth is terminal
and considered of small or moderate size (Hildebrand and Schroeder 1928).
Westneat (1995) suggests that tautog may have characteristics which are
primitive within the family, mostly based on its hard-prey diet and associated
jaw morphology.
Worldwide, there are 500-600 species in the Labridae (Nelson 1984).
In the United States, there are 12 genera (Robins et al. 1991).
Five labrid species occur in North Carolina waters (Smith 1907), but
only two species are commonly found north of Cape Hatteras: the cunner, Tautogolabrus
adspersus (Walbaum, 1792), and the tautog, Tautoga onitis (Linnaeus,
1758), (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953).
The genus Tautoga (Mitchill) consists of a single species: Tautoga
onitis. The following generic description is from Hildebrand and
Schroeder (1928): "Body is elongate, moderately deep and compressed;
anterior profile rather strongly arched, head nearly as deep as long;
eye small, placed high; mouth rather small; lips quite broad and thick" (Figure
1). "Teeth in the jaws strong, the anterior ones more or less conical
and incisor-like." Both the roof of the mouth and the floor of the
throat (i.e., pharynx) have a patch of knob-like teeth that
are used to crush and grind mollusk and crustacean prey (Bigelow and
Schroeder 1953; Liem and Sanderson 1986). The scales are small, with
about 70 being in the lateral series, but cheeks and operculums are
largely scaleless. The dorsal fin is long and continuous,with the soft
part short. The caudal fin is short and round to slightly truncate.
The anal fin has three stout spines, with the soft part similar to
that of the dorsal fin (Hildebrand and Schroeder 1928).
Normal coloring of this species is variably dull blackish, brownish,
blackish green, or blackish blue, with sides irregularly mottled or blotched.
The lips, chin, throat, and belly are often lighter. Larger males are
gray with white markings on the caudal, pelvic, and dorsal fins, and
on the chin -- a very conspicuous characteristic. Females and smaller
males are without white markings on fins and chin, and their fins are
often plain like the color of the body. Their eyes are green. Juveniles
are usually colored green or brown with more distinctive side mottling
and three or more darker bars. This coloring can vary with the visual
characteristics of the habitat they are using. In some areas, two color
patterns are often recognized; one being plain blackish and the other
having irregular blackish bars on a pale background. Fish are also observed
being dull gray-white. Some of these variations could reflect different
environmental factors, such as light or stress. (See the "Color
Modulation" section of the "Behavior" chapter.)
The tautog can be distinguished from its close and co-occurring relative,
the cunner, in several ways. The adult tautog is longer and stouter than
the adult cunner. The dorsal profile of its head is highly arched, while
in the cunner it is relatively straight. The caudal peduncle of the tautog
is proportionally wider, and the caudal fin is narrower, than those of
the cunner (Bigelow and Schroeder 1953; Leim and Scott 1966). Tautog
lack scales on the cheeks and operculums, while on cunner they are present
(Hildebrand and Schroeder 1928).
Subspecies
No subspecies are recognized. However, the differential juvenile growth
rates at the range extremes of the species, discussed in the "Growth" chapter,
have been thought possibly to reflect some degree of evolutionary divergence,
but this has not been supported in the laboratory (Martin 1993).
Common and Vernacular
Names
Tautog is the common name for T. onitis accepted by the American
Fisheries Society (Robins et al. 1991), but "blackfish" is also
widely used. Other regional common names include: Canada -- tautogue
noir (Leim and Scott 1966); Maine -- white chin (Bigelow and Schroeder
1953); New York -- blackfish (Goode 1887; Hildebrand and Schroeder 1928);
New Jersey -- smooth blackfish, tautog, and chub (Goode 1887); Maryland
-- black porgy, salt-water chub, chub, and blackfish (Hildebrand and
Schroeder 1928); Virginia -- moll and will-george (Goode 1887); and North
Carolina -- sea tench (in a 1709 usage) and oyster-fish (Goode 1887;
Smith 1907).
The Maine name "white chin" refers to the conspicuous white coloration
of the lower jaw of older males (Bigelow and Schroeder 1953; Hostetter
and Munroe 1993). Another name used in some places is "slippery bass" which
refers to the tautog's mucus covering and rough resemblance to a bass.
Jordan et al. (1930) also mention the names "cub" and "sea dog" being
used for the species.
DISTRIBUTION
AND HABITAT
GENERAL DISTRIBUTION
The tautog is a generally a coastal species found on the Atlantic coast
of North America, from the outer coast of Nova Scotia to South Carolina
(Bigelow and Schroeder 1953). It has been anecdotally, and perhaps dubiously,
reported on central Georges Bank from commercial fishery catch data (Chang
1990). The report of tautog on Georges Bank is not reliably supported
by Northeast Fisheries Science Center (NEFSC) bottom trawl survey records,
although it has been suggested that a few tautog may be collected near
the boundaries of the bank every decade (T. Azarovitz, pers. comm.1).
The tautog may be a "relict" species north of Massachusetts, found in
certain, deep, saltwater lakes and protected bays of Nova Scotia that
have waters that become warmer in summer and remain slightly warmer in
winter (Bleakney 1963).
The tautog is most abundant from Cape Cod to Chesapeake Bay. North of
Cape Cod, it is unusual to find tautog more than 6 km from land, or in
waters deeper than 18 m. South from Cape Cod to about New Jersey, it
can be found to 19 km offshore in up to 24 m of water, and occasionally
near the deep Great South Channel between Nantucket Shoals and Georges
Bank (Chang 1990). This offshore distance and depth range appear to increase
gradually towards the south and near Cape Hatteras (Chang 1990; Hostetter
and Munroe 1993). Tautog have been reported in brackish water, but not
in freshwater (Bigelow and Schroeder 1953). It has been reported up to
70 km upstream from the mouth of the Hudson River (New York) (Beebe and
Savidge 1988), and formerly in the Patapsco River in Baltimore, Maryland
(Fowler 1912).
It is uncertain if tautog populations at the northern and southern extremes
of its range were introduced by man. To the north, reports that the tautog
was introduced to Cape Ann (Massachusetts) in the late 1800s were countered
by other reports that it had been abundant many years previous to that
time. To the south, reports that it had been introduced to South Carolina
were countered by skepticism that its range could be artificially extended
southward (Goode 1887). Tautog are currently rarely observed or caught
in South Carolina waters (M. Bell, pers. comm.2).
In terms of the persistent, overall range of the species, the potential
effects of long-term global warming (or cooling) are unknown. Continued
global warming could theoretically increase northern coastal water temperatures
and potentially the presence of the species (and others) in the Gulf
of Maine, and perhaps restrict tautog to north of Cape Hatteras. Such
a northern expansion could have occurred since the last glacial period.
Empirically, however, it appears that recent warming of air temperatures
in the Arctic has hastened the melting of the polar ice cap which has,
in turn, increased the flow of cold freshwater into the North Atlantic.
This expansion of the North Atlantic "cold pool" has already reduced
the habitat for young Atlantic salmon, and may well be, or become, a
distribution-reducing factor for cold-intolerant fish species (K. Friedman,
pers comm.3).
DIFFERENTIAL
DISTRIBUTION
Spawning
Adult tautog generally migrate inshore in spring from coastal wintering
sites to spawn (Chenoweth 1963; Cooper 1966; Stolgitis 1970; Olla et
al. 1974; Briggs 1977). Spawning occurs primarily at or near the
mouths of estuaries and in inshore waters (Tatham et al. 1984;
Feigenbaum et al. 1989; Sogard et al. 1992; Able and Fahay
1998). Inside Narragansett Bay (Rhode Island), mature tautog returned
to the same spawning sites each year, usually in the upper estuary, but
dispersed throughout the bay after spawning (Cooper 1966; Dorf 1994;
Dorf and Powell 1997). Olla et al.'s (1980) tagging studies suggested,
however, that adult tautog did not always return to the same spawning
site in the spring, and that population mixing from different localities
occurred.
A portion of the adult population reported to remain offshore throughout
the year (Olla and Samet 1977; Eklund and Targett 1990; Adams 1993; Hostetter
and Munroe 1993), especially in the southern part of its range, was found
seasonally in spawning condition. For example, ripe fish were collected
by Eklund and Targett (1990) and Hostetter and Munroe (1993) on hard-bottom
sites 25-35 m in depth and 22-37 km off Maryland and Virginia. Collections
of eggs and larvae from Georges Bank to North Carolina through the NEFSC's
Marine Resources Monitoring, Assessment, and Prediction Program (MARMAP)
surveys, suggest that tautog spawning may also occur in continental shelf
waters and be concentrated off Southern New England (Sogard et al. 1992).
Dorf (1994), in contrast, speculates that the occurrence of offshore
eggs and larvae can also result from their being flushed from estuarine
spawning areas, at least in Southern New England.
Spawning is reported to follow a northward progression through the summer,
beginning in April in the southern part of the Middle Atlantic Bight,
and extending to the northern areas by May (Able and Fahay 1998). Peak
spawning in the central Bight is reported to occur in June and July,
and to decline by August (Berrien and Sibunka, in press).
Eggs and Larvae
Tautog eggs and larvae are collected on the inner continental shelf,
with highest concentrations being collected off Southern New England
and Long Island (New York) (Colton et al. 1979; Sogard et al. 1992;
Malchoff 1993). In the New York Bight (i.e., the continental shelf
off Long Island and New Jersey), larvae are reported to be part of a
summer coastal fish larval assemblage, perhaps closely associated with
spawning areas (Cowen et al. 1993). Tautog is the most abundant
of any larval species found recently in Narragansett Bay (Keller and
Klein-MacPhee 1992). In the Weweantic River Estuary (Massachusetts),
the greatest abundances of eggs and larvae were collected over eelgrass
(Zostera marina)-vegetated sites and near bottom (Stolgitis 1970).
As mentioned earlier, Dorf (1994) believes that eggs and larvae found
in offshore coastal waters can result from ex situ sources (i.e.,
flushed out and away from estuaries), but there are inadequate data to
support this hypothesis at present. Viable tautog eggs, although lacking
oil globules, are buoyant and found in greatest abundance at or near
the water surface (Merriman and Sclar 1952; Herman 1963; Stolgitis 1970;
Fritzsche 1978; Bourne and Govoni 1988). Nonviable eggs lack or lose
buoyancy and probably respond to currents differently, and thus probably
become distributed differently (Perry 1994).
Malchoff (1993) reported that tautog larvae migrate vertically in the
water column. The larvae, 2-4 mm in length, stay near the water surface
(less than 5 m deep) during the day, but go deeper at night. Older and
larger larvae spend more time at deeper depths as they grow. Malchoff
(1993) also believed the Hudson River plume strongly influences larval
tautog transport patterns in that part of the New York Bight. Sogard et
al. (1992) estimated that larval tautog in coastal New Jersey spend
about 3 wk in the plankton before settling to the benthos, but this period
can be a short as 17 days (Schroedinger and Epifanio 1997). This is a
relatively short planktonic period compared to that of other labrids.
This short period could possibly reflect an adaptation to cool-temperate
environments and a restricted period of optimum warmer conditions for
somatic growth.
Juveniles
Newly settled tautog inhabit shallow areas less than 1 m in depth (Warfel
and Merriman 1944; Sogard et al. 1992; Hostetter and Munroe 1993),
including tide pools (Breder 1922). Bigelow and Schroeder (1953) noted
that "fry" were often seined from Southern New England to Virginia. Sogard et
al. (1992) reported that as young-of-the-year (YOY) tautog grow,
they move to areas greater than 1 m deep. Cooper (1964) noted that juveniles
(no size range given) in Narragansett Bay were not observed in waters
deeper than about 9 m.
Several studies reported that young tautog (less than 10 cm) prefer
vegetated over unvegetated bottoms (Briggs and O'Connor 1971; Sogard et
al. 1992; Dorf 1994; Dorf and Powell 1997). These preferred, vegetated
habitats are reported to range from primarily eelgrass beds (Goode 1887;
Grover 1982; Orth and Heck 1980; Heck et al. 1989; Sogard et
al. 1992; Szedlmayer and Able 1996) or a mix of eelgrass and algal
associates [i.e., sea lettuce (Ulva lactuca), Enteromorpha sp.,
and Polysiphonia] (Briggs and O'Connor 1971), to beds of mostly Ulva (Nichols
and Breder 1926; Sogard and Able 1991). In the Great Bay - Mullica River
Estuary (New Jersey), early YOY preferred sea lettuce over eelgrass habitats
(Sogard 1989). Sogard (1989) reported that juvenile tautog also readily
used artificial sea grass as habitat. For YOY and 1-yr-old juveniles,
empty oyster and clam shells (Bigelow and Schroeder 1953) and shell and
sponge (Szedlmayer and Able 1996) have been reported to be used as habitat.
Dixon (1994) added small boulders as a preferred habitat type for juveniles.
The main habitat requirement and distribution factor, however, for juvenile
tautog (less than 25 cm) is the availability of cover (i.e., any
object that an individual can remain alongside, within, or under) (Olla et
al. 1974, 1975). Along these lines, Dixon (1994) noted that vertical
relief is an important attribute of juvenile habitat. Smith (1907) reported
that young fish were abundant around wharves in North Carolina; Able
and Fahay (1998) reported the same for New York Harbor. Larger juvenile
tautog are closely associated with hard- surface, reef-like habitats
(Olla et al. 1979). Zawacki (1971) reported that 15-20 cm fish
were common around a 1-mo-old, automobile-tire artificial reef in Shinnecock
Inlet, Long Island. Adams (1993) reported only juveniles greater than
12 cm total length (TL) recruited to southern, offshore reefs.
In Narragansett Bay, YOY tautog appear from late June through August
(Dorf 1994; Dorf and Powell 1997). During winter, juveniles remain inshore
(Cooper 1964; Stolgitis 1970; Olla et al. 1974). At this time,
especially when water temperatures are 5°C or less, YOY and 1-yr-old
tautog are found in discarded beverage cans and bottles, eelpots, empty
oyster and clam shells, and in crevices of vertical structures; they
are also found on their sides in nearby depressions in sediments, either
covered with a few millimeters of sand and silt (except head and gill
area) or exposed on the sediment surface (Bigelow and Schroeder 1953;
Cooper 1964, 1966; Olla et al. 1974, 1978, 1979, 1980). Juvenile
tautog (3-7 cm TL) were found to select a narrow range of hole sizes
(i.e., about 3-4.5 cm in diameter) in structured habitats, and
to occur usually near the bottom of any object or structure (Dixon 1994).
Dorf (1994) suggested that sheltering habitat could be a limiting distribution
factor in Narragansett Bay for juveniles less than 2 yr old. Although
the food-producing capability could be surmised as one of the reasons
that older juveniles prefer reef-like habitats, Dixon (1994) found that
neither the availability of attached food (e.g., small mussels)
nor the presence of small but not highly active predators (e.g.,
toadfish and sculpins) had much effect on habitat use.
Olla et al. (1979) reported that young tautog (less than 25 cm)
showed an affinity to particular shelters, establishing homesites from
which they ranged only a few meters during the day, and to which they
returned at night. They also suggested that use of such homesites by
some juveniles is seasonal. They observed a perennial homesite -- a basin
wall -- which was utilized year-round by part of the juvenile population.
At the onset of winter, other young tautog that had possibly dispersed
to other nearby summer sites, such as eelgrass or algae beds, rejoined
the juvenile colony at the wall to overwinter. In the spring, some of
this population again left the wall for other shelter.
The mouths of estuaries or inlets may be especially important habitat
areas for juvenile and adult tautog. Briggs (1975) reported that tautog
was the most frequently collected fish in traps used on the Kismet artificial
reef just inside Great South Bay, Long Island. The species ranked only
fourth in abundance on an artificial reef less than 5 km offshore.
Adults
Adult (i.e., mature) tautog (generally greater than 25 cm TL)
have the same basic habitat requirements as larger juveniles and are
found in vegetation, rocks, natural and artificial reefs, pilings, jetties
and groins, mussel and oyster beds, shipwrecks, submerged trees, logs
and timbers, and similar complexly structured coastal habitats [e.g.,
see Smith (1907) and Hildebrand and Schroeder (1928)]. The fish are extremely
local, so much so that when fishing for them "a few feet one way or the
other may mean the difference between success and failure" (Bigelow and
Schroeder 1953). Adams (1993) observed that the species preferred the
crest and outer edges of coastal reef habitats.
During summer, adults can be found inshore, co-existing with younger
fish. Olla et al. (1974) reported adults ranged up to 500 m away
from their homesite during the day, but generally returned to the same
general shelter area at night. This established a local population during
the summer.
In late fall, when water temperatures fall below 11°C, there is
an overall migration to perennial offshore areas with rugged topography
in waters 25-45 m deep (Cooper 1966). Individual tautog do not appear
to return annually to the same sites within an offshore area to overwinter
(Olla et al. 1979). On the other hand, some adults were found
to overwinter inshore, especially in the north.
The seasonal migrations of tautog do not seem to involve great distances
[e.g., Briggs (1974) reported that fish from southern Long Island
bays winter in deeper coastal waters off northern New Jersey]. Other
adult tautog in winter were observed in less than 10 m of water in eastern
Long Island Sound (Zawacki and Briggs 1976; Auster 1989), in 10 m of
water at an artificial reef 2.4 km off Delaware (Eklund and Targett 1991),
and in deeper areas of Chesapeake Bay (Hostetter and Munroe 1993). Eklund
and Targett (1990) noted populations in 25-35 m of water 22-37 km offshore
of Maryland and northern Virginia, and Hostetter and Munroe (1993) observed
populations in 10-75 m of water to 65 km offshore of southern Virginia.
At an artificial reef in about 20 m of water 15 km offshore of southern
Virginia, Adams (1993) observed that the largest tautog (greater than
75 cm in length) occurred in February when temperatures were about 6°C.
He considered tautog to be a "core resident species" on this reef, and
to be active year-round.
At very low temperatures, tautog enter a torpor-like state (Cooper 1966;
Briggs 1977). Curran (1992) reported that cunner go into true torpor
or hibernation during winter, and suggested tautog do likewise, at least
in the northern part of their range. She believed the ability of cunner
and probably tautog to hibernate may be the key to these two species,
of a basically tropical family, being able to tolerate cold water and
inhabit cool-temperate waters year-round.
DETERMINANTS
OF DISTRIBUTION
Juveniles
Olla et al. (1979) suggested that the distribution of habitat
use by juvenile tautog was somewhat seasonal and temperature mediated.
Juveniles seem to have a perennial site which they use during the winter.
In spring, some of the wintering groups dispersed to summer sites. Olla et
al. (1979) believed this springtime dispersal was possibly due to
increased aggression in the population or to other factors which made
the perennial winter site suboptimal. If a summer habitat became suboptimal,
because of elevated temperatures or inadequate vegetation, juvenile tautog
moved to other perennial summer sites.
Young tautog have strong adherence to homesites (Able and Fahay 1998),
but if shelter becomes suboptimal, they will move (Olla et al. 1979).
They will also move if a more attractive habitat is found. In the Great
Bay - Mullica River Estuary, Sogard (1989) found that in summer juveniles
moved from naturally vegetated homesites to newly planted, artificial
seagrass beds. The factors that made the artificial seagrass beds more
attractive are not known.
Adults
Adults also show an affinity to homesites, though this is not as strong
as for juveniles. Adults have been observed to leave a site readily if
suboptimal conditions develop. At an artificial reef in Delaware Bay,
tautog which were common in early summer were absent later that year,
coincident with a blue mussel (Mytilus edulis) kill, probably
caused by high water temperatures (personal observation by senior author).
Also, at an artificial reef 15 km off Virginia, Adams (1993) noted a
decrease in abundance of large tautog when water temperatures were above
20°C for an extended time. He suggested that large fish leave areas
with uncomfortably high temperatures for areas with cooler, deeper waters.
The adult fall offshore migration is triggered by bottom water temperature
dropping below 10°C (Cooper 1966; Olla et al. 1974; Lynch
1994). The spring inshore migration is associated with an increase in
bottom water temperatures to 11°C or above (Chenoweth 1963; Cooper
1966; Stolgitis 1970; Olla et al. 1974; Briggs 1977; Olla et
al. 1979). In laboratory experiments, Olla et al. (1980) confirmed
temperature, not changing photoperiod, as the leading factor in fall
migratory movements and distributions.
HABITAT NEEDS
Tautog are specifically associated with complexly structured habitats
in all post-larval stages of their life. As juveniles, these habitats
include submerged vegetation, shellfish beds, and three-dimensional objects
or structures with appropriately-sized crevices and holes for shelter.
(See the "Differential Distribution" section
in this chapter). As the fish grow, larger complex structures are needed
for shelter, and hard substrates are usually required to support the
epibenthic or encrusting invertebrates upon which the fish generally
feed. (See the "Feeding and Diet" chapter).
South of Long Island, beyond where rocks and boulders were deposited
during previous glacial periods, there are few natural rock outcroppings
in coastal marine waters to provide the "reef" habitat that tautog require,
although shellfish beds in euryhaline parts of estuaries serve as habitat
(Arve 1960). In this area, the man-made "reef" habitat created by coastal
jetties, groins, pilings, accidental shipwrecks, and intentional deposition
of solid material as artificial reefs is undoubtedly important to the
distribution of the species.
The availability of new macroalgal growth as cover in the late spring
to early summer period can be critical to settlement and survival of
post-larvae (Dorf 1994; Dorf and Powell 1997). This is especially true
for areas that do not support extensive eelgrass beds or complexly structured
habitats. Although macroalgae is normally degraded or swept into dense
beds later in the season, a healthy, new-growth, spring-summer macroalgal
community can be important to the initial survival of juvenile tautog
(Dorf 1994; Dorf and Powell 1997). Sogard and Able (1992) reported that
juvenile tautog in New Jersey appeared to prefer habitat where sea lettuce
(Ulva) was present. Ulva can have negative effects on certain
taxa or species, and the preference or tolerance of Ulva by tautog
can give it an advantage with competitors or predators (Dorf 1994). In
contrast to studies that report the importance of vegetation, Dixon (1994)
reported that YOY tautog preferred small boulders as habitat over cobbles,
vegetation, and other structure-based habitat options in an aquarium
study.
REPRODUCTION
SEXUALITY
In contrast to most labrids, tautog are not protogynous hermaphrodites
(Olla et al. 1981). They are heterosexual, but two different morphological
males are thought to be present in the population. One type of male is
dimorphic with a more pronounced mandible than found on the female, and
the other type is nondimorphic and resembles the female (Olla and Samet
1977; Hostetter and Munroe 1993). It has been suggested that the nondimorphic
male may be: 1) a sexual stage in the life of the tautog, 2) an indicator
of hermaphrodism, 3) a means to increase spawning opportunities, or 4)
coincident with a different reproductive behavior than the dimorphic
male (Hostetter and Munroe 1993).
FECUNDITY
Chenoweth (1963) found that tautog that were 21-68 cm in fork length
(FL), weighed 170-5207 g, and were between 3 and 20 yr old, contained
5000 to 637,500 mature eggs. He found that the number of eggs (Y) were
related to fork length (X) in millimeters by the regression: Y = -6.00307
+ 3.0960(X). The number of eggs (Y) was also related to weight (Z) in
grams by the regression: Y = 0.31492 + 1.07993(Z). The number of eggs
produced per unit of weight per ovary reaches a maximum in 7-9 yr old
fish which are about 34-39 cm TL (Cooper 1967). Thereafter, egg production
stabilizes until an age of about 16 yr, after which it declines (Chenoweth
1963; Cooper 1967).
SPAWNING
The ratio of the mass of gonadal tissue to the mass of all body tissues
[i.e., the gonadal-somatic index (GSI)] can be an indicator of
a fish's reproductive state, with the highest indices just prior to spawning,
and the lowest indices just after spawning. The GSIs for female and male
tautog off the coast of Maryland and Virginia were reported to peak between
April and June (Eklund and Targett 1991; Hostetter and Munroe 1993).
Spawning begins when water temperatures reach 9°C or above, generally
peaks about June, and continues throughout summer (Kuntz and Radcliffe
1918; Nichols and Breder 1926; Perlmutter 1939; Bigelow and Schroeder
1953; Wheatland 1956; Chenoweth 1963; Cooper 1964; Colton et al. 1979;
Eklund and Targett 1990; Monteleone 1992; Sogard et al. 1992;
Hostetter and Munroe 1993; Malchoff 1993). Sogard et al. (1992)
used NEFSC-MARMAP larval distribution (Berrien and Sibunka, in press)
and other data to show spawning began in May-June south of New York,
and reached its peak in June-
July off Southern New England. Dorf (1994) reported finding hatching
eggs in late May to late July in Narragansett Bay. Sandine (1984) reported
the occurrence of tautog eggs in Barnegat Bay (New Jersey) from March
to August, and questionably in October.
COURTSHIP
Tautog spawn in heterosexual pairs or in a group with a single female
being active simultaneously with several males (Olla and Samet 1977;
Dixon 1997). The mode of spawning is reported to depend on the number
of mates available for the female, the presence of a male-dominance hierarchy,
and environmental factors such as availability of shelter and food (Olla
and Samet 1977). In laboratory studies with two active males (one of
which was dominant) one female, and one shelter, mating occurred only
between the dominant male and the female (Olla and Samet 1977). Group
spawning was observed when there was either an increase in the number
of males, a lack of a dominant male, an increase in the male:shelter
ratio, an inability to control a territory, or an elevated temperature
(Olla and Samet 1977; Olla et al. 1981).
Several weeks before spawning, male aggressiveness toward females noticeably
decreased and was replaced by "rushing" (i.e., males quickly approached
females then veered off) (Olla and Samet 1977). Coincident with this
change in male behavior, females increased their girth with enlargement
of their ovaries.
In laboratory studies, courtship activity prior to spawning was observed
to continue for several hours. In paired spawning, the male rushed the
female frequently (Olla and Samet 1977), and male pigment bars darkened
while those of the female lightened (Bridges and Fahay 1968). Physical
contact between the sexes, including nuzzling and rubbing flanks, was
observed (Bridges and Fahay 1968). Courtship culminated with the paired
fish moving rapidly together within 1 m of the water's surface. The fish
then turned toward each other, arched their bodies, and released their
gametes near, or as they broke, the water surface (Olla and Samet 1977).
Spawning usually occurred in the afternoon, with as many as three spawnings
daily (Olla and Samet 1977, 1978). Spawning often continued into evening
in the wild (Ferraro 1980).
Olla and Samet (1977) observed slight differences in courtship during
group spawnings. The males did not rush the female as frequently, and "contact-clustering
behavior" occurred (i.e., two or more males clustered near the
female and contacted her with their flanks).
DEVELOPMENT
OVA
The development of the ova was described by Chenoweth (1963):
The ova arise from the germinal epithelium that lines
the interior wall of the ovary and fills a large part of the organ
through convolutions. At first the developing ova are opaque, relatively
hard, and average 0.36 mm in diameter. As development proceeds the
ovary enlarges and transparent, soft, mature ova appear interspersed
with the immature ova. These ova average 0.79 mm in diameter.
There are corresponding changes in the appearance of the ovary as the eggs
mature.
EGGS AND EMBRYOS
Spawned tautog eggs are about 1 mm in size, highly transparent, and spherical
in form (Herman 1963). The egg membrane is thin but tough and the yolk sphere
does not contain an oil globule (Kuntz and Radcliffe 1918). Egg size varies
from year to year, with diameters of 0.70-1.18 mm being reported (Nichols and
Breder 1926; Richards 1959; Chenoweth 1963; Lebida 1969). Egg size also decreases
with an increase in water temperature, and as the spawning season progresses
(Williams 1967). Therefore, it has been suggested that annual variation in
egg diameter reflects differences in sampling times. Eggs that were reported
in the literature with variable diameters could be explained by their being
collected at different times during the spawning season in different years
(Auster 1989).
Incubation of fertilized eggs at temperatures of about 20-22°C in the
laboratory was reported to take 42-48 hr to hatching (Kuntz and Radcliffe 1918;
Merriman and Sclar 1952; Perry 1994), but at 14.2-16.8°C, it took about
81 hr (Perry 1994). Kuntz and Radcliffe (1918) and Fahay (1983) described embryonic
development.
D. Perry (pers. comm.4)
reports that the proportion of normally developing, viable embryos among all
tautog embryos collected in central Long Island Sound near New Haven, Connecticut,
declined as the spawning season progressed. This decline had no significant
locality variance, nor appeared to be related to differences in habitat quality.
This progressive decline was also noted in laboratory spawned eggs and reared
embryos (Perry 1994). Olla and Samet (1978) found higher ambient temperatures
affected normal development of tautog embryos.
LARVAE
Newly hatched embryos, or yolk-sac larvae, are approximately 2.2 mm in length,
but can be slightly less. [Some variance in the lengths of specimens reported
in the literature and reported here can be the results of preservation which
usually causes some shrinkage. This shrinkage can vary with developmental stage
(e.g., degree of ossification).] The head is slightly deflected and
the yolk sac is relatively large, elliptically ovate, and unpigmented. The
vent is located about mid-length of the body. The depth of either dorsal or
ventral fin fold is less than the depth of the body just posterior to the vent.
The fin folds and the posterior caudal region of the body remain free of pigment.
One day after hatching at temperatures of 20-22°C, tautog larvae are about
2.0-3.0 mm in length. The yolk sac is greatly reduced and the head is no longer
deflected. The chromatophores increase in size and show well developed pigment
processes, but are fewer in number than in the newly hatched larvae; individual
pigment cells merge to form large chromatophores. The larvae have a distinct
blackish color (Able and Fahay 1998).
Four days after hatching is a critical period for this species. The yolk sac
has been absorbed and the mouth has been formed and is functional (Bigelow
and Schroeder 1953). These post-yolk-sac larvae, which are now 3.2-3.5 mm in
length, must begin planktonic feeding. Black chromatophores are uniformly distributed
over the dorsal and lateral aspects of the body, but the posterior caudal region
is free of pigment.
Larvae which are 5.0 mm in length show a relatively greater increase in body
depth and thickness than in length. The distribution of pigment remains essentially
the same as 4 days after hatching; however, the chromatophores increase in
size and number. Larvae which are 10 mm in length display well differentiated
dorsal, anal, and caudal fins. Pigment distribution remain the same as in earlier
ontogenetic stages of development, but the number of chromatophores and quantity
of pigment increase.
Malchoff (1993) found that larvae in the New York Bight ranged from about
2.0 to 7.0 mm TL. From his time-series collections, he developed a larval age-length
relationship: A = -1.877 + 8.535 L, where A = age in days, and L = length in
millimeters.
The mean duration of the larval phase of tautog was reported to be 25.4 ± 3.4
(SD) days for an undefined area of the Northwest Atlantic (Victor 1986). Sogard et
al. (1992) and Malchoff (1993), however, reported the larval phase to be
only about 20 days in the New York Bight and Great Bay - Mullica River Estuary.
Dorf (1994) and Dorf and Powell (1997) reported the same period for Narragansett
Bay, but it can be less (Schroedinger and Epifanio 1997).
JUVENILES
After the larval phase, settlement to benthic habitats occurs and an epibenthic
lifestyle begins. The transition between the larval pelagic and juvenile demersal
stages is evident in the otolith sagitta; inner increments are higher in contrast,
darker in appearance, and more circular than those outside the transition area
(Sogard et al. 1992).
At 30 mm TL, the fish are generally considered to be juveniles and show the
general morphological characteristics of adults, except for their often greenish
background color. The black chromatophores form heavily pigmented areas which
give the body a transversely banded appearance as well (Kuntz and Radcliffe
1918).
The fish lay down their first visible bone annulus in the spring of the following
year, usually in May. For example, in the opercular bone, this is evident as
a sharp transition from a translucent to an opaque zone. Annuli of this type
are formed on bony structures each spring (Cooper 1967).
ADULTS
Most male tautog mature by age 3, females by age 4 (Chenoweth 1963; Cooper
1967; Stolgitis 1970; Briggs 1977; Hostetter and Munroe 1993). Chenoweth (1963)
observed that 55% of males were already mature by age 2, and 90% were mature
by age 3, in Rhode Island. This latter age corresponds to a tautog length of
about 26 cm, as determined by Hostetter and Munroe (1993). The latter authors
found that few females were mature by age 2, but 80% were mature by age 3.
Precocious females, approximately 2 yr old, were observed in New York waters
(Olla and Samet 1977) and in Mt. Hope Bay, (Massachusetts) (Hostetter and Munroe
1993).
The influence of heavy fishing mortality on the genetic selection process
for sexual maturity is unknown at present; this pressure can favor the portion
of the population with genes for early maturation and spawning, although with
lower gamete production.
AGE AND GROWTH
LARVAE
Malchoff (1993) reported larval growth rates for this species to be 0.30 mm/day
in the New York Bight. This rate is lower than a mean 0.75 mm/day, back-calculated
rate reported for Narragansett Bay fish by Dorf (1994). Martin (1993) reported
that larval growth rates varied with latitude and were directly related to
water temperature.
JUVENILES
Sogard et al. (1992) used length-frequency progressions, otolith age
- fish size comparisons, and direct measurements of growth in cage experiments
to determine that YOY tautog grew at an average rate of about 0.5 mm/day during
the summer in the Great Bay - Mullica River Estuary. The same rate was found
in Narragansett Bay (Dorf 1994). In the Great Bay - Mullica River Estuary study,
mean growth rates varied slightly among different methods of estimation and
with habitat types where the fish were collected. Sogard et al. (1992)
also found minor growth also occurred in other seasons. Martin (1993) reported
that YOY grew up to 0.7 mm/day when fed mysids in the laboratory.
During the 2-3 yr of the juvenile phase of this species, annual growth is
usually rapid (Cooper 1967). Annual growth is reported to increase progressively
from northern to southern regions (Martin 1993). For example, in Narragansett
Bay, Cooper (1967) reported first-year growth in tautog to be 60-62 mm. This
was less than half the first-year growth of 134-146 mm by YOY tautog in southern
Virginia waters (Hostetter and Munroe 1993). Tracy (1910), however, reported
first-year growth to range between 72 and 288 mm in Rhode Island; the larger
value is obviously questionable (Fritzsche 1978).
In Narragansett Bay and the Great Bay - Mullica River Estuary, the greatest
annual growth increment, 200%, occurred during the second year for both sexes
(Cooper 1967; Sogard et al. 1992). In southern Virginia waters, the
greatest annual growth increment occurred in the first year, and the second
year's growth was only a 30% increase (Hostetter and Munroe 1993). It has been
hypothesized that the faster initial growth rates in southern waters reflect
the longer duration of warmer water temperatures, which may provide optimal
conditions for tautog growth during the first juvenile year, but for some reason
not the second year (Sogard et al. 1992; Hostetter and Munroe 1993).
Sogard (1992) observed that the mean seasonal growth in length of juvenile
tautog in New Jersey was higher in vegetation, but growth in mean weight was
higher on bare sand. Growth rates were higher in sea lettuce beds than in eelgrass
beds, and seemed to be directly related to prey density and not shelter type.
ADULTS
Although Cooper (1967) noted that Rhode Island females attained a slightly
greater mean length than males (62 mm compared to 60 mm) during their first
juvenile year, by age 3, as they became adults, males had faster annual growth
rates. At age 7 in Rhode Island, the male tautog's mean length was 348 mm,
while the female's mean length was 301 mm. Faster adult male annual growth
rates were also found by Simpson (1989) in Long Island Sound. In southern Virginia
waters, males also grew faster than females at all ages (i.e., K = 0.090
for males compared to K = 0.085 for females5)
(Hostetter and Munroe 1993).
Hostetter and Munroe (1993) found that tautog attained a relatively large
size slowly, and that their growth varied seasonally. These authors reported
that in southern Virginia's coastal waters, maximum somatic growth occurred
after spawning, from July to December. Slower growth occurred from January
to March because of decreased feeding associated with cool water temperatures
and the fish's associated torpor condition in colder northern waters. In northern
waters, this period of slower winter growth is also probably longer because
these waters cool sooner and remain cold longer. In southern Virginia waters,
the least somatic growth in adults occurred from March to June during gonadal
maturation and spawning.
Mean annual growth rates in length were similar for tautog in northern and
southern waters until about age 13, then growth rates decreased more rapidly
in northern waters. For males in Rhode Island, annual growth increments decreased
to less than 12 mm after age 12, and further declined to 2-4 mm after age 20.
For females in Rhode Island, annual growth increments decreased to less than
11 mm after age 13, and to 3-4 mm after age 17 (Cooper 1967). In southern Virginia
waters, increments in annual growth declined after age 13, but growth rates
were nearly double those of tautog in northern waters (Hostetter and Munroe
1993). The von Bertalanffy growth equation for tautog from both areas support
this finding: in northern waters, L= 506 mm for females and 664 mm for males
(Cooper 1966), compared with L= 733 mm for females and 732 mm for males in
southern waters (Hostetter and Munroe 1993).
Several studies reported variable length-weight relationships for whole and
eviscerated tautog from Narragansett Bay to southern Virginia waters (Table
1).
Tautog are a relatively long-lived fish, with the oldest fish examined by
the scientific community estimated to be a 34-yr-old male (Cooper 1967). A
91-cm (36.5-inch), 10.1-kg (22.5-lb) fish, caught off New York in 1876 and
reported by Goode (1887), could have been older. Hostetter and Munroe (1993)
reported that the world's record weight for the species is 10.9 kg (24 lb)
for an 81.9-cm fish caught recently off Virginia; the authors estimated the
fish to be about 30 yr old. Simpson (1989) suggested that males can live more
than 30 yr, and females about 25 yr.
FEEDING AND DIET
FEEDING BY LIFE STAGE
Larvae
No specific data were found; larval tautog probably feed on small motile crustaceans
in the water column such as small copepods [which were part of the diet of
small juvenile (30-40 mm) tautog reported by Grover (1982) and Dorf (1994)]
and other larval fish. D. Perry (pers. comm.6)
reported some success in getting larvae to eat dried food.
Juveniles
The diets of juvenile tautog are discussed below by arranging the available
information from north to south to best fit the gradual zoogeographical shifts
in diet that occur on this axis. Dorf (1994) found the diets of juveniles in
Narragansett Bay to consist of amphipods and copepods, with the copepods being
mostly harpacticoids. Richards (1963) notes that YOY and 1-yr-old juvenile
tautog in a sand-shell area of Long Island Sound ate pycnogonids (sea spiders),
razor clams (Ensis directus), and decapod and amphipod crustaceans.
Nichols and Breder (1926) also report "seaweed" as part of the diet of juvenile
tautog in this area.
Grover (1982) found that both caprellid and gammarid amphipods and small copepods
constituted 95% of the diet of YOY (31-71 mm TL) tautog in southern Long Island
waters. Frequency of occurrence (FO) in the digestive tract was 98% for amphipods,
94% for copepods, 29% for polychaetes, and 25% for isopods. This diet indicated
that juvenile tautog forage on benthic, as well as planktonic, prey. As they
grow, juvenile tautog rely less on planktonic food resources and feed primarily
on benthic prey, although the prey remain primarily crustaceans, such as decapods.
The general form and location of the tautog's mouth suggest that benthic organisms
would be a primary component of the adult's diet (Grover 1982). In Great South
Bay (New York), 2-3 yr old tautog (105-206 mm) did not feed mostly on crustaceans
throughout the year, but on blue mussels (Olla et al. 1975). This diet
difference is partially consistent with the laboratory prey selection findings
reported by Lankford et al. (1995) that juvenile tautog gradually shift
their diet from small crustaceans (e.g., amphipods) to small mussels
as the fish grow to and beyond a length of about 120 mm. Mussels that were
about half of their maximum size were primarily eaten.
In the Great Bay - Mullica River Estuary, Sogard (1992) found that copepods
were the preferred prey, with a 78% FO, in 31-85 mm YOY fish. Amphipods were
a close second in importance (FO = 75%), followed by other crustaceans (FO
= 40%), polychaetes (FO = 5%), and mollusks (FO = 5%). Festa (1979) reported
that isopods (Idotea sp. and Erichsonnella sp.), xanthid (mud)
crabs, and (to a lesser degree) several species of amphipods were the prey
of 60-160 mm tautog from Great Egg Harbor (New Jersey). He also reported that
larger subadult tautog, greater than 200 mm in length, ate predominantly xanthid
crabs.
In eelgrass beds of Chesapeake Bay, juvenile tautog (90-170 mm) fed on penaeid
shrimp, blue crabs (Callinectes sapidus), isopods, grass shrimp (Paleomontes sp.),
and detritus (Orth and Heck 1980).
Lindquist et al. (1985) reported that mytilid mollusks and gammarid
and caprellid amphipods dominated the diets of 22 tautog (95-270 mm) examined
from rock jetties near Wrightsville Beach (North Carolina); they also noted
the occurrence of algae in the stomachs. Troutman (1982) reported that the
dominant prey in these North Carolinian diets varied seasonally, and included
a venerid clam. MacKenzie (1977) also reported juvenile tautog to prey on a
juvenile venerid, the northern quahog (Mercenariamercenaria).
Adults
Adult tautog are durophagous and feed chiefly on blue mussels and other shellfish
throughout the year. Barnacles (Balanus sp.), brachyuran crabs, hermit
crabs, sand dollars, (bay?) scallops, amphipods, decapod shrimp, isopods, lobsters,
and probably nereid polychaetes were reported by Bigelow and Schroeder (1953)
to be part of the tautog's diet. Consistent with this broad summary, Osburn
(1921) reported that "tautog feed on bryozoa along with other hard shelled
organisms to which it was attached," and Verrill (1873) reported that tautog
fed on benthic tunicates. Bleakney (1963) reported tautog from Nova Scotia
to feed not on blue mussels, but on horse mussels (Modiolus modiolus)
and periwinkles (Littorinalittorea). Linton (1899) and Scott and Scott
(1988) also listed gastropods in the tautog diet. Auster (1989) included the
softshell (clam) (Mya arenaria) as a diet item.
South of Cape Cod, examination of digestive tracts of tautog in New York estuarine
waters showed that 70% of the fish contained 78-100% mussels by volume (Olla et
al. 1974). Steimle and Ogren (1982), however, found that Atlantic rock
crabs (Cancer irroratus) constituted greater than 78% by volume
of the diet of tautog collected from a coastal New York artificial reef. They
also found that sand dollars (Echinarachniusparma) (38%), Atlantic rock
crabs (28%), and blue mussels (9%) dominated the tautog diet on a volumetric
basis at a northern New Jersey artificial reef. Festa (1979) reported that
xanthid crabs dominated the diet of small adults (about 300 mm) in Great Egg
Harbor Estuary.
Unpublished data collected by the senior author showed that besides blue mussels
(54%), Metridium anemones (11%), Atlantic rock crabs (5%), and razor
clams (4%) dominated overall definable prey in 358 tautog stomachs, by volume,
in a 1990-94 Delaware Bay artificial reef study; these fish ranged from 110
to 580 mm, with a mean length of 320 mm. The proportional composition of the
diet varied among years (e.g., the mussels varied from 13 to 87% of
total diet volume). Various small mollusks, barnacles, decapod crabs, and other
crustaceans of suitable size were reported also to be the food of this species
in the Chesapeake Bay area (Hildebrand and Schroeder 1928).
Richards (1992) confirmed Smith's (1907) note that tautog will eat commercially
important Jonah crabs (Cancer borealis) and small American lobsters
(Homarus americanus). Predation on these species occurred, however,
when they were without shelter in aquaria.
Chao (1973) reports that the cunner lacks a well defined stomach, which is
a characteristic of most labrids, including tautog. The tautog's gastro-intestinal
tract is semi-transparent and remarkably thin for containing the broken shells
of its prey as it moves through to evacuation.
FEEDING BEHAVIOR
Tautog feed throughout the daytime. Beginning soon after sunrise, tautog were
reported to leave their shelters to forage for food, which involved scan-and-pick
feeding (Briggs 1969b; Olla et al. 1975). This activity sometimes took
the adults up to 500 m from their homesites (Wicklund 1966; Olla et al.1974).
Bigelow and Schroeder (1953) observed that tautog followed the flood tide up
above low water levels, around ledges, to prey on mussels in the intertidal
zone, and returned to deeper water during the ebb tide. Feeding continued to
evening twilight (Olla et al. 1974). Olla et al. (1974) reported
that tautog required about 8 hr to process and evacuate food.
In laboratory studies, Olla et al. (1974) observed that tautog grasped
mussels with their anterior teeth and tore them from their attached substrate
with a lateral shaking of the head. Small prey were swallowed whole (Bigelow
and Schroeder 1953), while larger, hard-shelled ones were crushed by pharyngeal
teeth before swallowing (Bigelow and Schroeder 1953; Olla et al. 1974).
The anterior teeth were not involved in the crushing process (Olla et al. 1974,
Liem and Sanderson 1986). It was observed that the tautog's mouth can accommodate
larger clumps of mussels than the pharyngeal teeth can process efficiently.
In this case, the fish ingests and egests the clump from its mouth, separating
it in the process into smaller, crushable sizes. The feeding and mastication
methods of this species are specialized and typical of labrids and cichlids
only (Liem and Sanderson 1986).
In the southern part of its range (i.e., below New Jersey), the blue
mussel, a dominant prey in colder waters, is at its warm-temperature limits
and has wide variability in recruitment and abundance (Foster et al. 1994).
Without strong periodic recruitment, mussel populations in these areas can
be preyed upon by tautog to near extirpation, as reported for Virginia (Chee
1977; Chesapeake Executive Council 1994). This near extirpation creates a change
in the prey field available to tautog in this habitat, and can cause at least
some of the tautog population to seek alternate or better foraging areas. This
change in prey field can be a factor in the local distribution of the population.
(See the "Adults" subsection, "Differential
Distribution" section of the "Distribution and
Habitat" chapter.)
FACTORS AFFECTING FEEDING
Tautog find prey visually and were reported not to feed at night (Olla et
al. 1974; Deacutis 1982). Neither do they actively feed in northern waters
during the coldest part of the year (Cooper 1966; Curran 1992). In Narragansett
Bay, Cooper (1966) observed that of 15 tautog, 13 had shrunken and emptied
digestive tracts in winter. The other two fish had remains of Atlantic rock
crabs in their stomachs. Curran (1992) found all feeding stopped in the closely
related cunner (and probably tautog) when water temperatures reached the
low levels that induce torpor and hibernation. Cunner, at least, survived
up to 6 mo without food, using glycogen, lipids, and proteins stored in their
livers (Curran 1992); tautog may do likewise.
In laboratory experiments, a decrease in feeding was observed with an increase
in water temperature above certain levels. Tautog which had been acclimated
to temperatures of 19°C and 21°C, respectively, decreased their ingestion
of food, when water temperatures were increased to 28.7-33.0°C over a certain
amount of days (Olla and Studholme 1975; McCormack 1976; Olla et al. 1978).
McCormack (1976) also reported that it required up to 7 days for field-collected
fish to begin feeding in laboratory aquaria. (For other environmental effects
on feeding, see the "Habitat Modification
and Loss" section of the "Natural and Human-Induced
Environmental Factors" chapter.)
Tautog were observed to vary, to some extent, their feeding in association
with their place in a group dominance hierarchy. In one laboratory study with
three fish, the dominant fish of the moment ate the greatest amount of food,
followed by the subordinate fish, in some order of rank (Olla et al. 1978).
In a related tank study, McCormack (1976), however, found no difference in
consumption of Atlantic surfclam (Spisula solidissima) meats between
the dominant and subordinate individuals of paired 160-280 mm tautog.
Food intake in tautog may decrease during spawning. Bridges and Fahay (1968)
reported possible courtship behavior of tautog in a laboratory study during
which no spawning occurred. In this study, 1 day prior to courtship behavior,
the male and female daily food (undefined) intake decreased from 40 to 1g.
During courtship behavior, which lasted 2 wk, the 300-mm female ceased eating,
but the 270-mm male increased food intake to 10 g/day. After courtship behavior
ceased, the female resumed eating, but less than 10 g/day. The time period
that this reduced level of feeding persisted, or if it changed, was not reported.
It is possible that some of this behavior was an artifact of the fish's confinement
in aquaria, as it has not been documented in the wild.
Deacutis (1982) found tautog did not have an acute sense of smell for detecting
prey, compared to red hake (Urophycis chuss), and were hesitant to explore
open bottom to find food they could not see.
BEHAVIOR
MIGRATION AND LOCAL MOVEMENTS
Many studies report limited, seasonal, onshore and offshore movements of this
species. (See the "Differential Distribution" section
of the "Distribution and Habitat" chapter.) Channels
may be important pathways of migration into coastal areas for spawning and
out again to deeper or offshore wintering areas (Cooper 1966). As noted previously,
Briggs (1977) reported that part of the southern Long Island tautog population
migrated to the area off northern New Jersey when water temperature declined
in the late fall; Nichols and Breder (1926) called this movement "heavy." At
a more local level, Merriman (1947) reported that small tautog do not move
with the high-tide levels into intertidal areas; this contrasts with Bigelow
and Schroeder's (1953) report of intertidal-associated movements of presumably
adults for feeding on tidally submerged mussels, as noted earlier.
SCHOOLING
Some degree of schooling is thought to occur as tau-tog are reported to congregate
during or just prior to spawning (Bigelow and Schroeder 1953; Cooper 1964;
Stolgitis 1970). Briggs (1969b) believed that tautog also congregate into some
type of a school before and during their movement to deeper water in the fall
off New York. This is supported by laboratory studies where tautog schooling
was observed when water temperatures declined to 6.3°C (Olla and Studholme
1975).
DAILY AND SEASONAL
ACTIVITY LEVELS
In laboratory studies, tautog showed a high level of activity (i.e.,
movement) within the first hour of daylight (Olla et al. 1978). This
activity decreased toward mid-morning, when the fish rested on the sand or
within shelters between bouts of activity (Frey 1963). Bigelow and Schroeder
(1953) noted that when tautog were not feeding, they gathered in holes or crevices,
being still or lying on their side, often grouped together, until a rising
tide stirred them to activity (for intertidal feeding, presumably). In field
investigations at night, Olla et al. (1974) found tautog to be inactive
in a shelter of some type, and in a state of such low responsiveness that individuals
were readily touched by divers. Curran (1992) suggested that this state represents
true sleep in cunner (and probably in tautog), and can be for energy conservation.
Tautog in northern waters overwinter in a torpid state within shelter (Nichols
and Breder 1926). Olla et al. (1974) observed small fish (less than
250 mm TL) in a torpid condition at water temperatures less than 6°C. Cooper
(1966) observed adult tautog being dormant, either lying on their sides or
in upright positions in crevices, at a water temperature of 7.5°C. In laboratory
studies, tautog activity decreased as temperatures declined to 5.2°C, then
tautog aggregated around shelters or burrowed under objects on the bottom (Olla et
al. 1980). At water temperatures of 2.1°C or below, activity ceased
and tautog remained in torpor. Curran (1992) considered this torpid state to
be true hibernation in cunner. Emergence from torpor occurred when temperature
increased to 4.0°C (Olla and Studholme 1975). In southern Virginia waters,
tautog were observed to remain active despite water temperatures as low as
7°C (Adams 1993). In laboratory studies, young and adult tautog responded
to an increase in temperature, from 19.1°C or 21.3°C to 28.7°C,
with a decrease in aggressiveness and activity, and an increase in the tendency
to aggregate (McCormack 1976; Olla et al. 1978). This lowered response
to stimuli at temperature extremes was supported by the sound-detection-threshold
experiments of Offutt (1971).
HIERARCHICAL DOMINANCE
In laboratory studies, a dominance hierarchy was observed among variably sized
pairs or a group of tautog (Olla and Studholme 1975; McCormack 1976; Olla et
al. 1978). In these studies, the larger, most aggressive or dominant fish
caused a less aggressive, usually smaller subordinate to assume a submissive
posture, tilting its dorsal surface toward the dominant fish at an angle. This
behavior usually occurred within 1-3 m of the dominant male. Field studies
have not confirmed these observations.
VOCALIZATION AND RESPONSE
TO SOUND
Fish (1954) found that tautog produced deep thumping, grunting, or barking
sounds, as well as sounds from crushing shells of their prey when feeding.
The nonfeeding sounds were heard in aquaria when fish were startled or exposed
to experimental electrical impulses, and were not thought to be produced commonly.
The air bladder was thought to be involved in this sound production. The purpose
of producing these sounds was unknown, but could be defensive. Parker (1910)
reported that tautog avoided sources of loud sounds. Offutt (1971) used conditioned,
cardiac responses to show that fish can respond to sound impulses.
COLOR MODULATION
Behre (1933) reported that small tautog responded to changes in light intensity
and color by changes in their body color or shading. Absence of light reduced
the intensity of body color, while high-intensity light enhanced the body's
dark coloration. Differently colored light and sequential combinations of colored
lights caused different responses in body coloration. These responses suggested
to Behre some ability by tautog to distinguish colors or shading. Regan et
al. (1982) reported that the photoreaction capacity of tautog was markedly
different from its close, but shorter-lived relative, the cunner. No reason
for this was suggested, however. McCormack (1976) reported that there were
body color or shading patterns evident during the interaction between dominant
and subordinate fish in an aquarium. Dominant fish often exhibited darker shading,
and submissive fish paled, during confrontations. Mallet (1972) reports color
changes from mottled-dark to pallid, gray-white as a stress response to hypoxia.
(For other stress responses, see the "Hydrographic" subsection, "Habitat
Modification and Loss" section of the "Natural
and Human-Induced Environmental Factors" chapter.)
MUTUALISM
Although many tropical and some temperate wrasses (e.g., European wrasse)
clean external parasites from larger fish of other species (Darwall et al. 1992),
this behavior has not been reported for any lifephase of tautog.
POPULATION STRUCTURE
SEX RATIO
Chenoweth (1963) noted that about 53% of the tautog he collected in Narragansett
Bay were female. In northern Virginia waters, however, the slightly skewed
sex ratio (0.86:1) was in favor of males (Eklund and Targett 1990). A significant
deviation from a 1:1 ratio was also observed in larger fish by Hostetter and
Munroe (1993), possibly because the older fish (greater than 18 yr old) were
predominantly males (Cooper 1967; Hostetter and Munroe 1993).
AGE COMPOSITION
As noted previously for Narragansett Bay, the oldest male fish found by Cooper
(1966) was estimated to be 34 yr, and the oldest female 22 yr. In southern
Virginia waters, the oldest male fish collected was estimated to be 25 yr,
and the oldest female 21, although the age of the oldest fish taken by rod
and reel from Virginia waters was estimated to be approximately 30 yr (Hostetter
and Munroe 1993).
In the early 1960s, 5- and 6-yr-old tautog were the most abundant age classes
in the Narragansett Bay population (Cooper 1967). The recent average age for
tautog in Virginia waters was 4 yr (Hostetter and Munroe 1993). In 1960s in
Narragansett Bay, 79% of the tautog were less than 10 yr old (Cooper 1967).
This population structure remained similar for Massachusetts to New York into
the mid-1990s (Lazar 1995). This age structure is also similar to southern
Virginia waters where 82% of the population was recently noted as being less
than 10 yr old (Hostetter and Munroe 1993).
NATURAL AND HUMAN-INDUCED ENVIRONMENTAL
FACTORS
PREDATORS
Perry (1994) reported from laboratory experiments that typical coastal copepods,
such as Acartia tonsa, can prey upon newly hatched, yolk-sac
tautog larvae. Bigelow and Schroeder (1953) reported the following fish species
were known to eat juvenile or adult tautog: smooth dogfish (Mustelus canis),
barndoor skate (Rajalaevis), red hake, sea raven (Hemitripterus americanus)
and goosefish (Lophius americanus). In analysis of 123 New York
striped bass (Morone saxatilis) stomachs, one tautog was found in a
greater-than-60-cm fish (Schaefer 1970). Striped bass were observed pursuing
young tautog in the same general area at a later time (Olla et al. 1974).
Zawacki (1971) reported that live small tautog were commonly used for surf
fishing for striped bass on Long Island, suggesting that the bass were familiar
with this prey. Schaefer (1960) reported that silver hake (Merlucciusbilinearis)
ate tautog in the "Mud Hole" off northern New Jersey. Wilk (1977) added bluefish
(Pomatomus saltatrix) to the list of tautog predators. Dixon (1994)
reported that toadfish (Opsanus tau) and longhorn sculpin (Myoxocephalus octodecemspinosus),
small cryptic predators common in coastal waters, as well as larger YOY bluefish,
preyed upon YOY tautog confined in aquaria. Other demersal piscivorous fish,
such as conger eel (Conger oceanicus), summer flounder (Paralichthys
dentatus), and various sharks, can be expected to prey upon small tautog.
Although American lobsters share the same habitat with tautog, and are active
at night when tautog are inactive, there are no reports of lobster predation
on tautog.
It is also likely that diving or stalking piscivorous birds, such as cormorants,
grebes, loons, herons, and egrets, also prey upon juvenile tautog in shallow
estuarine or coastal areas during daylight and the warm seasons when the fish
are outside their shelters (Whoriskey 1983). Bent (1919, 1922, 1923, 1926)
noted that small estuarine fish were eaten by these birds, with labrids being
specifically mentioned in the diets of the black-crowned night-heron (Nycticorax nycticorax)
and the double-crested cormorant (Phalacrocorax auritus). Nichols and
Breder (1926) noted a small tautog in the stomach of a red-throated loon (Gavia stellata).
Another observation of this predation can be recorded, with an approximately
12-15 cm tautog being observed in the beak and swallowed by a cormorant near
a rocky sea wall in Sandy Hook Bay (New Jersey) (personal observation by the
senior author in July 1992). The relative importance of this predation on population
structure, habitat use, and growth or mortality rates is unreported or unknown.
Humans as fishermen are also predators. Simpson (1989) estimated fishing mortality
for a Long Island Sound population to be F = 0.12, and to be about equal per
sex. He estimated natural adult mortality to be lower in males (M = 0.15) than
females (M = 0.20) for this population.
COMPETITORS
Competition between tautog on the one hand and cunner, sheepshead (Archosargusprobatocephalus),
and American lobster on the other hand has been suggested because all four
species use the same habitat and prey commonly on blue mussels, other small
mollusks, and crustaceans, and because the latter three species have ranges
that overlap that of tautog (Hildebrand and Schroeder 1928; Weiss 1970; Olla et
al. 1975). Botton and Ropes (1989) suggest that horseshoe crabs (Limulus
polyphemus) can consume large quantities of shellfish, including mussels,
and thus may also be a competitor for food. Sea stars, such as Asterias,
that prey heavily on mussels when they are available near the seabed, will
compete trophically with tautog for this food resource too (Richards 1963).
Other invertebrate predators of mussels, other shellfish, and barnacles can
be substantial tautog trophic competitors when present in abundance. Species
that eat copepods and decapod crustaceans should also be considered potential
competitors of juvenile tautog for this essential prey resource (Dorf 1994).
Competition with cunner was thought to be restricted to May and June, after
which cunner change their diet (Olla et al. 1975). In a laboratory study,
cunner seemed to be aggressively territorial in excluding similarly sized juvenile
tautog from certain habitats where they co-occur (McErlean 1963). The competition
between tautog and cunner can only seriously exist with juvenile tautog; larger
tautog use larger shelter and prey that are well beyond the range of cunner.
Dixon (1994) suggests, however, that YOY cunner and tautog have differing,
but overlapping, preferences in the location and use of their microhabitats
that can reduce competition.
DEFORMITIES AND ABNORMALITIES
Briggs (1966) reported on a mature female, estimated to be 9+ yr old, that
was pugheaded (i.e., exophthalmic eyes, shortened and broadened maxilla,
steep forehead, pushed-in snout, incomplete closure of jaws, extended mandibles).
Several 250-300 mm tautog collected by hook-and-line in Delaware Bay had notable
sigmoid vertical curvatures of their spines that compressed their body length
(personal observation by the senior author in June 1994). No ecological nor
environmental factor has been implicated in these deformities.
Tautog have been reported with small black spots within their muscle tissue.
Microscopic examination found that these spots were actually associated with
blood vessels, not the muscle tissue itself (I. Sunila, pers. comm.7).
These spots were found to be deposits of dark pigment surrounded by fibrosis.
This pigment originates from their high use of blue mussels as prey. These
spots are considered harmless to the fish and to consumers.
PATHOGENS, PARASITES,
AND BIOTOXINS
Tautog exposed to water in the apex of the New York Bight, near the then-active
dredged material and sewage sludge disposal sites, were found to have antigenic
responses to certain human enteric bacteria. These responses indicate that
tautog are immunologically responsive to exposure to sewage bacteria (Stolen et
al. 1983).
Finrot disease, thought to be caused by some microbial agent and anthropogenic
stress, was not found on tautog in the New York Bight, although the disease
was reported on 22 other common fish species in a study by Mahoney et al. (1973).
However, a subsequent study did report, without supporting details, this condition
on tautog (Murchelano and Ziskowski 1982).
Cooper (1964) observed that 10% of the tautog in his survey had scales that
were either pitted or regenerated. He considered this pitting or regeneration
to be due either to metacercaria of the trematode Cryptocotyl sp., or
to physical abrasions. Laird and Bullock (1969) found no positive evidence
of hematozoan parasites in New England and Canadian tautog. Cheung et al. (1979)
reported tautog to be infected lethally with a coccidian intestinal parasite.
Linton (1899) found immature trematode distomes to be encysted in abundance
on the skin, fins, and eyes of a tautog collected off Woods Hole (Massachusetts),
and also reported cestodes, nematodes, and trematodes to occur in cunner.
Perlmutter (1952) reported summer mortalities of "blackfish" along the ocean
side of Jones Beach, southern Long Island. These mortalities were speculated
to be related to a nearly concurrent, offshore bloom of the dinoflagellate Noctiluca sp.
A number of other fish species were reported affected at this time as well.
HABITAT MODIFICATION
AND LOSS
Structure
The dependence of tautog on specific coastal and estuarine habitats can make
any degradation or loss of these habitats a serious threat to the tautog resource
(Chesapeake Executive Council 1994). The loss of vegetated estuarine habitats
can impair the habitat value of estuaries as juvenile tautog (and other species)
nurseries. However, juvenile tautog have been observed using other habitats
and a variety of substrates, including artificial (e.g., pilings), that
provide shelter or concealment and access to food. (See the "Habitat
Needs" section of the "Distribution and Habitat" chapter.)
Rocky reef habitats used by tautog are very limited south of Long Island, and
shipwrecks and other subtidal, manmade material placed in this area have expanded
the distribution of this habitat into the common, open sandy habitats of the
Middle Atlantic coasts (Hastings 1978; Lindquist et al. 1985). Tautog
were collected around a sewer outfall under construction off southern Long
Island; their peak abundance was during October through December when bottom
water temperatures were 7-13°C (Briggs 1984). None of the tautog were found
with any obvious pathological conditions or other possible effects from this
physical habitat alteration and disturbance.
In the past, salvaging of certain metal shipwrecks or lowering of wreck heights
if they might be navigation hazards, reduced the value of shipwrecks as tautog
habitat. Before 1900, some wrecks off New Jersey were even dynamited to stun
and collect reef fish (Smith 1892). Recently, the use of certain, heavy, "rock-hopper-roller," bottom
trawl gear over many older, more fragile wrecks, lower-profile reefs, and mussel
beds (DiLernia 1993) also threatens habitat quality, including the destruction
of slow-growing colonies of the northern star coral (Astrangia sp.)
which provide biogenic reef habitat for tautog.
Reef habitats that lose structural height are more prone to siltation (Rothschild et
al. 1994) or burial by sediment movement. Briggs and O'Connor (1971)
report that tautog avoided estuarine areas that were sandfilled, as from
shoreline sand replenishment. The decline in the number, distribution, and
structure of oyster beds (Rothschild et al. 1994) is another threat
to the estuarine habitat needs of juvenile tautog and other species with
similar habitat needs (Chesapeake Executive Council 1994). The creation of
new "reef" habitat by the artificial reef programs of most states (McGurrin
1989) where the species occurs may be mitigating this habitat degradation
to some degree.
Hydrography
Auster (1989) reported that current velocity can affect small-scale spatial
distribution and can change foraging behavior in tautog and other species.
The high hydrographic energy caused by passage of a hurricane was reported
to disrupt juvenile tautog distributions in Narragansett Bay (Dorf 1994; Dorf
and Powell 1997).
Howell and Simpson (1994) reported that tautog are not tolerant of dissolved
oxygen (DO) concentrations below 2 mg/l, and prefer DO levels above 3 mg/l.
This intolerance is supported by the cardiac and preliminary respiration response
results of Mallet (1972) who showed that tautog are oxygen conformers under
normal conditions, and that there is no inflection threshold. He also showed
that these response rates increase with temperature and cadmium exposure, and
that hypoxia responses can then be detected at higher oxygen concentrations.
Mallet also observed several tautog stress responses to hypoxia. These responses
included a change in mottled-dark body color to a pallid white-gray, attempts
to jump out of experimental tanks, regurgitation of food, mucus secretion (especially
around the gills), and milt ejaculation in ripe males. Baldwin (1923) reported
some degree (undefined) of hypoxia "tolerance" at an unreported temperature
for this species.
In a fish kill off the northern New Jersey coast in 1968, possibly caused
by hypoxia, tautog were present in the area, but not found affected, although
its smaller relative, the cunner, were found dead or dying (Ogren and Chess
1969). Tautog mortalities, however, were reported by commercial fishers and
were observed on the beach during a major anoxia-hypoxia episode off New Jersey
in 1976 (Azarovitz et al. 1979).
Smith et al. (1980) reported that high water temperatures were incrementally
lethal in the laboratory for tautog gastrula-stage eggs: 10% mortality at 16.8°C
after 26 min, 50% mortality at 21.3°C after 50 min, and 90% mortality at
31.4°C after 55 min. These authors suggested that tautog eggs enduring
15 min of a once-through cooling system of a power plant, with a 10°C temperature
rise from a base temperature of 15°C, would suffer 10-50% mortality. Olla
and Samet (1978) reported that tautog eggs incubated above 20°C resulted
in embryos with anatomical deformities, and that those eggs incubated above
24°C resulted in increased mortalities. Normal embryonic development resumed
when water temperature decreased to 20°C or lower. Laurence (1973) reported
that tautog yolk-sac larvae ("prolarvae") may develop an energy deficit at
water temperatures above 19°C because of increased metabolic rate and use
of their yolk. This higher-than-normal rate of yolk use can result in smaller
larvae just when those larvae need to start feeding on plankton. He believed
these smaller larvae can be at a competitive disadvantage with larger siblings
and other plankton predators, and may be less successful in obtaining suitable
prey. This potential competitive disadvantage could affect survival or lead
to slower larval growth rates and lowered capacity to reach protected inshore
nursery habitats.
As noted in the "Eggs and Embryos" section
of the "Development" chapter, the viability of
tautog embryos has been reported to decline as the spawning season progresses.
The cause of this decline is not known, but could be environmental, such as
high temperatures (Olla and Samet 1978).
Pearce (1969) reported 31°C to be lethal to adult tautog in a 1-hr change
from ambient during an aquarium study. McCormack (1976), though, reported that
tautog do not lose equilibrium until about 33°C when water temperature
was raised more gradually. At above-ambient temperatures, tautog behavior is
altered; these alterations included changes in activity, dominance, habitat
use, and posture that suggest stress (Olla and Studholme 1975; McCormack 1976);
Olla and Studholme (1975) reported that the fish recover normal behavior when
water temperatures return to a normal ambient range.
Olla et al. (1980) found tautog to survive in waters as cold as 1.9°C.
The tautog's relative, the cunner, is reported to tolerate seawater temperatures
slightly below 0°C (Green and Farwell 1971). Temperature extremes have
been shown to reduce response thresholds in tautog (Liem and Sanderson 1986).
Tautog are sometimes killed if caught in shoal water by a quick drop to subfreezing
air temperatures. This type of mortality was reported for 1841, 1857, 1875,
and 1901 along Rhode Island and Massachusetts coasts (Bigelow and Schroeder
1953).
Water pH levels below 7.8 were reported stressful to this species, and prolonged
exposure to these levels can result in death (Behre 1933).
Contaminants
Metals
Mears and Eisler (1977) determined concentrations of chromium, copper, iron,
manganese, nickel, and zinc in tautog liver tissue. Their results suggested
patterns of concentrations of these metals that were related to body length
and sex. In males, chromium and copper concentrations decreased with length;
in both sexes, nickel concentrations decreased. In both sexes, iron, manganese,
and zinc concentrations were not related to body length. Hall et al. (1978)
also examined 15 metals in this species (collection location not defined) and
found low to average levels in muscle samples.
A recent study (National Marine Fisheries Service 1995) found muscle concentrations
of nine metals (i.e., silver, cadmium, chromium, copper, nickel, lead,
zinc, arsenic, and mercury) to be well below U.S. Food and Drug Administration
(FDA) action levels in tautog collected off Manasquan Inlet (New Jersey). Mercury
levels in tautog were higher than those in bluefish, black sea bass (Centropristis
striata), and summer flounder, but still well below FDA action levels.
Mallet (1972) reported that acute, sublethal (i.e., 20 ppm) cadmium
exposures and intoxication caused a change in tautog cardiac response to gradual
hypoxia; this response included higher variability in heart rates and lowered
rate-response sensitivity to oxygen levels, which suggest that the metal interferes
with the cardiac regulatory mechanism for hypoxia. Lethal cadmium exposures
(i.e., 400 ppm) resulted in death in about 5 hr, but produced no significant
histological damage to gills, liver, kidney, or intestine. In another study,
Mallet noted that cadmium levels on gill tissue can be very high.
Studies of the effect of longer-term experimental exposure of cadmium on the
closely related species, cunner, found that this metal did cause pathological
changes to a number of organs and tissues (National Marine Fisheries Service
1974; MacInnes et al. 1977).
Organics
Tautog fillets from the Hudson River - Raritan Bay Estuary, collected between
1980 and 1985, were examined for Chlordane, DDT, DDD, DDE, dibenzoanthracine,
Endine, Heptachlor, hexachlorobenzene, Lindane, and total polychlorinated biphenyls
(PCBs). Concentrations of these organics were found to be low, relative to
concentrations in some other fish species examined (New York State Department
of Environmental Conservation 1987). This general result was confirmed in a
recent study by the National Marine Fisheries Service (1995) which also found
that all concentrations of PCBs and pesticides in tautog tissue were below
an FDA action level of 2.0 ppm. Polycyclic aromatic hydrocarbons were largely
undetected in the tautog tissue.
No information was found of the effects of exposure to organics on tautog.
Several studies are available for cunner, however, the other labrid in the
Northeast, the results of which could be similar for tautog. Payne et al. (1978)
found little pathological effect in cunner from a 6-mo exposure to petroleum
hydrocarbons. Payne and May (1979) further reported that cunner were able to
metabolize petroleum hydrocarbons. Williams and Kiceniuk (1987) reported that
a prolonged exposure to crude oil is required to suppress feeding by cunner,
and that recovery can occur in a few weeks. Deacutis (1982) found, however,
that tautog would not hesitate to consume hard clam or mussel meats that were
contaminated with #2 fuel oil, and that tautog made little effort to avoid
oil-contaminated feeding areas. An environmental threat to tautog can exist
in petroleum- hydrocarbon-contaminated areas because bivalve mollusks, a common
prey, cannot metabolize polycyclic hydrocarbons and can accumulate petroleum
hydrocarbons to high levels and lose it slowly, maintaining a reservoir for
continued induction (Vandermeulen and Penrose 1978).
Klein-MacPhee et al. (1993) reported larval tautog survived, but had
significantly reduced feeding and growth, in an experimental exposure to starch
blended with vinyl alcohol copolymers (i.e., a biodegradable, potential
substitute for plastic packaging). The low nutritional value of starch to carnivorous
fish and a low DO concentration were suggested as being involved in these results.
ECOLOGICAL ROLES
The tautog is an important member of the resident, coastal, three-dimensionally
structured habitat community in the Northeast's marine environment. Although
the species' overall ecological role in the community has not been well studied,
we do have some information, such as their predative and competitive interactions
as discussed in the "Feeding and Diet" chapter
and in the "Competitors" section of the "Natural
and Human-Induced Environmental Factors" chapter, or we can suggest reasonable
probabilities. These probable roles require more study, verification, and quantification,
if possible. Some consideration should be given also to the species that could
irreversibly replace tautog in the north-temperate-reef ecosystem, if tautog
abundance is reduced to the point where the species loses its ability to compete
for its former habitats and ecological role.
One example of this role is the species' predation on mussels and other encrusting
and epifaunal organisms found in reef-like habitats, as discussed in the "Feeding
and Diet" chapter. This predation can be important to the biodiversity
of the invertebrate community of these habitats, and to the mesoscale landscape
ecology of these habitats. Tautog grazing on dense sets of mussels and barnacles
can help to reopen habitat space for renewed colonization by other species
necessary for maintaining biodiversity and habitat patchiness. That is, the
tautog may be a "keystone" species in the same manner as Paines' (1969) sea
star was in defining the concept. Its occasional feeding on anemones, sponges,
bryozoans, and other long-lived, relatively unproductive, surface-encrusting
taxa can also open space for colonization by more productive species, such
as mussels or oysters, that can also be used as food by other fisheries resources
or be harvested by humans. The tautog's feeding can be important in controlling
certain predators or competitors, such as crabs, gastropods, or tunicates,
of juvenile harvestable shellfish. Heavy tautog predation on shellfish beds
or other encrusting fauna can also increase the rate of recycling of nutrients,
which accumulate in these filter-feeding organisms, back into the water. Its
preference for mussels and shellfish can be a problem, however, if it feeds
on shellfish populations that are cultured or harvested by humans.
Tautog will resort to shellfish prey from the surface of open habitats, such
as small clams in muddy or sandy areas near and away from shelter, when epifaunal
food resources of the shelter become scarce. This alternate use of off-reef
prey can thus affect soft-bottom community ecology, as well, in the vicinity
of a reef habitat with a tautog population.
By crushing the shells of their molluscan prey during feeding, tautog also
contribute to available habitat types and use by altering a coarse, empty-whole-shell-rich
habitat that would remain if mussels only died of adverse environmental conditions
or age, to a finer, shell-hash-sediment habitat. The coarse shell or the shell
hash creates different habitats used or favored by different invertebrate or
post-larval fish and epifaunal species assemblages.
Some indirect effects of tautog populations on habitats, such as that of certain
harvesting gear targeting tautog, are discussed in the "Structure" subsection, "Habitat
Modification and Loss" section of the "Natural
and Human-Induced Environmental Factors" chapter.
Although small tautog are preyed upon (see the "Predators" section
of the "Natural and Human-Induced Environmental Factors" chapter),
it does not appear that any predator is highly dependent on them as food, or
that tautog serve as a key forage species. Their localized populations, slow
growth, and relatively long life can provide stability to resident fish communities
within which they occur.
RESEARCH NEEDS
This review shows that there is much known about the life history and habitat
requirements of the tautog, although there are still many weak areas or gaps
that need attention before it can be claimed that we know enough. Some of our
knowledge of the species comes from laboratory studies; some effort must be
made to confirm these controlled-environment results in the field before they
are used, without question, in planning. In turn, some field observations need
to be examined in the laboratory to define better the cause-and-effect relationships.
The variations or differences in some results from multiple studies also need
to be resolved, or the reasons for the differences understood.
This review suggests there are areas for which life history and habitat requirement
information is nonexistent or weak for tautog. Some of these key areas that
require special research attention are presented below, noting in many cases
the implications for fishery resource and habitat managers:
- Defining specific prespawning and spawning aggregation areas used by all
major local populations, as well as defining the criteria for, or times
of, use of these areas. It will be critical to protect these areas from degradation,
and to protect these populations against excessive exploitation during
their
use of these areas.
- Defining specific wintering areas used by juveniles
and adults of all major local populations, as well as defining the criteria
for,
or times of, use of these areas. It will be critical to protect these
areas from degradation, and to protect these juveniles and adults against
excessive
exploitation during their use of these areas.
- Defining specific migration
routes used by tautog to get to and from spawning and wintering areas.
It will be critical to protect these migration routes from degradation,
and to protect
these fish against excessive exploitation during their use of these
routes.
- Defining sources of offshore eggs and larvae -- are the eggs and
larvae from in
situ sources,
or have they been washed out of coastal spawning areas? If offshore
eggs and larvae do come from coastal spawning areas, then weather
may be a
critical factor in recruitment.
- Defining the extent, condition (e.g.,
optimum, suitable), and trends of juvenile habitats. It will be critical
to protect
these habitats, or to stimulate their restoration or enhancement,
if required.
- Exploring possible genetic differences within the overall
tautog population, noting
their geographical distribution and trends, and relating them to
recruitment,
growth,
and exploitation rates. Such knowledge of different/local genetic
components within the overall population could support more effective
regional
management of this species.
- Confirming that tautog, like cunner, "hibernate" in
the winter, and if they do, determining in what areas, for how long,
and with
what special
habitat needs. It would be important to understand such behavior,
particularly as it might affect harvesting availability and vulnerability.
- Defining
susceptibility of juveniles to coastal contamination and contaminant
effects. Such knowledge
would be important for assessing and managing habitat/population
damage.
- Defining
the role of prey type and availability in local juvenile/adult population
dynamics. Such knowledge could help to explain differences in local
abundance, movements,
growth, fecundity, contaminant burdens, etc.
- Defining larval tautog
diets and prey availability requirements.
- Better defining and quantifying
the
ecological role of tautog in coastal "reef," or grass bed, communities.
ENDNOTES
- NEFSC, Woods Hole, MA; October 1995.
- South
Carolina Wildlife and Marine Resources Commission, Charleston, SC;
1995.
- NOAA-UMass
Cooperative Marine Education and Research Program, Amherst, MA; May
1999.
- NEFSC,
Milford, CT; 1995.
- The "K" term is a component of
the von Bertalanffy growth equation: lt = L [1-e-K(t-t0)],
where lt = length at age t, L = ultimate length
associated with a particular stock or population, K = coefficient of
growth, e = base
of natural logarithms, and t0 = age when length would theoretically
be zero.
- NEFSC, Milford, CT; 1995.
- Bureau
of Aquaculture, Connecticut Department of Agriculture, Milford, CT;
1998.
ACKNOWLEDGMENTS
We
thank Claire Steimle and Judy Berrien for their assistance in finding
much of the literature used here, and to Tina Berger, Dean Perry, Barbara
Dorf, the
Atlantic States Marine Fisheries Commission's Tautog Technical Committee, and
others for sharing information with us.
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Acronyms |
| DO |
= |
dissolved oxygen |
| FDA |
= |
[U.S. Department of Health and Human Services'] Food and Drug
Administration |
| FL |
= |
fork length |
| FO |
= |
frequency of occurrence |
| GSI |
= |
gonadal-somatic index |
| MARMAP |
= |
[NEFSC's] Marine Resources Monitoring, Assessment, and Prediction
Program |
| NEFSC |
= |
[National Marine Fisheries Service's] Northeast Fisheries Science
Center |
| PCB |
= |
polychlorinated biphenyl |
| TL |
= |
total length |
| YOY |
= |
young of the year |
