NOAA Technical Memorandum NMFS-NE-118

This series represents a secondary level of scientific publishing. All issues employ thorough internal scientific

review; some issues employ external scientific review. Reviews are -- by design -- transparent collegial

reviews, not anonymous peer reviews. All issues may be cited in formal scientific communications.

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).

___________________

a

Robins, 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.

b

Turgeon, 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.

c

Williams, 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.

d

Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal Spec. Publ. 4; 231 p.

e

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised

classification. Fish. Bull. (U.S.) 96:686-726.
Page iii

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 = [National Marine Fisheries Service’s] Northeast Fisheries Science Center

PCB = polychlorinated biphenyl

TL = total length

YOY = young of the year
Page iv
Page v

ABSTRACT

This report compiles and summarizes available information on the tautog (Tautoga onitis), covering nomenclature and

taxonomy, distribution and habitat, reproduction, development, growth, feeding and diet, behavior, population structure,

natural and human-induced environmental factors, and ecological roles. The report also identifies research needs and

includes an extensive bibliography.

Recent declines in this species’ abundance and certain known aspects of its life history and specific habitat requirements have caused coastal fishery resource and habitat managers to believe the species may need further conservation

measures. Essential to developing an effective conservation management strategy for this species is a thorough summary of

what is known of the species’ life history and of the habitat requirements for all life stages. This information will be important

for developing holistic approaches to managing a sustained population and fishery for this species and closely associated

members of its ecological community.

This review shows that although much is known about the species, and studies on its life history and ecology are

ongoing, there are important gaps and conflicting or unconfirmed results in our understanding of the species, and its needs,

which should be addressed.
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.
Page 1

INTRODUCTION

The tautog (Tautoga onitis) is a valuable recreational

and commercial fishery resource from Massachusetts to

Virginia in the Northwest Atlantic. It is commonly found on

complexly structured, vegetated, or reef-like habitats during post-larval stages. Fishery interest in the species has

increased in recent years, and additional management measures for this fishery are being considered. To support the

development of information necessary to produce good

public policy, an assemblage of existing information on the

species is needed (Atlantic States Marine Fisheries Commission 1995). This report compiles and summarizes available information on the life history and habitat requirements

of, and natural and human-induced environmental threats

to, tautog. It builds upon and substantially expands the

previous efforts of Auster (1989), Gray (1992), and others.

This review is needed because of: 1) recent increased fishing effort for, and resource user conflicts over, tautog

(DiLernia 1993); 2) sensitivity of reef fish such as tautog to

exploitation (Hostetter and Munroe 1993); 3) the species’

particular habitat needs and the threats to this habitat; and

4) the possible need to exercise additional management of

at least certain localized populations of this species

(Hostetter and Munroe 1993). Information on regional-level

stock abundance, and detailed discussion of state-level

populations and harvests, are not included here, but have

been compiled by Lazar (1995).

As there are incidental references to this species in

many documents and papers, and as there are ongoing studies wholly or partially involving this species, such a review

cannot be definitive. It can, however, serve as a stepping

stone to adequate knowledge for fishery resource and/or

habitat management, or for fishery research planning, as

have previous reviews. Because of the scarcity or absence

of certain information on tautog, relevant information from

studies of the its close labrid relative, the cunner

(Tautogolabrus adspersus) -- which has similar habitat needs

-- has sometimes been considered and discussed as probably an appropriate estimate for tautog.

NOMENCLATURE AND TAXONOMY

NOMENCLATURE

Valid Name

Tautoga onitis (Linnaeus, 1758) is the name recognized by the American Fisheries Society (Robins et al. 1991).

The generic term is the original name applied to this fish

species supposedly by Narragansett Indians. The species

term is derived from the Latin, onitis, which means “a kind

of plant”; but its application by Linnaeus is unclear (Smith

1907).

Synonymy

The species has a rich synonymic history which, following Jordan and Evermann (1896-1900) and Jordan et al.

(1930), includes: Labrus onitis (Linnaeus 1758 and 1766);

L. hiatula (Linnaeus 1766); L. carolinus (Bonnaterre 1788);

L. blackfish (Schopf 1788); L. subfuscus (Walbaum 1792); L.

tesselatus (Block 1792); Hiatula gardeniana (Lacepede

1800); L. americanus (Block and Schneider 1801); L. tautoga

rubens (Mitchill 1814, 1815); L. tautoga alia (Mitchill 1814,

1815); Tautoga niger (Mitchill 1814, 1815); L. tautoga fusca

(Mitchill 1815); T. tessellata (Cuvier and Valenciennes 1839);

T. americana (DeKay 1842); T. onitis (Gunther 1862; Uhler

and Lugger 1876; Yarrow 1877; Jordan and Evermann 1898;

Evermann and Hildebrand 1910; Fowler 1912); H. onitis (Jordan 1886; Jenkins 1887; Bean 1891); and H. hiatula (Goode

and Bean 1885).

TAXONOMY

Description and Affinities

This species is a member of the Labridae, a family of

lipped fishes commonly called wrasses. This family takes

its name from the presence of conspicuous, thick, longitudinally folded lips which, along with other characteristics

such as the form of the jaws, give the mouth a peculiar

appearance (Smith 1907; Liem and Sanderson 1986). The

mouth is terminal and considered of small or moderate size

(Hildebrand and Schroeder 1928). Westneat (1995) suggests that tautog may have characteristics which are primitive within the family, mostly based on its hard-prey diet

and associated jaw morphology.

Worldwide, there are 500-600 species in the Labridae

(Nelson 1984). In the United States, there are 12 genera

(Robins et al. 1991). Five labrid species occur in North

Carolina waters (Smith 1907), but only two species are commonly found north of Cape Hatteras: the cunner,

Tautogolabrus adspersus (Walbaum, 1792), and the tautog,

Tautoga onitis (Linnaeus, 1758), (Hildebrand and Schroeder

1928; Bigelow and Schroeder 1953).

The genus Tautoga (Mitchill) consists of a single

species: Tautoga onitis. The following generic description

is from Hildebrand and Schroeder (1928): “Body is elongate, moderately deep and compressed; anterior profile

rather strongly arched, head nearly as deep as long; eye

small, placed high; mouth rather small; lips quite broad and

thick” (Figure 1). “Teeth in the jaws strong, the anterior

ones more or less conical and incisor-like.” Both the roof of

the mouth and the floor of the throat (i.e., pharynx) have a

patch of knob-like teeth that are used to crush and grind

mollusk and crustacean prey (Bigelow and Schroeder 1953;

Liem and Sanderson 1986). The scales are small, with about

70 being in the lateral series, but cheeks and operculums are

largely scaleless. The dorsal fin is long and continuous,
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 blackish, brownish, blackish green, or blackish blue, with sides

irregularly mottled or blotched. The lips, chin, throat, and

belly are often lighter. Larger males are gray with white

markings on the caudal, pelvic, and dorsal fins, and on the

chin -- a very conspicuous characteristic. Females and

smaller males are without white markings on fins and chin,

and their fins are often plain like the color of the body. Their

eyes are green. Juveniles are usually colored green or brown

with more distinctive side mottling and three or more darker

bars. This coloring can vary with the visual characteristics

of the habitat they are using. In some areas, two color

patterns are often recognized; one being plain blackish and

the other having irregular blackish bars on a pale background. Fish are also observed being dull gray-white. Some

of these variations could reflect different environmental factors, such as light or stress. (See the “Color Modulation”

section of the “Behavior” chapter.)

The tautog can be distinguished from its close and

co-occurring relative, the cunner, in several ways. The adult

tautog is longer and stouter than the adult cunner. The

dorsal profile of its head is highly arched, while in the cunner it is relatively straight. The caudal peduncle of the

tautog is proportionally wider, and the caudal fin is narrower, than those of the cunner (Bigelow and Schroeder

1953; Leim and Scott 1966). Tautog lack scales on the cheeks

and operculums, while on cunner they are present

(Hildebrand and Schroeder 1928).

Subspecies

No subspecies are recognized. However, the differential juvenile growth rates at the range extremes of the

species, discussed in the “Growth” chapter, have been

thought possibly to reflect some degree of evolutionary

divergence, but this has not been supported in the laboratory (Martin 1993).

Common and Vernacular Names

Tautog is the common name for T. onitis accepted by

the American Fisheries Society (Robins et al. 1991), but

“blackfish” is also widely used. Other regional common

names include: Canada -- tautogue noir (Leim and Scott

1966); Maine -- white chin (Bigelow and Schroeder 1953);

New York -- blackfish (Goode 1887; Hildebrand and

Schroeder 1928); New Jersey -- smooth blackfish, tautog,

and chub (Goode 1887); Maryland -- black porgy, salt-water

chub, chub, and blackfish (Hildebrand and Schroeder 1928);

Virginia -- moll and will-george (Goode 1887); and North

Carolina -- sea tench (in a 1709 usage) and oyster-fish

(Goode 1887; Smith 1907).

The Maine name “white chin” refers to the conspicuous white coloration of the lower jaw of older males (Bigelow

and Schroeder 1953; Hostetter and Munroe 1993). Another

name used in some places is “slippery bass” which refers to

the tautog’s mucus covering and rough resemblance to a

bass. Jordan et al. (1930) also mention the names “cub”

and “sea dog” being used for the species.

DISTRIBUTION AND HABITAT

GENERAL DISTRIBUTION

The tautog is a generally a coastal species found on

the Atlantic coast of North America, from the outer coast of

Nova Scotia to South Carolina (Bigelow and Schroeder 1953).

It has been anecdotally, and perhaps dubiously, reported

on central Georges Bank from commercial fishery catch data

(Chang 1990). The report of tautog on Georges Bank is not

reliably supported by Northeast Fisheries Science Center

(NEFSC) bottom trawl survey records, although it has been

suggested that a few tautog may be collected near the

boundaries of the bank every decade (T. Azarovitz, pers.

comm.1

). The tautog may be a “relict” species north of

Massachusetts, found in certain, deep, saltwater lakes and

protected bays of Nova Scotia that have waters that become warmer in summer and remain slightly warmer in winter (Bleakney 1963).

The tautog is most abundant from Cape Cod to Chesapeake Bay. North of Cape Cod, it is unusual to find tautog

more than 6 km from land, or in waters deeper than 18 m.

South from Cape Cod to about New Jersey, it can be found

to 19 km offshore in up to 24 m of water, and occasionally

near the deep Great South Channel between Nantucket

Shoals and Georges Bank (Chang 1990). This offshore distance and depth range appear to increase gradually towards

the south and near Cape Hatteras (Chang 1990; Hostetter

and Munroe 1993). Tautog have been reported in brackish

water, but not in freshwater (Bigelow and Schroeder 1953).

It has been reported up to 70 km upstream from the mouth

of the Hudson River (New York) (Beebe and Savidge 1988),

and formerly in the Patapsco River in Baltimore, Maryland

(Fowler 1912).

It is uncertain if tautog populations at the northern

and southern extremes of its range were introduced by man.

To the north, reports that the tautog was introduced to

Cape Ann (Massachusetts) in the late 1800s were countered by other reports that it had been abundant many years

previous to that time. To the south, reports that it had been

introduced to South Carolina were countered by skepticism

that its range could be artificially extended southward

(Goode 1887). Tautog are currently rarely observed or

caught in South Carolina waters (M. Bell, pers. comm.2

).
Page 3

In terms of the persistent, overall range of the species, the potential effects of long-term global warming (or

cooling) are unknown. Continued global warming could

theoretically increase northern coastal water temperatures

and potentially the presence of the species (and others) in

the Gulf of Maine, and perhaps restrict tautog to north of

Cape Hatteras. Such a northern expansion could have occurred since the last glacial period. Empirically, however, it

appears that recent warming of air temperatures in the Arctic has hastened the melting of the polar ice cap which has,

in turn, increased the flow of cold freshwater into the North

Atlantic. This expansion of the North Atlantic “cold pool”

has already reduced the habitat for young Atlantic salmon,

and may well be, or become, a distribution-reducing factor

for cold-intolerant fish species (K. Friedman, pers comm.3

).

DIFFERENTIAL DISTRIBUTION

Spawning

Adult tautog generally migrate inshore in spring from

coastal wintering sites to spawn (Chenoweth 1963; Cooper

1966; Stolgitis 1970; Olla et al. 1974; Briggs 1977). Spawning occurs primarily at or near the mouths of estuaries and

in inshore waters (Tatham et al. 1984; Feigenbaum et al.

1989; Sogard et al. 1992; Able and Fahay 1998). Inside

Narragansett Bay (Rhode Island), mature tautog returned

to the same spawning sites each year, usually in the upper

estuary, but dispersed throughout the bay after spawning

(Cooper 1966; Dorf 1994; Dorf and Powell 1997). Olla et

al.’s (1980) tagging studies suggested, however, that adult

tautog did not always return to the same spawning site in

the spring, and that population mixing from different localities occurred.

A portion of the adult population reported to remain

offshore throughout the year (Olla and Samet 1977; Eklund

and Targett 1990; Adams 1993; Hostetter and Munroe 1993),

especially in the southern part of its range, was found seasonally in spawning condition. For example, ripe fish were

collected by Eklund and Targett (1990) and Hostetter and

Munroe (1993) on hard-bottom sites 25-35 m in depth and

22-37 km off Maryland and Virginia. Collections of eggs

and larvae from Georges Bank to North Carolina through

the NEFSC’s Marine Resources Monitoring, Assessment,

and Prediction Program (MARMAP) surveys, suggest that

tautog spawning may also occur in continental shelf waters

and be concentrated off Southern New England (Sogard et

al. 1992). Dorf (1994), in contrast, speculates that the occurrence of offshore eggs and larvae can also result from

their being flushed from estuarine spawning areas, at least

in Southern New England.

Spawning is reported to follow a northward progression through the summer, beginning in April in the southern part of the Middle Atlantic Bight, and extending to the

northern areas by May (Able and Fahay 1998). Peak spawning in the central Bight is reported to occur in June and July,

and to decline by August (Berrien and Sibunka, in press).

Eggs and Larvae

Tautog eggs and larvae are collected on the inner

continental shelf, with highest concentrations being collected off Southern New England and Long Island (New

York) (Colton et al. 1979; Sogard et al. 1992; Malchoff 1993).

In the New York Bight (i.e., the continental shelf off Long

Island and New Jersey), larvae are reported to be part of a

summer coastal fish larval assemblage, perhaps closely associated with spawning areas (Cowen et al. 1993). Tautog

is the most abundant of any larval species found recently in

Narragansett Bay (Keller and Klein-MacPhee 1992). In the

Weweantic River Estuary (Massachusetts), the greatest

abundances of eggs and larvae were collected over eelgrass (Zostera marina)-vegetated sites and near bottom

(Stolgitis 1970).

As mentioned earlier, Dorf (1994) believes that eggs

and larvae found in offshore coastal waters can result from

ex situ sources (i.e., flushed out and away from estuaries),

but there are inadequate data to support this hypothesis at

present. Viable tautog eggs, although lacking oil globules,

are buoyant and found in greatest abundance at or near the

water surface (Merriman and Sclar 1952; Herman 1963;

Stolgitis 1970; Fritzsche 1978; Bourne and Govoni 1988).

Nonviable eggs lack or lose buoyancy and probably respond to currents differently, and thus probably become

distributed differently (Perry 1994).

Malchoff (1993) reported that tautog larvae migrate

vertically in the water column. The larvae, 2-4 mm in length,

stay near the water surface (less than 5 m deep) during the

day, but go deeper at night. Older and larger larvae spend

more time at deeper depths as they grow. Malchoff (1993)

also believed the Hudson River plume strongly influences

larval tautog transport patterns in that part of the New York

Bight. Sogard et al. (1992) estimated that larval tautog in

coastal New Jersey spend about 3 wk in the plankton before settling to the benthos, but this period can be a short

as 17 days (Schroedinger and Epifanio 1997). This is a

relatively short planktonic period compared to that of other

labrids. This short period could possibly reflect an adaptation to cool-temperate environments and a restricted period

of optimum warmer conditions for somatic growth.

Juveniles

Newly settled tautog inhabit shallow areas less than

1 m in depth (Warfel and Merriman 1944; Sogard et al. 1992;

Hostetter and Munroe 1993), including tide pools (Breder

1922). Bigelow and Schroeder (1953) noted that “fry” were

often seined from Southern New England to Virginia. Sogard

et al. (1992) reported that as young-of-the-year (YOY) 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 reported to range from primarily eelgrass beds (Goode 1887;

Grover 1982; Orth and Heck 1980; Heck et al. 1989; Sogard

et al. 1992; Szedlmayer and Able 1996) or a mix of eelgrass

and algal associates [i.e., sea lettuce (Ulva lactuca),

Enteromorpha sp., and Polysiphonia] (Briggs and

O’Connor 1971), to beds of mostly Ulva (Nichols and Breder

1926; Sogard and Able 1991). In the Great Bay - Mullica

River Estuary (New Jersey), early YOY preferred sea lettuce

over eelgrass habitats (Sogard 1989). Sogard (1989) reported that juvenile tautog also readily used artificial sea

grass as habitat. For YOY and 1-yr-old juveniles, empty

oyster and clam shells (Bigelow and Schroeder 1953) and

shell and sponge (Szedlmayer and Able 1996) have been

reported to be used as habitat. Dixon (1994) added small

boulders as a preferred habitat type for juveniles.

The main habitat requirement and distribution factor,

however, for juvenile tautog (less than 25 cm) is the availability of cover (i.e., any object that an individual can remain alongside, within, or under) (Olla et al. 1974, 1975).

Along these lines, Dixon (1994) noted that vertical relief is

an important attribute of juvenile habitat. Smith (1907) reported that young fish were abundant around wharves in

North Carolina; Able and Fahay (1998) reported the same

for New York Harbor. Larger juvenile tautog are closely

associated with hard- surface, reef-like habitats (Olla et al.

1979). Zawacki (1971) reported that 15-20 cm fish were common around a 1-mo-old, automobile-tire artificial reef in

Shinnecock Inlet, Long Island. Adams (1993) reported only

juveniles greater than 12 cm total length (TL) recruited to

southern, offshore reefs.

In Narragansett Bay, YOY tautog appear from late

June through August (Dorf 1994; Dorf and Powell 1997).

During winter, juveniles remain inshore (Cooper 1964;

Stolgitis 1970; Olla et al. 1974). At this time, especially

when water temperatures are 5°C or less, YOY and 1-yr-old

tautog are found in discarded beverage cans and bottles,

eelpots, empty oyster and clam shells, and in crevices of

vertical structures; they are also found on their sides in

nearby depressions in sediments, either covered with a few

millimeters of sand and silt (except head and gill area) or

exposed on the sediment surface (Bigelow and Schroeder

1953; Cooper 1964, 1966; Olla et al. 1974, 1978, 1979, 1980).

Juvenile tautog (3-7 cm TL) were found to select a narrow

range of hole sizes (i.e., about 3-4.5 cm in diameter) in structured habitats, and to occur usually near the bottom of any

object or structure (Dixon 1994). Dorf (1994) suggested

that sheltering habitat could be a limiting distribution factor

in Narragansett Bay for juveniles less than 2 yr old. Although the food-producing capability could be surmised

as one of the reasons that older juveniles prefer reef-like

habitats, Dixon (1994) found that neither the availability of

attached food (e.g., small mussels) nor the presence of small

but not highly active predators (e.g., toadfish and sculpins)

had much effect on habitat use.

Olla et al. (1979) reported that young tautog (less

than 25 cm) showed an affinity to particular shelters, establishing homesites from which they ranged only a few meters

during the day, and to which they returned at night. They

also suggested that use of such homesites by some juveniles is seasonal. They observed a perennial homesite -- a

basin wall -- which was utilized year-round by part of the

juvenile population. At the onset of winter, other young

tautog that had possibly dispersed to other nearby summer

sites, such as eelgrass or algae beds, rejoined the juvenile

colony at the wall to overwinter. In the spring, some of this

population again left the wall for other shelter.

The mouths of estuaries or inlets may be especially

important habitat areas for juvenile and adult tautog. Briggs

(1975) reported that tautog was the most frequently collected fish in traps used on the Kismet artificial reef just

inside Great South Bay, Long Island. The species ranked

only fourth in abundance on an artificial reef less than 5 km

offshore.

Adults

Adult (i.e., mature) tautog (generally greater than 25

cm TL) have the same basic habitat requirements as larger

juveniles and are found in vegetation, rocks, natural and

artificial reefs, pilings, jetties and groins, mussel and oyster

beds, shipwrecks, submerged trees, logs and timbers, and

similar complexly structured coastal habitats [e.g., see Smith

(1907) and Hildebrand and Schroeder (1928)]. The fish are

extremely local, so much so that when fishing for them “a

few feet one way or the other may mean the difference between success and failure” (Bigelow and Schroeder 1953).

Adams (1993) observed that the species preferred the crest

and outer edges of coastal reef habitats.

During summer, adults can be found inshore, co-existing with younger fish. Olla et al. (1974) reported adults

ranged up to 500 m away from their homesite during the

day, but generally returned to the same general shelter area

at night. This established a local population during the

summer.

In late fall, when water temperatures fall below 11°C,

there is an overall migration to perennial offshore areas

with rugged topography in waters 25-45 m deep (Cooper

1966). Individual tautog do not appear to return annually to

the same sites within an offshore area to overwinter (Olla et

al. 1979). On the other hand, some adults were found to

overwinter inshore, especially in the north.

 The seasonal migrations of tautog do not seem to

involve great distances [e.g., Briggs (1974) reported that

fish from southern Long Island bays winter in deeper coastal

waters off northern New Jersey]. Other adult tautog in
Page 5

winter were observed in less than 10 m of water in eastern

Long Island Sound (Zawacki and Briggs 1976; Auster 1989),

in 10 m of water at an artificial reef 2.4 km off Delaware

(Eklund and Targett 1991), and in deeper areas of Chesapeake Bay (Hostetter and Munroe 1993). Eklund and Targett

(1990) noted populations in 25-35 m of water 22-37 km offshore of Maryland and northern Virginia, and Hostetter and

Munroe (1993) observed populations in 10-75 m of water to

65 km offshore of southern Virginia. At an artificial reef in

about 20 m of water 15 km offshore of southern Virginia,

Adams (1993) observed that the largest tautog (greater than

75 cm in length) occurred in February when temperatures

were about 6°C. He considered tautog to be a “core resident species” on this reef, and to be active year-round.

At very low temperatures, tautog enter a torpor-like

state (Cooper 1966; Briggs 1977). Curran (1992) reported

that cunner go into true torpor or hibernation during winter,

and suggested tautog do likewise, at least in the northern

part of their range. She believed the ability of cunner and

probably tautog to hibernate may be the key to these two

species, of a basically tropical family, being able to tolerate

cold water and inhabit cool-temperate waters year-round.

DETERMINANTS OF DISTRIBUTION

Juveniles

Olla et al. (1979) suggested that the distribution of

habitat use by juvenile tautog was somewhat seasonal and

temperature mediated. Juveniles seem to have a perennial

site which they use during the winter. In spring, some of

the wintering groups dispersed to summer sites. Olla et al.

(1979) believed this springtime dispersal was possibly due

to increased aggression in the population or to other factors which made the perennial winter site suboptimal. If a

summer habitat became suboptimal, because of elevated

temperatures or inadequate vegetation, juvenile tautog

moved to other perennial summer sites.

Young tautog have strong adherence to homesites

(Able and Fahay 1998), but if shelter becomes suboptimal,

they will move (Olla et al. 1979). They will also move if a

more attractive habitat is found. In the Great Bay - Mullica

River Estuary, Sogard (1989) found that in summer juveniles moved from naturally vegetated homesites to newly

planted, artificial seagrass beds. The factors that made the

artificial seagrass beds more attractive are not known.

Adults

Adults also show an affinity to homesites, though

this is not as strong as for juveniles. Adults have been

observed to leave a site readily if suboptimal conditions

develop. At an artificial reef in Delaware Bay, tautog which

were common in early summer were absent later that year,

coincident with a blue mussel (Mytilus edulis) kill, probably caused by high water temperatures (personal observation by senior author). Also, at an artificial reef 15 km off

Virginia, Adams (1993) noted a decrease in abundance of

large tautog when water temperatures were above 20°C for

an extended time. He suggested that large fish leave areas

with uncomfortably high temperatures for areas with cooler,

deeper waters.

The adult fall offshore migration is triggered by bottom water temperature dropping below 10°C (Cooper 1966;

Olla et al. 1974; Lynch 1994). The spring inshore migration

is associated with an increase in bottom water temperatures

to 11°C or above (Chenoweth 1963; Cooper 1966; Stolgitis

1970; Olla et al. 1974; Briggs 1977; Olla et al. 1979). In

laboratory experiments, Olla et al. (1980) confirmed temperature, not changing photoperiod, as the leading factor

in fall migratory movements and distributions.

HABITAT NEEDS

Tautog are specifically associated with complexly

structured habitats in all post-larval stages of their life. As

juveniles, these habitats include submerged vegetation,

shellfish beds, and three-dimensional objects or structures

with appropriately-sized crevices and holes for shelter. (See

the “Differential Distribution” section in this chapter). As

the fish grow, larger complex structures are needed for shelter, and hard substrates are usually required to support the

epibenthic or encrusting invertebrates upon which the fish

generally feed. (See the “Feeding and Diet” chapter). South

of Long Island, beyond where rocks and boulders were

deposited during previous glacial periods, there are few

natural rock outcroppings in coastal marine waters to provide the “reef” habitat that tautog require, although shellfish beds in euryhaline parts of estuaries serve as habitat

(Arve 1960). In this area, the man-made “reef” habitat created by coastal jetties, groins, pilings, accidental shipwrecks,

and intentional deposition of solid material as artificial reefs

is undoubtedly important to the distribution of the species.

The availability of new macroalgal growth as cover

in the late spring to early summer period can be critical to

settlement and survival of post-larvae (Dorf 1994; Dorf and

Powell 1997). This is especially true for areas that do not

support extensive eelgrass beds or complexly structured

habitats. Although macroalgae is normally degraded or

swept into dense beds later in the season, a healthy, newgrowth, spring-summer macroalgal community can be important to the initial survival of juvenile tautog (Dorf 1994;

Dorf and Powell 1997). Sogard and Able (1992) reported

that juvenile tautog in New Jersey appeared to prefer habitat where sea lettuce (Ulva) was present. Ulva can have

negative effects on certain taxa or species, and the preference or tolerance of Ulva by tautog can give it an advantage with competitors or predators (Dorf 1994). In contrast

to studies that report the importance of vegetation, Dixon
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 resembles the female (Olla and Samet 1977; Hostetter and

Munroe 1993). It has been suggested that the nondimorphic

male may be: 1) a sexual stage in the life of the tautog, 2) an

indicator of hermaphrodism, 3) a means to increase spawning opportunities, or 4) coincident with a different reproductive behavior than the dimorphic male (Hostetter and

Munroe 1993).

FECUNDITY

Chenoweth (1963) found that tautog that were 21-68

cm in fork length (FL), weighed 170-5207 g, and were between 3 and 20 yr old, contained 5000 to 637,500 mature

eggs. He found that the number of eggs (Y) were related to

fork length (X) in millimeters by the regression: Y = -6.00307

+ 3.0960(X). The number of eggs (Y) was also related to

weight (Z) in grams by the regression: Y = 0.31492 +

1.07993(Z). The number of eggs produced per unit of weight

per ovary reaches a maximum in 7-9 yr old fish which are

about 34-39 cm TL (Cooper 1967). Thereafter, egg production stabilizes until an age of about 16 yr, after which it

declines (Chenoweth 1963; Cooper 1967).

SPAWNING

The ratio of the mass of gonadal tissue to the mass

of all body tissues [i.e., the gonadal-somatic index (GSI)]

can be an indicator of a fish’s reproductive state, with the

highest indices just prior to spawning, and the lowest indices just after spawning. The GSIs for female and male tautog off the coast of Maryland and Virginia were reported to

peak between April and June (Eklund and Targett 1991;

Hostetter and Munroe 1993).

Spawning begins when water temperatures reach 9°C

or above, generally peaks about June, and continues

throughout summer (Kuntz and Radcliffe 1918; Nichols and

Breder 1926; Perlmutter 1939; Bigelow and Schroeder 1953;

Wheatland 1956; Chenoweth 1963; Cooper 1964; Colton et

al. 1979; Eklund and Targett 1990; Monteleone 1992; Sogard

et al. 1992; Hostetter and Munroe 1993; Malchoff 1993).

Sogard et al. (1992) used NEFSC-MARMAP larval distribution (Berrien and Sibunka, in press) and other data to show

spawning began in May-June south of New York, and

reached its peak in JuneJuly off Southern New England. Dorf (1994) reported

finding hatching eggs in late May to late July in

Narragansett Bay. Sandine (1984) reported the occurrence

of tautog eggs in Barnegat Bay (New Jersey) from March to

August, and questionably in October.

COURTSHIP

Tautog spawn in heterosexual pairs or in a group

with a single female being active simultaneously with several males (Olla and Samet 1977; Dixon 1997). The mode of

spawning is reported to depend on the number of mates

available for the female, the presence of a male-dominance

hierarchy, and environmental factors such as availability of

shelter and food (Olla and Samet 1977). In laboratory studies with two active males (one of which was dominant) one

female, and one shelter, mating occurred only between the

dominant male and the female (Olla and Samet 1977). Group

spawning was observed when there was either an increase

in the number of males, a lack of a dominant male, an increase in the male:shelter ratio, an inability to control a territory, or an elevated temperature (Olla and Samet 1977; Olla

et al. 1981).

Several weeks before spawning, male aggressiveness

toward females noticeably decreased and was replaced by

“rushing” (i.e., males quickly approached females then

veered off) (Olla and Samet 1977). Coincident with this

change in male behavior, females increased their girth with

enlargement of their ovaries.

In laboratory studies, courtship activity prior to

spawning was observed to continue for several hours. In

paired spawning, the male rushed the female frequently (Olla

and Samet 1977), and male pigment bars darkened while

those of the female lightened (Bridges and Fahay 1968).

Physical contact between the sexes, including nuzzling and

rubbing flanks, was observed (Bridges and Fahay 1968).

Courtship culminated with the paired fish moving rapidly

together within 1 m of the water’s surface. The fish then

turned toward each other, arched their bodies, and released

their gametes near, or as they broke, the water surface (Olla

and Samet 1977). Spawning usually occurred in the afternoon, with as many as three spawnings daily (Olla and Samet

1977, 1978). Spawning often continued into evening in the

wild (Ferraro 1980).

Olla and Samet (1977) observed slight differences in

courtship during group spawnings. The males did not rush

the female as frequently, and “contact-clustering behavior”

occurred (i.e., two or more males clustered near the female

and contacted her with their flanks).
Page 7

DEVELOPMENT

OVA

The development of the ova was described by

Chenoweth (1963):

The ova arise from the germinal epithelium

that lines the interior wall of the ovary and

fills a large part of the organ through convolutions. At first the developing ova are opaque,

relatively hard, and average 0.36 mm in diameter. As development proceeds the ovary enlarges and transparent, soft, mature ova appear interspersed with the immature ova. These

ova average 0.79 mm in diameter.

There are corresponding changes in the appearance of the

ovary as the eggs mature.

EGGS AND EMBRYOS

Spawned tautog eggs are about 1 mm in size, highly

transparent, and spherical in form (Herman 1963). The egg

membrane is thin but tough and the yolk sphere does not

contain an oil globule (Kuntz and Radcliffe 1918). Egg size

varies from year to year, with diameters of 0.70-1.18 mm

being reported (Nichols and Breder 1926; Richards 1959;

Chenoweth 1963; Lebida 1969). Egg size also decreases

with an increase in water temperature, and as the spawning

season progresses (Williams 1967). Therefore, it has been

suggested that annual variation in egg diameter reflects

differences in sampling times. Eggs that were reported in

the literature with variable diameters could be explained by

their being collected at different times during the spawning

season in different years (Auster 1989).

Incubation of fertilized eggs at temperatures of about

20-22°C in the laboratory was reported to take 42-48 hr to

hatching (Kuntz and Radcliffe 1918; Merriman and Sclar

1952; Perry 1994), but at 14.2-16.8°C, it took about 81 hr

(Perry 1994). Kuntz and Radcliffe (1918) and Fahay (1983)

described embryonic development.

D. Perry (pers. comm.4

) reports that the proportion of

normally developing, viable embryos among all tautog embryos collected in central Long Island Sound near New

Haven, Connecticut, declined as the spawning season progressed. This decline had no significant locality variance,

nor appeared to be related to differences in habitat quality.

This progressive decline was also noted in laboratory

spawned eggs and reared embryos (Perry 1994). Olla and

Samet (1978) found higher ambient temperatures affected

normal development of tautog embryos.

LARVAE

Newly hatched embryos, or yolk-sac larvae, are approximately 2.2 mm in length, but can be slightly less. [Some

variance in the lengths of specimens reported in the literature and reported here can be the results of preservation

which usually causes some shrinkage. This shrinkage can

vary with developmental stage (e.g., degree of ossification).]

The head is slightly deflected and the yolk sac is relatively

large, elliptically ovate, and unpigmented. The vent is located about mid-length of the body. The depth of either

dorsal or ventral fin fold is less than the depth of the body

just posterior to the vent. The fin folds and the posterior

caudal region of the body remain free of pigment.

One day after hatching at temperatures of 20-22°C,

tautog larvae are about 2.0-3.0 mm in length. The yolk sac is

greatly reduced and the head is no longer deflected. The

chromatophores increase in size and show well developed

pigment processes, but are fewer in number than in the

newly hatched larvae; individual pigment cells merge to

form large chromatophores. The larvae have a distinct blackish color (Able and Fahay 1998).

Four days after hatching is a critical period for this

species. The yolk sac has been absorbed and the mouth

has been formed and is functional (Bigelow and Schroeder

1953). These post-yolk-sac larvae, which are now 3.2-3.5

mm in length, must begin planktonic feeding. Black chromatophores are uniformly distributed over the dorsal and

lateral aspects of the body, but the posterior caudal region

is free of pigment.

Larvae which are 5.0 mm in length show a relatively

greater increase in body depth and thickness than in length.

The distribution of pigment remains essentially the same as

4 days after hatching; however, the chromatophores increase

in size and number. Larvae which are 10 mm in length display well differentiated dorsal, anal, and caudal fins. Pigment distribution remain the same as in earlier ontogenetic

stages of development, but the number of chromatophores

and quantity of pigment increase.

Malchoff (1993) found that larvae in the New York

Bight ranged from about 2.0 to 7.0 mm TL. From his timeseries collections, he developed a larval age-length relationship: A = -1.877 + 8.535 L, where A = age in days, and L

= length in millimeters.

The mean duration of the larval phase of tautog was

reported to be 25.4 ± 3.4 (SD) days for an undefined area of

the Northwest Atlantic (Victor 1986). Sogard et al. (1992)

and Malchoff (1993), however, reported the larval phase to

be only about 20 days in the New York Bight and Great Bay

- Mullica River Estuary. Dorf (1994) and Dorf and Powell

(1997) reported the same period for Narragansett Bay, but it

can be less (Schroedinger and Epifanio 1997).
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 characteristics of adults, except for their often greenish background

color. The black chromatophores form heavily pigmented

areas which give the body a transversely banded appearance as well (Kuntz and Radcliffe 1918).

The fish lay down their first visible bone annulus in

the spring of the following year, usually in May. For example, in the opercular bone, this is evident as a sharp

transition from a translucent to an opaque zone. Annuli of

this type are formed on bony structures each spring (Cooper 1967).

ADULTS

Most male tautog mature by age 3, females by age 4

(Chenoweth 1963; Cooper 1967; Stolgitis 1970; Briggs 1977;

Hostetter and Munroe 1993). Chenoweth (1963) observed

that 55% of males were already mature by age 2, and 90%

were mature by age 3, in Rhode Island. This latter age

corresponds to a tautog length of about 26 cm, as determined by Hostetter and Munroe (1993). The latter authors

found that few females were mature by age 2, but 80% were

mature by age 3. Precocious females, approximately 2 yr

old, were observed in New York waters (Olla and Samet

1977) and in Mt. Hope Bay, (Massachusetts) (Hostetter and

Munroe 1993).

The influence of heavy fishing mortality on the genetic selection process for sexual maturity is unknown at

present; this pressure can favor the portion of the population with genes for early maturation and spawning, although

with lower gamete production.

AGE AND GROWTH

LARVAE

Malchoff (1993) reported larval growth rates for this

species to be 0.30 mm/day in the New York Bight. This rate

is lower than a mean 0.75 mm/day, back-calculated rate reported for Narragansett Bay fish by Dorf (1994). Martin

(1993) reported that larval growth rates varied with latitude

and were directly related to water temperature.

JUVENILES

Sogard et al. (1992) used length-frequency progressions, otolith age - fish size comparisons, and direct measurements of growth in cage experiments to determine that

YOY tautog grew at an average rate of about 0.5 mm/day

during the summer in the Great Bay - Mullica River Estuary.

The same rate was found in Narragansett Bay (Dorf 1994).

In the Great Bay - Mullica River Estuary study, mean growth

rates varied slightly among different methods of estimation

and with habitat types where the fish were collected. Sogard

et al. (1992) also found minor growth also occurred in other

seasons. Martin (1993) reported that YOY grew up to 0.7

mm/day when fed mysids in the laboratory.

During the 2-3 yr of the juvenile phase of this species, annual growth is usually rapid (Cooper 1967). Annual

growth is reported to increase progressively from northern

to southern regions (Martin 1993). For example, in

Narragansett Bay, Cooper (1967) reported first-year growth

in tautog to be 60-62 mm. This was less than half the firstyear growth of 134-146 mm by YOY tautog in southern Virginia waters (Hostetter and Munroe 1993). Tracy (1910),

however, reported first-year growth to range between 72

and 288 mm in Rhode Island; the larger value is obviously

questionable (Fritzsche 1978).

In Narragansett Bay and the Great Bay - Mullica River

Estuary, the greatest annual growth increment, 200%, occurred during the second year for both sexes (Cooper 1967;

Sogard et al. 1992). In southern Virginia waters, the greatest annual growth increment occurred in the first year, and

the second year’s growth was only a 30% increase (Hostetter

and Munroe 1993). It has been hypothesized that the faster

initial growth rates in southern waters reflect the longer

duration of warmer water temperatures, which may provide

optimal conditions for tautog growth during the first juvenile year, but for some reason not the second year (Sogard

et al. 1992; Hostetter and Munroe 1993).

Sogard (1992) observed that the mean seasonal

growth in length of juvenile tautog in New Jersey was higher

in vegetation, but growth in mean weight was higher on

bare sand. Growth rates were higher in sea lettuce beds

than in eelgrass beds, and seemed to be directly related to

prey density and not shelter type.

ADULTS

Although Cooper (1967) noted that Rhode Island females attained a slightly greater mean length than males (62

mm compared to 60 mm) during their first juvenile year, by

age 3, as they became adults, males had faster annual growth

rates. At age 7 in Rhode Island, the male tautog’s mean

length was 348 mm, while the female’s mean length was 301

mm. Faster adult male annual growth rates were also found
Page 9

by Simpson (1989) in Long Island Sound. In southern Virginia waters, males also grew faster than females at all ages

(i.e., K = 0.090 for males compared to K = 0.085 for females5)

(Hostetter and Munroe 1993).

Hostetter and Munroe (1993) found that tautog attained a relatively large size slowly, and that their growth

varied seasonally. These authors reported that in southern

Virginia’s coastal waters, maximum somatic growth occurred

after spawning, from July to December. Slower growth occurred from January to March because of decreased feeding associated with cool water temperatures and the fish’s

associated torpor condition in colder northern waters. In

northern waters, this period of slower winter growth is also

probably longer because these waters cool sooner and remain cold longer. In southern Virginia waters, the least

somatic growth in adults occurred from March to June during gonadal maturation and spawning.

Mean annual growth rates in length were similar for

tautog in northern and southern waters until about age 13,

then growth rates decreased more rapidly in northern waters. For males in Rhode Island, annual growth increments

decreased to less than 12 mm after age 12, and further declined to 2-4 mm after age 20. For females in Rhode Island,

annual growth increments decreased to less than 11 mm

after age 13, and to 3-4 mm after age 17 (Cooper 1967). In

southern Virginia waters, increments in annual growth declined after age 13, but growth rates were nearly double

those of tautog in northern waters (Hostetter and Munroe

1993). The von Bertalanffy growth equation for tautog from

both areas support this finding: in northern waters, L = 506

mm for females and 664 mm for males (Cooper 1966), compared with L = 733 mm for females and 732 mm for males in

southern waters (Hostetter and Munroe 1993).

Several studies reported variable length-weight relationships for whole and eviscerated tautog from

Narragansett Bay to southern Virginia waters (Table 1).

Tautog are a relatively long-lived fish, with the oldest fish examined by the scientific community estimated to

be a 34-yr-old male (Cooper 1967). A 91-cm (36.5-inch),

10.1-kg (22.5-lb) fish, caught off New York in 1876 and reported by Goode (1887), could have been older. Hostetter

and Munroe (1993) reported that the world’s record weight

for the species is 10.9 kg (24 lb) for an 81.9-cm fish caught

recently off Virginia; the authors estimated the fish to be

about 30 yr old. Simpson (1989) suggested that males can

live more than 30 yr, and females about 25 yr.

FEEDING AND DIET

FEEDING BY LIFE STAGE

Larvae

No specific data were found; larval tautog probably

feed on small motile crustaceans in the water column such

as small copepods [which were part of the diet of small

juvenile (30-40 mm) tautog reported by Grover (1982) and

Dorf (1994)] and other larval fish. D. Perry (pers. comm.6)

reported some success in getting larvae to eat dried food.

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 report “seaweed” as part of the diet of juvenile tautog in this

area.

Grover (1982) found that both caprellid and gammarid

amphipods and small copepods constituted 95% of the diet

of YOY (31-71 mm TL) tautog in southern Long Island waters. Frequency of occurrence (FO) in the digestive tract

was 98% for amphipods, 94% for copepods, 29% for polychaetes, and 25% for isopods. This diet indicated that juvenile tautog forage on benthic, as well as planktonic, prey.

As they grow, juvenile tautog rely less on planktonic food

resources and feed primarily on benthic prey, although the

prey remain primarily crustaceans, such as decapods. The

general form and location of the tautog’s mouth suggest

that benthic organisms would be a primary component of

the adult’s diet (Grover 1982). In Great South Bay (New

York), 2-3 yr old tautog (105-206 mm) did not feed mostly on

crustaceans throughout the year, but on blue mussels (Olla

et al. 1975). This diet difference is partially consistent with

the laboratory prey selection findings reported by Lankford

et al. (1995) that juvenile tautog gradually shift their diet

from small crustaceans (e.g., amphipods) to small mussels

as the fish grow to and beyond a length of about 120 mm.

Mussels that were about half of their maximum size were

primarily eaten.

In the Great Bay - Mullica River Estuary, Sogard

(1992) found that copepods were the preferred prey, with a

78% FO, in 31-85 mm YOY fish. Amphipods were a close

second in importance (FO = 75%), followed by other crustaceans (FO = 40%), polychaetes (FO = 5%), and mollusks

(FO = 5%). Festa (1979) reported that isopods (Idotea sp.

and Erichsonnella sp.), xanthid (mud) crabs, and (to a lesser

degree) several species of amphipods were the prey of 60-

160 mm tautog from Great Egg Harbor (New Jersey). He

also reported that larger subadult tautog, greater than 200

mm in length, ate predominantly xanthid crabs.

In eelgrass beds of Chesapeake Bay, juvenile tautog

(90-170 mm) fed on penaeid shrimp, blue crabs (Callinectes

sapidus), isopods, grass shrimp (Paleomontes sp.), and

detritus (Orth and Heck 1980).

Lindquist et al. (1985) reported that mytilid mollusks

and gammarid and caprellid amphipods dominated the diets

of 22 tautog (95-270 mm) examined from rock jetties near

Wrightsville Beach (North Carolina); they also noted the

occurrence of algae in the stomachs. Troutman (1982) reported that the dominant prey in these North Carolinian

diets varied seasonally, and included a venerid clam.

MacKenzie (1977) also reported juvenile tautog to prey on

a juvenile venerid, the northern quahog (Mercenaria

mercenaria).

Adults

Adult tautog are durophagous and feed chiefly on

blue mussels and other shellfish throughout the year. Barnacles (Balanus sp.), brachyuran crabs, hermit crabs, sand

dollars, (bay?) scallops, amphipods, decapod shrimp, isopods, lobsters, and probably nereid polychaetes were reported by Bigelow and Schroeder (1953) to be part of the

tautog’s diet. Consistent with this broad summary, Osburn

(1921) reported that “tautog feed on bryozoa along with

other hard shelled organisms to which it was attached,”

and Verrill (1873) reported that tautog fed on benthic tunicates. Bleakney (1963) reported tautog from Nova Scotia to

feed not on blue mussels, but on horse mussels (Modiolus

modiolus) and periwinkles (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 Atlantic rock crabs (Cancerirroratus) 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 volumetric basis at a northern New Jersey artificial reef. Festa

(1979) reported that xanthid crabs dominated the diet of

small adults (about 300 mm) in Great Egg Harbor Estuary.

Unpublished data collected by the senior author

showed that besides blue mussels (54%), Metridium anemones (11%), Atlantic rock crabs (5%), and razor clams (4%)

dominated overall definable prey in 358 tautog stomachs,

by volume, in a 1990-94 Delaware Bay artificial reef study;

these fish ranged from 110 to 580 mm, with a mean length of

320 mm. The proportional composition of the diet varied

among years (e.g., the mussels varied from 13 to 87% of

total diet volume). Various small mollusks, barnacles, decapod crabs, and other crustaceans of suitable size were reported also to be the food of this species in the Chesapeake

Bay area (Hildebrand and Schroeder 1928).

Richards (1992) confirmed Smith’s (1907) note that

tautog will eat commercially important Jonah crabs (Cancer

borealis) and small American lobsters (Homarus

americanus). Predation on these species occurred, however, when they were without shelter in aquaria.

Chao (1973) reports that the cunner lacks a well defined stomach, which is a characteristic of most labrids,

including tautog. The tautog’s gastro-intestinal tract is semi-
Page 11

transparent and remarkably thin for containing the broken

shells of its prey as it moves through to evacuation.

FEEDING BEHAVIOR

Tautog feed throughout the daytime. Beginning soon

after sunrise, tautog were reported to leave their shelters to

forage for food, which involved scan-and-pick feeding

(Briggs 1969b; Olla et al. 1975). This activity sometimes

took the adults up to 500 m from their homesites (Wicklund

1966; Olla et al.1974). Bigelow and Schroeder (1953) observed that tautog followed the flood tide up above low

water levels, around ledges, to prey on mussels in the intertidal zone, and returned to deeper water during the ebb tide.

Feeding continued to evening twilight (Olla et al. 1974).

Olla et al. (1974) reported that tautog required about 8 hr to

process and evacuate food.

In laboratory studies, Olla et al. (1974) observed that

tautog grasped mussels with their anterior teeth and tore

them from their attached substrate with a lateral shaking of

the head. Small prey were swallowed whole (Bigelow and

Schroeder 1953), while larger, hard-shelled ones were

crushed by pharyngeal teeth before swallowing (Bigelow

and Schroeder 1953; Olla et al. 1974). The anterior teeth

were not involved in the crushing process (Olla et al. 1974,

Liem and Sanderson 1986). It was observed that the tautog’s

mouth can accommodate larger clumps of mussels than the

pharyngeal teeth can process efficiently. In this case, the

fish ingests and egests the clump from its mouth, separating it in the process into smaller, crushable sizes. The feeding and mastication methods of this species are specialized

and typical of labrids and cichlids only (Liem and Sanderson

1986).

In the southern part of its range (i.e., below New

Jersey), the blue mussel, a dominant prey in colder waters,

is at its warm-temperature limits and has wide variability in

recruitment and abundance (Foster et al. 1994). Without

strong periodic recruitment, mussel populations in these

areas can be preyed upon by tautog to near extirpation, as

reported for Virginia (Chee 1977; Chesapeake Executive

Council 1994). This near extirpation creates a change in the

prey field available to tautog in this habitat, and can cause

at least some of the tautog population to seek alternate or

better foraging areas. This change in prey field can be a

factor in the local distribution of the population. (See the

“Adults” subsection, “Differential Distribution” section of

the “Distribution and Habitat” chapter.)

FACTORS AFFECTING FEEDING

Tautog find prey visually and were reported not to

feed at night (Olla et al. 1974; Deacutis 1982). Neither do

they actively feed in northern waters during the coldest

part of the year (Cooper 1966; Curran 1992). In Narragansett

Bay, Cooper (1966) observed that of 15 tautog, 13 had

shrunken and emptied digestive tracts in winter. The other

two fish had remains of Atlantic rock crabs in their stomachs. Curran (1992) found all feeding stopped in the closely

related cunner (and probably tautog) when water temperatures reached the low levels that induce torpor and hibernation. Cunner, at least, survived up to 6 mo without food,

using glycogen, lipids, and proteins stored in their livers

(Curran 1992); tautog may do likewise.

In laboratory experiments, a decrease in feeding was

observed with an increase in water temperature above certain levels. Tautog which had been acclimated to temperatures of 19°C and 21°C, respectively, decreased their ingestion of food, when water temperatures were increased to

28.7-33.0°C over a certain amount of days (Olla and

Studholme 1975; McCormack 1976; Olla et al. 1978).

McCormack (1976) also reported that it required up to 7

days for field-collected fish to begin feeding in laboratory

aquaria. (For other environmental effects on feeding, see

the “Habitat Modification and Loss” section of the “Natural and Human-Induced Environmental Factors” chapter.)

Tautog were observed to vary, to some extent, their

feeding in association with their place in a group dominance hierarchy. In one laboratory study with three fish,

the dominant fish of the moment ate the greatest amount of

food, followed by the subordinate fish, in some order of

rank (Olla et al. 1978). In a related tank study, McCormack

(1976), however, found no difference in consumption of

Atlantic surfclam (Spisula solidissima) meats between the

dominant and subordinate individuals of paired 160-280 mm

tautog.

Food intake in tautog may decrease during spawning. Bridges and Fahay (1968) reported possible courtship

behavior of tautog in a laboratory study during which no

spawning occurred. In this study, 1 day prior to courtship

behavior, the male and female daily food (undefined) intake

decreased from 40 to 1g. During courtship behavior, which

lasted 2 wk, the 300-mm female ceased eating, but the 270-

mm male increased food intake to 10 g/day. After courtship

behavior ceased, the female resumed eating, but less than

10 g/day. The time period that this reduced level of feeding

persisted, or if it changed, was not reported. It is possible

that some of this behavior was an artifact of the fish’s confinement in aquaria, as it has not been documented in the

wild.

Deacutis (1982) found tautog did not have an acute

sense of smell for detecting prey, compared to red hake

(Urophycis chuss), and were hesitant to explore open bottom to find food they could not see.

BEHAVIOR

MIGRATION AND LOCAL MOVEMENTS

Many studies report limited, seasonal, onshore and

offshore movements of this species. (See the “Differential

Distribution” section of the “Distribution and Habitat” chap-
Page 12

ter.) Channels may be important pathways of migration into

coastal areas for spawning and out again to deeper or offshore wintering areas (Cooper 1966). As noted previously,

Briggs (1977) reported that part of the southern Long Island tautog population migrated to the area off northern

New Jersey when water temperature declined in the late fall;

Nichols and Breder (1926) called this movement “heavy.”

At a more local level, Merriman (1947) reported that small

tautog do not move with the high-tide levels into intertidal

areas; this contrasts with Bigelow and Schroeder’s (1953)

report of intertidal-associated movements of presumably

adults for feeding on tidally submerged mussels, as noted

earlier.

SCHOOLING

Some degree of schooling is thought to occur as tautog are reported to congregate during or just prior to spawning (Bigelow and Schroeder 1953; Cooper 1964; Stolgitis

1970). Briggs (1969b) believed that tautog also congregate

into some type of a school before and during their movement to deeper water in the fall off New York. This is supported by laboratory studies where tautog schooling was

observed when water temperatures declined to 6.3°C (Olla

and Studholme 1975).

DAILY AND SEASONAL ACTIVITY

LEVELS

In laboratory studies, tautog showed a high level of

activity (i.e., movement) within the first hour of daylight

(Olla et al. 1978). This activity decreased toward mid-morning, when the fish rested on the sand or within shelters

between bouts of activity (Frey 1963). Bigelow and

Schroeder (1953) noted that when tautog were not feeding,

they gathered in holes or crevices, being still or lying on

their side, often grouped together, until a rising tide stirred

them to activity (for intertidal feeding, presumably). In field

investigations at night, Olla et al. (1974) found tautog to be

inactive in a shelter of some type, and in a state of such low

responsiveness that individuals were readily touched by

divers. Curran (1992) suggested that this state represents

true sleep in cunner (and probably in tautog), and can be

for energy conservation.

Tautog in northern waters overwinter in a torpid state

within shelter (Nichols and Breder 1926). Olla et al. (1974)

observed small fish (less than 250 mm TL) in a torpid condition at water temperatures less than 6°C. Cooper (1966)

observed adult tautog being dormant, either lying on their

sides or in upright positions in crevices, at a water temperature of 7.5°C. In laboratory studies, tautog activity decreased as temperatures declined to 5.2°C, then tautog aggregated around shelters or burrowed under objects on the

bottom (Olla et al. 1980). At water temperatures of 2.1°C or

below, activity ceased and tautog remained in torpor. Curran

(1992) considered this torpid state to be true hibernation in

cunner. Emergence from torpor occurred when temperature

increased to 4.0°C (Olla and Studholme 1975). In southern

Virginia waters, tautog were observed to remain active despite water temperatures as low as 7°C (Adams 1993). In

laboratory studies, young and adult tautog responded to

an increase in temperature, from 19.1°C or 21.3°C to 28.7°C,

with a decrease in aggressiveness and activity, and an increase in the tendency to aggregate (McCormack 1976; Olla

et al. 1978). This lowered response to stimuli at temperature extremes was supported by the sound-detection-threshold experiments of Offutt (1971).

HIERARCHICAL DOMINANCE

In laboratory studies, a dominance hierarchy was observed among variably sized pairs or a group of tautog (Olla

and Studholme 1975; McCormack 1976; Olla et al. 1978). In

these studies, the larger, most aggressive or dominant fish

caused a less aggressive, usually smaller subordinate to

assume a submissive posture, tilting its dorsal surface toward the dominant fish at an angle. This behavior usually

occurred within 1-3 m of the dominant male. Field studies

have not confirmed these observations.

VOCALIZATION AND RESPONSE TO

SOUND

Fish (1954) found that tautog produced deep thumping, grunting, or barking sounds, as well as sounds from

crushing shells of their prey when feeding. The nonfeeding

sounds were heard in aquaria when fish were startled or

exposed to experimental electrical impulses, and were not

thought to be produced commonly. The air bladder was

thought to be involved in this sound production. The purpose of producing these sounds was unknown, but could

be defensive. Parker (1910) reported that tautog avoided

sources of loud sounds. Offutt (1971) used conditioned,

cardiac responses to show that fish can respond to sound

impulses.

COLOR MODULATION

Behre (1933) reported that small tautog responded to

changes in light intensity and color by changes in their

body color or shading. Absence of light reduced the intensity of body color, while high-intensity light enhanced the

body’s dark coloration. Differently colored light and sequential combinations of colored lights caused different

responses in body coloration. These responses suggested

to Behre some ability by tautog to distinguish colors or

shading. Regan et al. (1982) reported that the photoreac-
Page 13

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 Human-Induced Environmental Factors” chapter.)

MUTUALISM

Although many tropical and some temperate wrasses

(e.g., European wrasse) clean external parasites from larger

fish of other species (Darwall et al. 1992), this behavior has

not been reported for any lifephase of tautog.

POPULATION STRUCTURE

SEX RATIO

Chenoweth (1963) noted that about 53% of the tautog he collected in Narragansett Bay were female. In northern Virginia waters, however, the slightly skewed sex ratio

(0.86:1) was in favor of males (Eklund and Targett 1990). A

significant deviation from a 1:1 ratio was also observed in

larger fish by Hostetter and Munroe (1993), possibly because the older fish (greater than 18 yr old) were predominantly males (Cooper 1967; Hostetter and Munroe 1993).

AGE COMPOSITION

As noted previously for Narragansett Bay, the oldest

male fish found by Cooper (1966) was estimated to be 34 yr,

and the oldest female 22 yr. In southern Virginia waters, the

oldest male fish collected was estimated to be 25 yr, and the

oldest female 21, although the age of the oldest fish taken

by rod and reel from Virginia waters was estimated to be

approximately 30 yr (Hostetter and Munroe 1993).

In the early 1960s, 5- and 6-yr-old tautog were the

most abundant age classes in the Narragansett Bay population (Cooper 1967). The recent average age for tautog in

Virginia waters was 4 yr (Hostetter and Munroe 1993). In

1960s in Narragansett Bay, 79% of the tautog were less than

10 yr old (Cooper 1967). This population structure remained

similar for Massachusetts to New York into the mid-1990s

(Lazar 1995). This age structure is also similar to southern

Virginia waters where 82% of the population was recently

noted as being less than 10 yr old (Hostetter and Munroe

1993).

NATURAL AND HUMAN-INDUCED

ENVIRONMENTAL FACTORS

PREDATORS

Perry (1994) reported from laboratory experiments that

typical coastal copepods, such as Acartia tonsa, can prey

upon newly hatched, yolk-sac tautog larvae. Bigelow and

Schroeder (1953) reported the following fish species were

known to eat juvenile or adult tautog: smooth dogfish

(Mustelus canis), barndoor skate (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 cryptic predators common in coastal waters, as well as larger

YOY bluefish, preyed upon YOY tautog confined in aquaria.

Other demersal piscivorous fish, such as conger eel (Conger oceanicus), summer flounder (Paralichthys dentatus),

and various sharks, can be expected to prey upon small

tautog. Although American lobsters share the same habitat with tautog, and are active at night when tautog are

inactive, there are no reports of lobster predation on tautog.

It is also likely that diving or stalking piscivorous

birds, such as cormorants, grebes, loons, herons, and egrets,

also prey upon juvenile tautog in shallow estuarine or

coastal areas during daylight and the warm seasons when

the fish are outside their shelters (Whoriskey 1983). Bent

(1919, 1922, 1923, 1926) noted that small estuarine fish were

eaten by these birds, with labrids being specifically mentioned in the diets of the black-crowned night-heron

(Nycticorax nycticorax) and the double-crested cormorant

(Phalacrocorax auritus). Nichols and Breder (1926) noted

a small tautog in the stomach of a red-throated loon (Gavia

stellata). Another observation of this predation can be

recorded, with an approximately 12-15 cm tautog being observed in the beak and swallowed by a cormorant near a

rocky sea wall in Sandy Hook Bay (New Jersey) (personal

observation by the senior author in July 1992). The relative

importance of this predation on population structure, habitat use, and growth or mortality rates is unreported or unknown.

Humans as fishermen are also predators. Simpson

(1989) estimated fishing mortality for a Long Island Sound
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 commonly on blue mussels, other small mollusks, and crustaceans, and because the latter three species have ranges

that overlap that of tautog (Hildebrand and Schroeder 1928;

Weiss 1970; Olla et al. 1975). Botton and Ropes (1989)

suggest that horseshoe crabs (Limulus polyphemus) can

consume large quantities of shellfish, including mussels,

and thus may also be a competitor for food. Sea stars, such

as Asterias, that prey heavily on mussels when they are

available near the seabed, will compete trophically with tautog for this food resource too (Richards 1963). Other invertebrate predators of mussels, other shellfish, and barnacles

can be substantial tautog trophic competitors when present

in abundance. Species that eat copepods and decapod

crustaceans should also be considered potential competitors of juvenile tautog for this essential prey resource (Dorf

1994).

Competition with cunner was thought to be restricted

to May and June, after which cunner change their diet (Olla

et al. 1975). In a laboratory study, cunner seemed to be

aggressively territorial in excluding similarly sized juvenile

tautog from certain habitats where they co-occur (McErlean

1963). The competition between tautog and cunner can

only seriously exist with juvenile tautog; larger tautog use

larger shelter and prey that are well beyond the range of

cunner. Dixon (1994) suggests, however, that YOY cunner

and tautog have differing, but overlapping, preferences in

the location and use of their microhabitats that can reduce

competition.

DEFORMITIES AND ABNORMALITIES

Briggs (1966) reported on a mature female, estimated

to be 9+ yr old, that was pugheaded (i.e., exophthalmic eyes,

shortened and broadened maxilla, steep forehead, pushedin snout, incomplete closure of jaws, extended mandibles).

Several 250-300 mm tautog collected by hook-and-line in

Delaware Bay had notable sigmoid vertical curvatures of

their spines that compressed their body length (personal

observation by the senior author in June 1994). No ecological nor environmental factor has been implicated in these

deformities.

Tautog have been reported with small black spots

within their muscle tissue. Microscopic examination found

that these spots were actually associated with blood vessels, not the muscle tissue itself (I. Sunila, pers. comm.7

).

These spots were found to be deposits of dark pigment

surrounded by fibrosis. This pigment originates from their

high use of blue mussels as prey. These spots are considered harmless to the fish and to consumers.

PATHOGENS, PARASITES, AND

BIOTOXINS

Tautog exposed to water in the apex of the New York

Bight, near the then-active dredged material and sewage

sludge disposal sites, were found to have antigenic responses to certain human enteric bacteria. These responses

indicate that tautog are immunologically responsive to exposure to sewage bacteria (Stolen et al. 1983).

Finrot disease, thought to be caused by some microbial agent and anthropogenic stress, was not found on tautog in the New York Bight, although the disease was reported on 22 other common fish species in a study by

Mahoney et al. (1973). However, a subsequent study did

report, without supporting details, this condition on tautog

(Murchelano and Ziskowski 1982).

Cooper (1964) observed that 10% of the tautog in his

survey had scales that were either pitted or regenerated.

He considered this pitting or regeneration to be due either

to metacercaria of the trematode Cryptocotyl sp., or to physical abrasions. Laird and Bullock (1969) found no positive

evidence of hematozoan parasites in New England and Canadian tautog. Cheung et al. (1979) reported tautog to be

infected lethally with a coccidian intestinal parasite. Linton

(1899) found immature trematode distomes to be encysted

in abundance on the skin, fins, and eyes of a tautog collected off Woods Hole (Massachusetts), and also reported

cestodes, nematodes, and trematodes to occur in cunner.

Perlmutter (1952) reported summer mortalities of

“blackfish” along the ocean side of Jones Beach, southern

Long Island. These mortalities were speculated to be related to a nearly concurrent, offshore bloom of the dinoflagellate Noctiluca sp. A number of other fish species were

reported affected at this time as well.

HABITAT MODIFICATION AND LOSS

Structure

The dependence of tautog on specific coastal and

estuarine habitats can make any degradation or loss of these

habitats a serious threat to the tautog resource (Chesapeake Executive Council 1994). The loss of vegetated estuarine habitats can impair the habitat value of estuaries as

juvenile tautog (and other species) nurseries. However,

juvenile tautog have been observed using other habitats

and a variety of substrates, including artificial (e.g., pilings),

that provide shelter or concealment and access to food.
Page 15

(See the “Habitat Needs” section of the “Distribution and

Habitat” chapter.) Rocky reef habitats used by tautog are

very limited south of Long Island, and shipwrecks and other

subtidal, manmade material placed in this area have expanded

the distribution of this habitat into the common, open sandy

habitats of the Middle Atlantic coasts (Hastings 1978;

Lindquist et al. 1985). Tautog were collected around a sewer

outfall under construction off southern Long Island; their

peak abundance was during October through December

when bottom water temperatures were 7-13°C (Briggs 1984).

None of the tautog were found with any obvious pathological conditions or other possible effects from this physical habitat alteration and disturbance.

In the past, salvaging of certain metal shipwrecks or

lowering of wreck heights if they might be navigation hazards, reduced the value of shipwrecks as tautog habitat.

Before 1900, some wrecks off New Jersey were even dynamited to stun and collect reef fish (Smith 1892). Recently,

the use of certain, heavy, “rock-hopper-roller,” bottom trawl

gear over many older, more fragile wrecks, lower-profile reefs,

and mussel beds (DiLernia 1993) also threatens habitat quality, including the destruction of slow-growing colonies of

the northern star coral (Astrangia sp.) which provide biogenic reef habitat for tautog.

Reef habitats that lose structural height are more

prone to siltation (Rothschild et al. 1994) or burial by sediment movement. Briggs and O’Connor (1971) report that

tautog avoided estuarine areas that were sandfilled, as from

shoreline sand replenishment. The decline in the number,

distribution, and structure of oyster beds (Rothschild et al.

1994) is another threat to the estuarine habitat needs of

juvenile tautog and other species with similar habitat needs

(Chesapeake Executive Council 1994). The creation of new

“reef” habitat by the artificial reef programs of most states

(McGurrin 1989) where the species occurs may be mitigating this habitat degradation to some degree.

Hydrography

Auster (1989) reported that current velocity can affect small-scale spatial distribution and can change foraging behavior in tautog and other species. The high hydrographic energy caused by passage of a hurricane was reported to disrupt juvenile tautog distributions in

Narragansett Bay (Dorf 1994; Dorf and Powell 1997).

Howell and Simpson (1994) reported that tautog are

not tolerant of dissolved oxygen (DO) concentrations below 2 mg/l, and prefer DO levels above 3 mg/l. This intolerance is supported by the cardiac and preliminary respiration response results of Mallet (1972) who showed that

tautog are oxygen conformers under normal conditions, and

that there is no inflection threshold. He also showed that

these response rates increase with temperature and cadmium exposure, and that hypoxia responses can then be

detected at higher oxygen concentrations. Mallet also observed several tautog stress responses to hypoxia. These

responses included a change in mottled-dark body color to

a pallid white-gray, attempts to jump out of experimental

tanks, regurgitation of food, mucus secretion (especially

around the gills), and milt ejaculation in ripe males. Baldwin

(1923) reported some degree (undefined) of hypoxia “tolerance” at an unreported temperature for this species.

In a fish kill off the northern New Jersey coast in

1968, possibly caused by hypoxia, tautog were present in

the area, but not found affected, although its smaller relative, the cunner, were found dead or dying (Ogren and Chess

1969). Tautog mortalities, however, were reported by commercial fishers and were observed on the beach during a

major anoxia-hypoxia episode off New Jersey in 1976

(Azarovitz et al. 1979).

Smith et al. (1980) reported that high water temperatures were incrementally lethal in the laboratory for tautog

gastrula-stage eggs: 10% mortality at 16.8°C after 26 min,

50% mortality at 21.3°C after 50 min, and 90% mortality at

31.4°C after 55 min. These authors suggested that tautog

eggs enduring 15 min of a once-through cooling system of

a power plant, with a 10°C temperature rise from a base

temperature of 15°C, would suffer 10-50% mortality. Olla

and Samet (1978) reported that tautog eggs incubated above

20°C resulted in embryos with anatomical deformities, and

that those eggs incubated above 24°C resulted in increased

mortalities. Normal embryonic development resumed when

water temperature decreased to 20°C or lower. Laurence

(1973) reported that tautog yolk-sac larvae (“prolarvae”)

may develop an energy deficit at water temperatures above

19°C because of increased metabolic rate and use of their

yolk. This higher-than-normal rate of yolk use can result in

smaller larvae just when those larvae need to start feeding

on plankton. He believed these smaller larvae can be at a

competitive disadvantage with larger siblings and other

plankton predators, and may be less successful in obtaining suitable prey. This potential competitive disadvantage

could affect survival or lead to slower larval growth rates

and lowered capacity to reach protected inshore nursery

habitats.

As noted in the “Eggs and Embryos” section of the

“Development” chapter, the viability of tautog embryos has

been reported to decline as the spawning season progresses.

The cause of this decline is not known, but could be environmental, such as high temperatures (Olla and Samet 1978).

Pearce (1969) reported 31°C to be lethal to adult tautog in a 1-hr change from ambient during an aquarium study.

McCormack (1976), though, reported that tautog do not

lose equilibrium until about 33°C when water temperature

was raised more gradually. At above-ambient temperatures,

tautog behavior is altered; these alterations included

changes in activity, dominance, habitat use, and posture

that suggest stress (Olla and Studholme 1975; McCormack

1976); Olla and Studholme (1975) reported that the fish recover normal behavior when water temperatures return to a

normal ambient range.
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 concentrations decreased with length; in both sexes, nickel concentrations decreased. In both sexes, iron, manganese, and

zinc concentrations were not related to body length. Hall et

al. (1978) also examined 15 metals in this species (collection

location not defined) and found low to average levels in

muscle samples.

A recent study (National Marine Fisheries Service

1995) found muscle concentrations of nine metals (i.e., silver, cadmium, chromium, copper, nickel, lead, zinc, arsenic,

and mercury) to be well below U.S. Food and Drug Administration (FDA) action levels in tautog collected off

Manasquan Inlet (New Jersey). Mercury levels in tautog

were higher than those in bluefish, black sea bass

(Centropristis striata), and summer flounder, but still well

below FDA action levels.

Mallet (1972) reported that acute, sublethal (i.e., 20

ppm) cadmium exposures and intoxication caused a change

in tautog cardiac response to gradual hypoxia; this response

included higher variability in heart rates and lowered rateresponse sensitivity to oxygen levels, which suggest that

the metal interferes with the cardiac regulatory mechanism

for hypoxia. Lethal cadmium exposures (i.e., 400 ppm) resulted in death in about 5 hr, but produced no significant

histological damage to gills, liver, kidney, or intestine. In

another study, Mallet noted that cadmium levels on gill

tissue can be very high.

Studies of the effect of longer-term experimental exposure of cadmium on the closely related species, cunner,

found that this metal did cause pathological changes to a

number of organs and tissues (National Marine Fisheries

Service 1974; MacInnes et al. 1977).

Organics

Tautog fillets from the Hudson River - Raritan Bay

Estuary, collected between 1980 and 1985, were examined

for Chlordane, DDT, DDD, DDE, dibenzoanthracine, Endine,

Heptachlor, hexachlorobenzene, Lindane, and total polychlorinated biphenyls (PCBs). Concentrations of these organics were found to be low, relative to concentrations in

some other fish species examined (New York State Department of Environmental Conservation 1987). This general

result was confirmed in a recent study by the National Marine Fisheries Service (1995) which also found that all concentrations of PCBs and pesticides in tautog tissue were

below an FDA action level of 2.0 ppm. Polycyclic aromatic

hydrocarbons were largely undetected in the tautog tissue.

No information was found of the effects of exposure

to organics on tautog. Several studies are available for

cunner, however, the other labrid in the Northeast, the results of which could be similar for tautog. Payne et al.

(1978) found little pathological effect in cunner from a 6-mo

exposure to petroleum hydrocarbons. Payne and May (1979)

further reported that cunner were able to metabolize petroleum hydrocarbons. Williams and Kiceniuk (1987) reported

that a prolonged exposure to crude oil is required to suppress feeding by cunner, and that recovery can occur in a

few weeks. Deacutis (1982) found, however, that tautog

would not hesitate to consume hard clam or mussel meats

that were contaminated with #2 fuel oil, and that tautog

made little effort to avoid oil-contaminated feeding areas.

An environmental threat to tautog can exist in petroleumhydrocarbon-contaminated areas because bivalve mollusks,

a common prey, cannot metabolize polycyclic hydrocarbons

and can accumulate petroleum hydrocarbons to high levels

and lose it slowly, maintaining a reservoir for continued

induction (Vandermeulen and Penrose 1978).

Klein-MacPhee et al. (1993) reported larval tautog

survived, but had significantly reduced feeding and growth,

in an experimental exposure to starch blended with vinyl

alcohol copolymers (i.e., a biodegradable, potential substitute for plastic packaging). The low nutritional value of

starch to carnivorous fish and a low DO concentration were

suggested as being involved in these results.

ECOLOGICAL ROLES

The tautog is an important member of the resident,

coastal, three-dimensionally structured habitat community

in the Northeast’s marine environment. Although the species’ overall ecological role in the community has not been

well studied, we do have some information, such as their

predative and competitive interactions as discussed in the

“Feeding and Diet” chapter and in the “Competitors” section of the “Natural and Human-Induced Environmental

Factors” chapter, or we can suggest reasonable probabilities. These probable roles require more study, verification,
Page 17

and quantification, if possible. Some consideration should

be given also to the species that could irreversibly replace

tautog in the north-temperate-reef ecosystem, if tautog

abundance is reduced to the point where the species loses

its ability to compete for its former habitats and ecological

role.

One example of this role is the species’ predation on

mussels and other encrusting and epifaunal organisms found

in reef-like habitats, as discussed in the “Feeding and Diet”

chapter. This predation can be important to the biodiversity

of the invertebrate community of these habitats, and to the

mesoscale landscape ecology of these habitats. Tautog

grazing on dense sets of mussels and barnacles can help to

reopen habitat space for renewed colonization by other

species necessary for maintaining biodiversity and habitat

patchiness. That is, the tautog may be a “keystone” species in the same manner as Paines’ (1969) sea star was in

defining the concept. Its occasional feeding on anemones,

sponges, bryozoans, and other long-lived, relatively unproductive, surface-encrusting taxa can also open space

for colonization by more productive species, such as mussels or oysters, that can also be used as food by other

fisheries resources or be harvested by humans. The tautog’s

feeding can be important in controlling certain predators or

competitors, such as crabs, gastropods, or tunicates, of

juvenile harvestable shellfish. Heavy tautog predation on

shellfish beds or other encrusting fauna can also increase

the rate of recycling of nutrients, which accumulate in these

filter-feeding organisms, back into the water. Its preference

for mussels and shellfish can be a problem, however, if it

feeds on shellfish populations that are cultured or harvested

by humans.

Tautog will resort to shellfish prey from the surface of

open habitats, such as small clams in muddy or sandy areas

near and away from shelter, when epifaunal food resources

of the shelter become scarce. This alternate use of off-reef

prey can thus affect soft-bottom community ecology, as

well, in the vicinity of a reef habitat with a tautog population.

By crushing the shells of their molluscan prey during feeding, tautog also contribute to available habitat types

and use by altering a coarse, empty-whole-shell-rich habitat that would remain if mussels only died of adverse environmental conditions or age, to a finer, shell-hash-sediment

habitat. The coarse shell or the shell hash creates different

habitats used or favored by different invertebrate or postlarval fish and epifaunal species assemblages.

Some indirect effects of tautog populations on habitats, such as that of certain harvesting gear targeting tautog, are discussed in the “Structure” subsection, “Habitat

Modification and Loss” section of the “Natural and Human-Induced Environmental Factors” chapter.

Although small tautog are preyed upon (see the

“Predators” section of the “Natural and Human-Induced

Environmental Factors” chapter), it does not appear that

any predator is highly dependent on them as food, or that

tautog serve as a key forage species. Their localized populations, slow growth, and relatively long life can provide

stability to resident fish communities within which they

occur.

RESEARCH NEEDS

This review shows that there is much known about

the life history and habitat requirements of the tautog, although there are still many weak areas or gaps that need

attention before it can be claimed that we know enough.

Some of our knowledge of the species comes from laboratory studies; some effort must be made to confirm these

controlled-environment results in the field before they are

used, without question, in planning. In turn, some field

observations need to be examined in the laboratory to define better the cause-and-effect relationships. The variations or differences in some results from multiple studies

also need to be resolved, or the reasons for the differences

understood.

This review suggests there are areas for which life

history and habitat requirement information is nonexistent

or weak for tautog. Some of these key areas that require

special research attention are presented below, noting in

many cases the implications for fishery resource and habitat managers:

1. Defining specific prespawning and spawning aggregation areas used by all major local populations, as well as

defining the criteria for, or times of, use of these areas.

It will be critical to protect these areas from degradation, and to protect these populations against excessive exploitation during their use of these areas.

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 degradation, and to protect these fish against excessive exploitation 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 overall tautog population, noting their geographical distribution and trends, and relating them to recruitment,

growth, and exploitation rates. Such knowledge of different/local genetic components within the overall population could support more effective regional management of this species.

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 vulnerability.

8. Defining susceptibility of juveniles to coastal contamination and contaminant effects. Such knowledge would

be important for assessing and managing habitat/population 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 requirements.

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 Commission, Charleston, SC; 1995.

3. NOAA-UMass Cooperative Marine Education and Research 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 Committee, and others for sharing information with us.

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