HomeMy WebLinkAboutTautog life history NMFSNOAA Technical Memorandum NMFS-NE-118
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Tautog (Tautoga onitis)
Life History and Habitat Requirements
Frank W. Steimle1 and Patricia A. Shaheen2
1National Marine Fisheries Serv., Highlands, NJ 07732
2Rutgers Univ., New Brunswick, NJ 07102
U.S. DEPARTMENT OF COMMERCE
William Daley, Secretary
National Oceanic and Atmospheric Administration
D. James Baker, Administrator
National Marine Fisheries Service
Penelope D. Dalton, Assistant Administrator for Fisheries
Northeast Region
Northeast Fisheries Science Center
Woods Hole, Massachusetts
May 1999
Note on Species Names
The NMFS Northeast Region's policy on the use of species names in all technical communications is generally to follow the
American Fisheries Society's lists of scientific and common names for fishes (i.e., Robins et al. 1991a), mollusks (i.e., Turgeon
et al. 1998b), a decapod crustaceans (i.e., Williams et al. 1989c), and to follow the Society for Marine Mammalogy's guidance
on scientific and common names for marine mammals (i.e., Rice 1998d). Exceptions to this policy occur when there are
subsequent compelling revisions in the classifications of species, resulting in changes in the names of species (e.g., Cooper
and Chapleau 1998e).
___________________
aRobins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. Common and scientific names
of fishes from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ.20; 183 p.
bTurgeon, D.D. (chair); Quinn, J.F.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;
Rosenberg, G.; Roth, B.; Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic
invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. Publ. 26; 526 p.
cWilliams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common
and scientific names of aquatic invertebrates from the United States and Canada: decapod crustraceans. Amer. Fish. Soc. Spec. Publ. 17;
77 p.
dRice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal Spec. Publ. 4; 231 p.
eCooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised
classification. Fish. Bull. (U.S.) 96:686-726.
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Contents
Abstract ................................................................................................................................................................................ v
Introduction ...........................................................................................................................................................................1
Nomenclature and Taxonomy.................................................................................................................................................1
Distribution and Habitat ........................................................................................................................................................2
Reproduction .........................................................................................................................................................................6
Development..........................................................................................................................................................................7
Age and Growth.....................................................................................................................................................................8
Feeding and Diet....................................................................................................................................................................9
Behavior............................................................................................................................................................................... 11
Population Structure ............................................................................................................................................................ 13
Natural and Human-Induced Environmental Factors ........................................................................................................... 13
Ecological Roles................................................................................................................................................................... 16
Research Needs ................................................................................................................................................................... 17
Endnotes .............................................................................................................................................................................. 18
Acknowledgments ............................................................................................................................................................... 18
References Cited .................................................................................................................................................................. 18
Tables
Table 1. Reported length-weight relations for tautog.......................................................................................................9
Figures
Figure 1.Drawing of tautog by H.L. Todd based on specimen collected by Vinal N.
Edwards at Woods Hole, Massachusetts, on December 13, 1875......................................................................vi
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 = [N ational Marine Fisheries Service’s] Northeast Fisheries Science Center
PCB = polychlorinated biphenyl
TL = total length
YOY = young of the year
Page iv
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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 require-
ments 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.
Page vi
Figure 1. Drawing of tautog by H.L. Todd based on specimen collected by Vinal N. Edwards at Woods Hole, Massachusetts, on December 13, 1875.
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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 dur-
ing post-larval stages. Fishery interest in the species has
increased in recent years, and additional management mea-
sures 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 Com-
mission 1995). This report compiles and summarizes avail-
able 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 fish-
ing 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 stud-
ies 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 prob-
ably an appropriate estimate for tautog.
NOMENCLATURE AND TAXONOMY
NOMENCLATURE
Valid Name
Tautoga onitis (Linnaeus, 1758) is the name recog-
nized 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, fol-
lowing 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 (Jor-
dan 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, longitu-
dinally 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) sug-
gests that tautog may have characteristics which are primi-
tive 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 com-
monly 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 elon-
gate, 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,
Page 2
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 black-
ish, 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 back-
ground. Fish are also observed being dull gray-white. Some
of these variations could reflect different environmental fac-
tors, 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 cun-
ner it is relatively straight. The caudal peduncle of the
tautog is proportionally wider, and the caudal fin is nar-
rower, 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 differ-
ential 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 labora-
tory (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 conspicu-
ous 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 be-
come warmer in summer and remain slightly warmer in win-
ter (Bleakney 1963).
The tautog is most abundant from Cape Cod to Chesa-
peake 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 dis-
tance 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 coun-
tered 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).
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In terms of the persistent, overall range of the spe-
cies, 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 oc-
curred since the last glacial period. Empirically, however, it
appears that recent warming of air temperatures in the Arc-
tic 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). Spawn-
ing 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 locali-
ties 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 sea-
sonally 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 oc-
currence 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 progres-
sion through the summer, beginning in April in the south-
ern part of the Middle Atlantic Bight, and extending to the
northern areas by May (Able and Fahay 1998). Peak spawn-
ing 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 col-
lected 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 as-
sociated 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 eel-
grass (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 re-
spond 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 be-
fore 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 adapta-
tion 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) tau-
Page 4
tog 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 re-
ported 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) re-
ported 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 avail-
ability of cover (i.e., any object that an individual can re-
main 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) re-
ported 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 com-
mon 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 struc-
tured 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. Al-
though 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, estab-
lishing 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 juve-
niles 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 col-
lected 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 be-
tween 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-ex-
isting 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
5Page
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 Chesa-
peake Bay (Hostetter and Munroe 1993). Eklund and Targett
(1990) noted populations in 25-35 m of water 22-37 km off-
shore 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 resi-
dent 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 fac-
tors 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 juve-
niles 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, prob-
ably caused by high water temperatures (personal observa-
tion 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 bot-
tom 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 tem-
perature, 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 shel-
ter, 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 pro-
vide the “reef” habitat that tautog require, although shell-
fish beds in euryhaline parts of estuaries serve as habitat
(Arve 1960). In this area, the man-made “reef” habitat cre-
ated 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 im-
portant 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 habi-
tat where sea lettuce (Ulva) was present. Ulva can have
negative effects on certain taxa or species, and the prefer-
ence or tolerance of Ulva by tautog can give it an advan-
tage with competitors or predators (Dorf 1994). In contrast
to studies that report the importance of vegetation, Dixon
Page 6
(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 re-
sembles 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 spawn-
ing opportunities, or 4) coincident with a different repro-
ductive 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 be-
tween 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 produc-
tion 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 indi-
ces just after spawning. The GSIs for female and male tau-
tog 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 distribu-
tion (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 sev-
eral 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 stud-
ies 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 in-
crease in the male:shelter ratio, an inability to control a ter-
ritory, 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 after-
noon, 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).
7Page
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 convo-
lutions. At first the developing ova are opaque,
relatively hard, and average 0.36 mm in diam-
eter. As development proceeds the ovary en-
larges and transparent, soft, mature ova ap-
pear 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 em-
bryos collected in central Long Island Sound near New
Haven, Connecticut, declined as the spawning season pro-
gressed. 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 ap-
proximately 2.2 mm in length, but can be slightly less. [Some
variance in the lengths of specimens reported in the litera-
ture 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 lo-
cated 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 black-
ish 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 chro-
matophores 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 dis-
play well differentiated dorsal, anal, and caudal fins. Pig-
ment 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 rela-
tionship: 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).
Page 8
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 characteris-
tics of adults, except for their often greenish background
color. The black chromatophores form heavily pigmented
areas which give the body a transversely banded appear-
ance 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 ex-
ample, 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 (Coo-
per 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 deter-
mined 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 ge-
netic selection process for sexual maturity is unknown at
present; this pressure can favor the portion of the popula-
tion 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 re-
ported 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 progres-
sions, otolith age - fish size comparisons, and direct mea-
surements 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 spe-
cies, 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 Vir-
ginia 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%, oc-
curred during the second year for both sexes (Cooper 1967;
Sogard et al. 1992). In southern Virginia waters, the great-
est 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 juve-
nile 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 fe-
males 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
9Page
by Simpson (1989) in Long Island Sound. In southern Vir-
ginia 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 at-
tained 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 oc-
curred from January to March because of decreased feed-
ing 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 re-
main cold longer. In southern Virginia waters, the least
somatic growth in adults occurred from March to June dur-
ing 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 wa-
ters. For males in Rhode Island, annual growth increments
decreased to less than 12 mm after age 12, and further de-
clined 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 de-
clined 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), com-
pared with L = 733 mm for females and 732 mm for males in
southern waters (Hostetter and Munroe 1993).
Several studies reported variable length-weight rela-
tionships for whole and eviscerated tautog from
Narragansett Bay to southern Virginia waters (Table 1).
Tautog are a relatively long-lived fish, with the old-
est 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 re-
ported 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.
Table 1.Reported length-weight relations for tautog. [W = weight (g); L = length (mm); Wo = weight (oz); Li = length
(inches); and (E) = eviscerated weight.]
Area Sex Relationship Source
Rhode Island male Log W(E) = -4.357 + 2.776 log L Cooper (1967)
female Log W(E) = -4.804 + 3.016 log L Cooper (1967)
Long Island both Log Wo = -5.992 + 2.916 log Li Briggs (1969a)
New York Bight male Log W = -5.203 + 3.206 log L Wilk et al. (1978)
female Log W = -5.444 + 3.230 log L Wilk et al. (1978)
both Log W = -4.721 + 3.020 log L Wilk et al. (1978)
Virginia both Log W = -4.632 + 2.979 log L Hostetter and Munroe (1993)
Page 10
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 re-
port “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 wa-
ters. Frequency of occurrence (FO) in the digestive tract
was 98% for amphipods, 94% for copepods, 29% for poly-
chaetes, and 25% for isopods. This diet indicated that ju-
venile 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 crusta-
ceans (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) re-
ported 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 (Mercenaria
mercenaria).
Adults
Adult tautog are durophagous and feed chiefly on
blue mussels and other shellfish throughout the year. Bar-
nacles (Balanus sp.), brachyuran crabs, hermit crabs, sand
dollars, (bay?) scallops, amphipods, decapod shrimp, iso-
pods, lobsters, and probably nereid polychaetes were re-
ported 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 tuni-
cates. Bleakney (1963) reported tautog from Nova Scotia to
feed not on blue mussels, but on horse mussels (Modiolus
modiolus) and periwinkles (Littorina littorea). 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 At-
lantic 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
(Echinarachnius parma) (38%), Atlantic rock crabs (28%),
and blue mussels (9%) dominated the tautog diet on a volu-
metric 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 anemo-
nes (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, deca-
pod crabs, and other crustaceans of suitable size were re-
ported 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, how-
ever, when they were without shelter in aquaria.
Chao (1973) reports that the cunner lacks a well de-
fined stomach, which is a characteristic of most labrids,
including tautog. The tautog’s gastro-intestinal tract is semi-
11Page
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) ob-
served that tautog followed the flood tide up above low
water levels, around ledges, to prey on mussels in the inter-
tidal 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, separat-
ing it in the process into smaller, crushable sizes. The feed-
ing 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 stom-
achs. Curran (1992) found all feeding stopped in the closely
related cunner (and probably tautog) when water tempera-
tures reached the low levels that induce torpor and hiberna-
tion. 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 cer-
tain levels. Tautog which had been acclimated to tempera-
tures of 19°C and 21°C, respectively, decreased their inges-
tion 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 “Natu-
ral and Human-Induced Environmental Factors” chapter.)
Tautog were observed to vary, to some extent, their
feeding in association with their place in a group domi-
nance 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 spawn-
ing. 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 con-
finement 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 bot-
tom 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” chap-
Page 12
ter.) Channels may be important pathways of migration into
coastal areas for spawning and out again to deeper or off-
shore wintering areas (Cooper 1966). As noted previously,
Briggs (1977) reported that part of the southern Long Is-
land 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 spawn-
ing (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 move-
ment to deeper water in the fall off New York. This is sup-
ported 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-morn-
ing, 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 condi-
tion 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 tempera-
ture of 7.5°C. In laboratory studies, tautog activity de-
creased as temperatures declined to 5.2°C, then tautog ag-
gregated 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 de-
spite 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 in-
crease in the tendency to aggregate (McCormack 1976; Olla
et al. 1978). This lowered response to stimuli at tempera-
ture extremes was supported by the sound-detection-thresh-
old experiments of Offutt (1971).
HIERARCHICAL DOMINANCE
In laboratory studies, a dominance hierarchy was ob-
served 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 to-
ward 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 thump-
ing, 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 pur-
pose 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 inten-
sity of body color, while high-intensity light enhanced the
body’s dark coloration. Differently colored light and se-
quential 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 photoreac-
13Page
tion 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 Hu-
man-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 tau-
tog he collected in Narragansett Bay were female. In north-
ern 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 be-
cause the older fish (greater than 18 yr old) were predomi-
nantly 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 popu-
lation (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 (Raja laevis), 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 (Merluccius bilinearis) 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 cryp-
tic 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 (Con-
ger oceanicus), summer flounder (Paralichthys dentatus),
and various sharks, can be expected to prey upon small
tautog. Although American lobsters share the same habi-
tat with tautog, and are active at night when tautog are
inactive, there are no reports of lobster predation on tau-
tog.
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 men-
tioned 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 ob-
served 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, habi-
tat use, and growth or mortality rates is unreported or un-
known.
Humans as fishermen are also predators. Simpson
(1989) estimated fishing mortality for a Long Island Sound
Page 14
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 (Archosargus probatocephalus), and
American lobster on the other hand has been suggested
because all four species use the same habitat and prey com-
monly on blue mussels, other small mollusks, and crusta-
ceans, 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 tau-
tog for this food resource too (Richards 1963). Other inver-
tebrate 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 competi-
tors 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 ecologi-
cal 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 ves-
sels, 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 consid-
ered 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 re-
sponses to certain human enteric bacteria. These responses
indicate that tautog are immunologically responsive to ex-
posure to sewage bacteria (Stolen et al. 1983).
Finrot disease, thought to be caused by some micro-
bial agent and anthropogenic stress, was not found on tau-
tog in the New York Bight, although the disease was re-
ported 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 physi-
cal abrasions. Laird and Bullock (1969) found no positive
evidence of hematozoan parasites in New England and Ca-
nadian 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 col-
lected 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 re-
lated to a nearly concurrent, offshore bloom of the dinoflagel-
late 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 (Chesa-
peake Executive Council 1994). The loss of vegetated es-
tuarine 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.
15Page
(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 patho-
logical conditions or other possible effects from this physi-
cal habitat alteration and disturbance.
In the past, salvaging of certain metal shipwrecks or
lowering of wreck heights if they might be navigation haz-
ards, reduced the value of shipwrecks as tautog habitat.
Before 1900, some wrecks off New Jersey were even dyna-
mited 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 qual-
ity, including the destruction of slow-growing colonies of
the northern star coral (Astrangia sp.) which provide bio-
genic reef habitat for tautog.
Reef habitats that lose structural height are more
prone to siltation (Rothschild et al. 1994) or burial by sedi-
ment 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 mitigat-
ing this habitat degradation to some degree.
Hydrography
Auster (1989) reported that current velocity can af-
fect small-scale spatial distribution and can change forag-
ing behavior in tautog and other species. The high hydro-
graphic energy caused by passage of a hurricane was re-
ported 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 be-
low 2 mg/l, and prefer DO levels above 3 mg/l. This intoler-
ance is supported by the cardiac and preliminary respira-
tion 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 cad-
mium exposure, and that hypoxia responses can then be
detected at higher oxygen concentrations. Mallet also ob-
served 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 “toler-
ance” 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 rela-
tive, the cunner, were found dead or dying (Ogren and Chess
1969). Tautog mortalities, however, were reported by com-
mercial 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 tempera-
tures 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 obtain-
ing 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 envi-
ronmental, such as high temperatures (Olla and Samet 1978).
Pearce (1969) reported 31°C to be lethal to adult tau-
tog 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 re-
cover normal behavior when water temperatures return to a
normal ambient range.
Page 16
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 concentra-
tions decreased with length; in both sexes, nickel concen-
trations 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., sil-
ver, cadmium, chromium, copper, nickel, lead, zinc, arsenic,
and mercury) to be well below U.S. Food and Drug Admin-
istration (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) re-
sulted 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 ex-
posure 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 poly-
chlorinated biphenyls (PCBs). Concentrations of these or-
ganics were found to be low, relative to concentrations in
some other fish species examined (New York State Depart-
ment of Environmental Conservation 1987). This general
result was confirmed in a recent study by the National Ma-
rine Fisheries Service (1995) which also found that all con-
centrations 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 re-
sults 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 petro-
leum hydrocarbons. Williams and Kiceniuk (1987) reported
that a prolonged exposure to crude oil is required to sup-
press 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 substi-
tute 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 spe-
cies’ 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” sec-
tion of the “Natural and Human-Induced Environmental
Factors” chapter, or we can suggest reasonable probabili-
ties. These probable roles require more study, verification,
17Page
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” spe-
cies 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 un-
productive, surface-encrusting taxa can also open space
for colonization by more productive species, such as mus-
sels 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 popula-
tion.
By crushing the shells of their molluscan prey dur-
ing feeding, tautog also contribute to available habitat types
and use by altering a coarse, empty-whole-shell-rich habi-
tat that would remain if mussels only died of adverse envi-
ronmental 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 habi-
tats, such as that of certain harvesting gear targeting tau-
tog, are discussed in the “Structure” subsection, “Habitat
Modification and Loss” section of the “Natural and Hu-
man-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 popu-
lations, 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, al-
though 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 labora-
tory 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 de-
fine better the cause-and-effect relationships. The varia-
tions 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 habi-
tat managers:
1. Defining specific prespawning and spawning aggrega-
tion 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 degrada-
tion, and to protect these populations against exces-
sive exploitation during their use of these areas.
2. 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.
3. 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 degrada-
tion, and to protect these fish against excessive exploi-
tation during their use of these routes.
4. 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.
5. 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.
Page 18
6. Exploring possible genetic differences within the over-
all tautog population, noting their geographical distri-
bution and trends, and relating them to recruitment,
growth, and exploitation rates. Such knowledge of dif-
ferent/local genetic components within the overall popu-
lation could support more effective regional manage-
ment of this species.
7. 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 vulnerabil-
ity.
8. Defining susceptibility of juveniles to coastal contami-
nation and contaminant effects. Such knowledge would
be important for assessing and managing habitat/popu-
lation damage.
9. 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.
10. Defining larval tautog diets and prey availability require-
ments.
11. Better defining and quantifying the ecological role of
tautog in coastal “reef,” or grass bed, communities.
ENDNOTES
1. NEFSC, Woods Hole, MA; October 1995.
2. South Carolina Wildlife and Marine Resources Com-
mission, Charleston, SC; 1995.
3. NOAA-UMass Cooperative Marine Education and Re-
search Program, Amherst, MA; May 1999.
4. NEFSC, Milford, CT; 1995.
5. 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.
6. NEFSC, Milford, CT; 1995.
7. 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 Commit-
tee, and others for sharing information with us.
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