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123

CHAPTER 10. Nekton

CHAPTER 10.

Nekton

Kenneth B. Raposa

124

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

125

CHAPTER 10. Nekton

Nekton

Introduction

Nekton generally refers to free-swimming

organisms including invertebrates, ﬁ sh, and marine

turtles and mammals. In estuaries, however, this

term typically refers to ﬁ shes and decapod crusta-

ceans. Nekton is a critical functional component of

estuarine ecosystems. Some estuarine nekton species

are commercially and recreationally important,

while others provide food for birds, mammals, and

larger ﬁ sh (Friedland et al., 1988; Sekiguchi, 1995;

Smith, 1997). Some species of nekton can physi-

cally transfer organic materials between intertidal

and subtidal estuarine habitats (Cicchetti, 1998),

and as a guild, nekton can be used as an indicator of

estuarine condition (Deegan et al., 1997). In some

situations, nekton can exert substantial top-down

control over estuarine system processes (Silliman

and Bertness, 2002). Nekton is also a charismatic

group of species that the public can easily relate to;

it therefore can provide an important link between

estuarine science and education or policy.

Narragansett Bay provides refuge, spawning,

and foraging habitats for a diverse assemblage of

nekton. Due to its location in southern New Eng-

land, Narragansett Bay supports species from north-

ern, boreal areas as well as species from subtropical

and tropical climates over an annual cycle. These

species include permanent and seasonal residents,

seasonal and occasional visitors, anadromous and

catadromous species, and accidentals and strays.

Narragansett Bay provides support functions for all

life history stages of nekton, including planktonic,

larval, juvenile, and adult stages. When present in

Narragansett Bay, these nekton have available to

them a wide variety of habitats that include open

water, unvegetated bottoms, intertidal beaches, salt

and brackish marshes, SAVs, tidal freshwater creeks,

rocky reefs, and human-modiﬁ ed shorelines.

Many species of nekton in Narragansett Bay

support commercial or recreational ﬁ sheries (DeAl-

teris et al., 2000) and thus have been the focus of

numerous research and monitoring programs. Based

on data from several ongoing nekton monitoring

programs, a great deal is known about the long-term

trends in species abundance and biomass as well as

distribution patterns over time. Aside from this, sur-

prisingly little research has actually been done that

speciﬁ cally examines the ecology and functional

role of most ﬁ sh species in Narragansett Bay. For

example, Keller et al. (1996) indicates that we still

do not fully understand why the abundance of some

species varies considerably over time independent of

ﬁ shing pressure.

The goal of this chapter is to provide an

ecological overview of nekton from two major

zones of the Bay (open water and shore) and another

overview focusing on ichthyoplankton. Open water

nekton include those species that typically are found

in the deepwater areas of the Bay, either in pelagic

or demersal habitats, and those that are typically

captured with a trawl. Shore-zone or intertidal nek-

ton include those species that are found in shallow

water habitats of the Bay that include salt marshes,

eelgrass beds, coves, embayments and unvegetated

shallows.

Open-water Nekton

One of the ﬁ rst studies that focused on

ﬁ shes in the open waters of Narragansett Bay was

conducted over 30 years ago by Oviatt and Nixon

(1973). These authors used a trawl to sample from

nine regular and 13 occasional stations in Narra-

gansett Bay for one year. Forty-four species were

documented in Narragansett Bay. Although typical

of temperate estuaries, a small number of species

dominated the catch (in this study, the 10 most abun-

dant species made up 91 percent of the catch). This

study also demonstrated that:

• The composition of the ﬁ sh community in

Narragansett Bay is comparable to those in

Block Island and Long Island sounds.

• Fish abundance and biomass per unit area are

comparable to other New England coastal

and offshore areas, although standing crop was

much less than in kelp forests, coral reefs, and

salt marshes.

• Winter ﬂ ounder (Pseudopleuronectes

americanus) was easily the most abundant

species, making up 36 percent of the catch.

• Spatial patterns in ﬁ sh distribution were not

apparent except that diversity was highest near

the mouth of the Bay.

• The demersal ﬁ sh in Narragansett Bay may

be important in regulating the diversity and

abundance of the benthos.

Oviatt and Nixon’s work was limited in that

it only documented the ﬁ sh of Narragansett Bay

at one point in time. For example, although win-

ter ﬂ ounder dominated in 1971–72, this and other

126

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Figure 10.1. Locations of sampling stations that are part of the seasonal and monthly ﬁ sh trawl survey, the juvenile ﬁ nﬁ sh survey, and

the Keller et al. ichthyoplankton survey that are discussed in this chapter.

127

CHAPTER 10. Nekton

demersal species have declined dramatically in sub-

sequent decades (Oviatt et al., 2003). In recognition

of the need for detailed ﬁ sheries data over time, two

long-term monitoring programs were initiated in the

open waters of Narragansett Bay. These programs

are the RIDEM sportﬁ sh trawl survey throughout

Narragansett Bay and in Rhode Island and Block

Island sounds (e.g., Lynch, 2000), and the GSO ﬁ sh

trawl survey (Jefferies and Johnson, 1974; Jeffries

and Terceiro, 1985; Jeffries et al., 1989). The GSO

trawl survey is the longer running of the two, dat-

ing back to 1959; however, this survey is spatially

limited since samples are only collected from two

stations in the West Passage of Narragansett Bay. In

contrast, the RIDEM trawl survey began 20 years

later in 1979, but it samples throughout the entire

Bay (Fig. 10.1) and thus provides a

more comprehensive dataset in terms

of combining temporal and spatial

coverage. The RIDEM program has

two components: a monthly survey

at 12 ﬁ xed stations in the Bay that

began in 1990, and a seasonal survey

in spring and fall at approximately

50 stations (selected randomly from

approximately 265 stations located

throughout the Bay) that began in

1979.

From 1979 through 2003,

107 species (mostly ﬁ sh, a few crustaceans, and one

bivalve species) have been collected from the com-

bined efforts of the RIDEM monthly and seasonal

ﬁ sh trawls. However, the mean number of species

in any given year is much less, averaging 57 species

per year from the monthly program and 45 species

per year from the seasonal program (Fig 10.2). This

illustrates the value of the two programs—more

species are observed annually with the monthly ef-

fort, which provides a more comprehensive overall

view of ﬁ sh community composition and structure,

while the seasonal program provides more infor-

mation on the Bay-wide distribution of common

species because more stations are sampled. Based

on abundance from the seasonal data, ﬁ ve species

make up greater than 90 percent of the community

found in Narragansett Bay since 1979. In decreasing

abundance, these species include bay anchovy (An-

choa mitchilli, 51 percent of total abundance), scup

(Stenotomus chrysops, 19 percent), longﬁ n squid

(Loligo pealei, 8 percent), menhaden (Brevoortia

tyrannus, 6 percent), and butterﬁ sh (Peprilus tria-

canthus, 5 percent) (Appendix 10.1). Using the same

data, but considering biomass, 13 species make up

over 90 percent of the total nekton biomass. In de-

creasing order, these species are scup (19 percent),

winter ﬂ ounder (18 percent), American lobster (9

percent), skates (Rajidae, 9 percent), windowpane

ﬂ ounder (Scophthalmus aquosus, 6 percent), longﬁ n

squid (6 percent), tautog (Tautoga onitis, 6 percent),

butterﬁ sh (5 percent), summer ﬂ ounder (Paralich-

thys dentatus, 4 percent), bay anchovy (3 percent),

weakﬁ sh (Cynoscion regalis, 2 percent), Atlantic

herring (Clupea harengus, 2 percent), and blueﬁ sh

(Pomotomus saltatrix, 2 percent). Based on biomass,

the nekton species that dominate Narragansett Bay

are primarily demersal species such as ﬂ ounders,

lobster, and skates. However, based on abundance,

the opposite is true where the dominant species are

mostly small, schooling, pelagic species.

The data from the RIDEM trawl programs

are particularly useful for observing trends in ﬁ sh

over time and at a Bay-wide scale.

There is no clear trend in the annual

number of species in Narragansett Bay

(Fig. 10.2), nor is there a trend in total

ﬁ sh biomass over time (Fig. 10.3). In

contrast, total abundance is tending

to increase over time, mostly due to

increases in small pelagic schooling

ﬁ sh such as Atlantic menhaden and bay

anchovy. In fact, these data have docu-

mented a shift in species abundance

patterns in Narragansett Bay. The Bay

is undergoing a shift from a community

dominated by demersal species to a system domi-

nated by pelagic species that may be due to climate

and bottom-trawl ﬁ shing (Oviatt et al., 2003). Fur-

ther, data from the seasonal trawl survey illustrate

that this trend is occurring on a Bay-wide scale. For

example, using GIS, it is clear that the abundance

of the commercially important winter ﬂ ounder has

been in steady decline since at least the beginning

of the survey, and this decline is evident throughout

Narragansett Bay (Fig. 10.4). Similar patterns have

been observed for other demersal species, including

those that are not exposed to ﬁ shing pressure (e.g.,

hogchoker, Trinectes maculatus) (Lynch, personal

communication).

In contrast to the abundance of long-term

monitoring data, surprisingly little research on open-

water nekton in Narragansett Bay has been con-

ducted, especially recently. However, there are some

notable recent examples. Durbin and Durbin (1998)

used a bioenergetic model to examine the effects of

menhaden predation on phytoplankton in Narragan-

sett Bay. DeAlteris et al. (2000) used monitoring

and landing data to summarize the status and trends

of many of Narragansett Bay’s commercial ﬁ sher-

ies. Lapolla (2001a, 2001b) examined a number

of population characteristics of the bay anchovy in

128

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Figure 10.2. The average annual number of species of nekton in Narragansett Bay as determined from the RIDEM

seasonal and monthly ﬁ sh trawl program. Nearly all the species are ﬁ shes; relatively few are invertebrates.

grams per trawl; BPUE) between 1979 and 2003 from the RIDEM seasonal ﬁ sh trawl.

129

CHAPTER 10. Nekton

Figure 10.4. Winter ﬂ ounder abundance and distribution

in Narragansett Bay in three time blocks (a = 1979–1985;

b = 1986–1995; c = 1996–2003). For each ﬁ gure, mean

CPUE is shown, where one dot equals nine ﬁ sh. Stations

that are sampled by the trawl program during each time

block are outlined in black.

130

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Narragansett Bay, including population structure,

growth, mortality, and spawning season. Meng et

al. (2001) found that winter ﬂ ounder growth rates

in Narragansett Bay were lower in the upper Bay,

suggesting that anthropogenically lowered dissolved

oxygen levels had a negative impact on this species.

More recently, Oviatt et al. (2003) used historic and

current data to demonstrate the dramatic effects

humans have had on the ﬁ shes of Narragansett Bay

through ﬁ shing pressures, and Castro and Angell

(2000), Castro et al. (2005), and Cobb and Castro

(2006) have examined aspects of the emergence,

spread, and severity of lobster shell disease in the

Bay.

Shore-zone and Intertidal Nekton

Shallow estuarine waters provide critical

nursery habitats for juvenile estuarine ﬁ sh and per-

manent habitats for some abundant forage species.

These types of habitats are often at risk, however,

due to their proximity to the land and thus the

activities of humans. Nekton in shallow, shore-zone

habitats are monitored monthly from June through

October by RIDEM with a juvenile ﬁ nﬁ sh sein-

ing survey at 20 nearshore stations in Narragansett

Bay (Fig. 10.5). Since the inception of this program

in 1990, 78 species (or undifferentiated species

within the same family, e.g., gobidae, bothidae)

have been collected from nearshore and shore-zone

habitats as part of this monitoring program. Based

on abundance, the most common species include

Atlantic menhaden (62 percent of total abundance),

silversides (Menidia spp., 8 percent), river herring

species (6 percent), blueﬁ sh (Pomotomus saltatrix, 4

percent), winter ﬂ ounder (3 percent), striped killiﬁ sh

(Fundulus majalis, 3 percent), sea herring species

(3 percent), and bay anchovy (2 percent) (Appendix

10.2). Meng and Powell (1999) used these data to

explore relationships between ﬁ sh communities and

habitats. This study found that separate analyses

of ﬁ sh communities and their habitats correlated

well. In addition, it was found that total abundance,

species richness, and the number of winter ﬂ oun-

der were highest at an upper Bay station. This is

contrary to the ﬁ ndings of Oviatt and Nixon (1973);

however, the two studies used different gears to

sample different age classes of ﬁ sh, and the two

studies were conducted over 25 years apart. Dorf

and Powell (1997) used these same seining data to

document the distribution and habitat preferences of

juvenile tautog, a recreationally important species,

in Narragansett Bay. More recently, DeLong et al.

(2001) used data from this survey in a model to

examine the effects of density and environmental

conditions on the growth of juvenile winter ﬂ ounder.

Nekton has also been sampled extensively

from salt marsh habitats around Narragansett Bay

and the south shore of Rhode Island (Fig. 10.6).

As with salt marshes elsewhere, marshes in Rhode

Island clearly support highly abundant and produc-

Figure 10.5. Researchers

conducting the RIDEM

juvenile ﬁ nﬁ sh seine

survey. Photo by J.

Christopher Powell,

RIDEM.

131

CHAPTER 10. Nekton

tive nekton communities (Raposa, 2002; Meng et

al., 2004). Quantitative data collected from three

salt marshes around Rhode Island show that these

marshes are consistently dominated by very few

species (i.e., species diversity is low). These spe-

cies include the common mummichog (Fundulus

heteroclitus), striped killiﬁ sh, sheepshead min-

now (Cyprinodon variegatus), Atlantic and inland

silversides (Menidia menidia and Menidia beryllina,

respectively), and grass shrimp (Palaemonetes spp.)

(Appendix 10.3). Less abundant, though ecologi-

cally important, species that also use Narragansett

Bay salt marshes include juvenile winter ﬂ ounder,

sticklebacks (e.g., three-spined Gasterosteus aculea-

tus, fourspine Apeltes quadracus, and nine-spined

Pungitius pungitius), American eel (Anguilla rostra-

ta), and blue crab (Callinectes sapidus). The data in

Appendix 10.3 further indicate that while general

patterns of species composition are similar among

marshes, large differences in density exist (e.g.,

Palaemonetes pugio). Halpin (1997) also noted

substantial differences in mummichog use among

different Narragansett Bay salt marshes. The factors

that contribute to differences in nekton composition

and abundance among salt marshes in Narragansett

Bay are largely unknown and need to be identiﬁ ed

and examined, especially in light of ongoing and

future marsh restoration efforts.

Marsh nekton species can move among and

utilize multiple marsh habitats (e.g., creeks, pools,

vegetated marsh surface) depending on life history

stage and tide stage. Roman et al. (2003) showed

that more species were found in subtidal creeks and

pools when compared to intertidal marsh habitats in

the Sachuest Point salt marsh in Middletown, R.I.

Data from Raposa (2002) in the Galilee, R.I., salt

marsh indicate that nekton tend to be more abundant

in subtidal, rather than intertidal, marsh creeks. In

nearby Cape Cod, Mass., Raposa (2003) showed

that mummichogs moved into soft-substrate pools in

fall where they burrowed into the sediments to over-

winter. A given marsh is a dynamic place with mul-

tiple habitats interacting to support nekton. Threats

to some of these habitats in Rhode Island marshes

include the invasion of high marsh by the common

reed, Phragmites australis, the loss of marsh pools

due to historic ditching, and tidal restrictions that

limit nekton access to marsh surface habitats, which

are used for foraging, nursery, and refuge.

The restoration of tide-restricted salt marshes

around Narragansett Bay is clearly returning natural

and abundant nekton communities to marshes that

supported a dysfunctional and depleted community.

Studies indicate that removing tide-restricting struc-

tures results in improved nekton function, and that

the more severe the restriction, the more negatively

affected the nekton community is, and the more

positive the response is after restoration (Raposa,

2002; Raposa, unpublished data; Raposa and Ro-

man, 2003; Roman et al., 2003). A consortium of

agencies, including the R.I. Coastal Resources

Management Council, the Narragansett Bay Estu-

ary Program, and Save The Bay, among others, has

identiﬁ ed salt marshes around Narragansett Bay that

are in need of restoration, and some of these efforts

are under way. If previous results hold true, these

restoration efforts should continue to return nekton

communities to more natural conditions represen-

tative of unrestricted salt marshes. In addition to

removing tidal restrictions, efforts should seek to

restore pool habitats that were lost from ditching.

Salt marsh pools can support dense nekton assem-

blages (Raposa and Roman, 2001), and if the pools

are shallow enough, this nekton provides attractive

forage for wading birds.

Ichthyoplankton

Ichthyoplankton (eggs and larvae) are early

life-history stages of nekton that are useful for

understanding adult spawning patterns and temporal

ﬂ uctuations in the abundance of juvenile and adult

nekton. Ichthyoplankton are particularly abundant

in estuaries in part due to the use of these areas as

spawning and nursery grounds by nekton species.

In recognition of this, and to help ﬁ ll a critical

data gap, multiple surveys and ichthyoplankton

monitoring programs were initiated in Narragansett

Bay. The ﬁ rst survey occurred in 1957–1958 and

included sampling in the lower East Passage of

Narragansett Bay and in Mount Hope Bay (Herman,

1963). Another survey occurred in 1972–1973 and

included 160 total stations divided among 10 sectors

in Narragansett Bay (Bourne and Govoni, 1988;

hereafter referred to as the MRI (Marine Research

Inc.) survey). Almost 20 years later, similar methods

were used by Keller et al. (1999; hereafter referred

to as the Keller survey) to collect newer data from

1989–1990 and to explore changes in ichthyoplank-

ton composition and abundance over time. The

most recent effort is a partnership between URI and

RIDEM to collect annual data beginning in 2002 to

observe ichthyoplankton trends over an even longer

time period (Klein-MacPhee et al., 2002). The

combined data from these programs provide a base-

line for examining trends in composition, relative

abundance, distribution, and seasonal abundance of

ichthyoplankton in Narragansett Bay.

132

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Ichthyoplankton on the whole display a

clear seasonal pattern in abundance, with a distinct

peak in eggs in June and in larvae slightly later in

July. This pattern was observed in both the MRI and

Keller surveys. The total number of ichthyoplankton

species was also similar between the two surveys

(43 in the MRI survey; 41 in the Keller survey), but

differences in the abundance of dominant species

were apparent. In 1972–73 the most abundant spe-

cies included cunner (Tautogolabrus adspersus),

tautog, bay anchovy, Atlantic menhaden, scup, and

weakﬁ sh; in 1989–90 the dominant species included

bay anchovy, tautog, and cunner, but menhaden,

scup, and weakﬁ sh were not abundant. Egg and lar-

val (all species combined) densities were consider-

ably lower in 1989–90 compared to the MRI survey.

Abundance of some species declined substantially

in the highly impacted upper Bay, Providence

River, and Greenwich Bay areas. In fact, Keller et

al. (1999) indicate that there was a general shift in

ichthyoplankton distribution down-Bay away from

these impacted areas. It was not clear whether this

was due to reduced adult spawning in the upper Bay

regions, or to higher mortality of ichthyoplankton

while in these areas. In either case, upper Bay re-

gions that were known as important historic spawn-

ing and nursery areas for some important nekton

species now seem to have lost some of that value,

perhaps due to impacts from human activities.

Summary

In addition to the impacts to ichthyoplankton

outlined above, the abundance, distribution, growth,

and survival of juvenile and adult nekton in Nar-

ragansett Bay are also affected by human activities.

Commercial ﬁ shing has depleted many ﬁ sh popula-

tions over at least a century (Oviatt et al., 2003),

and ﬁ shing pressures continue to exert considerable

inﬂ uence. Substantial areas of important nursery

habitats such as eelgrass and salt marshes have been

extensively degraded or lost. Eutrophication and

the resultant increase in the frequency and dura-

tion of hypoxia forces ﬁ sh to either move out of the

affected areas or suffer negative impacts. Meng et

al. (2001) demonstrated that winter ﬂ ounder growth

and survival decreased in upper Bay areas where

water quality and dissolved oxygen conditions are

poor. In the summer of 2003, a large ﬁ sh kill (over 1

million Atlantic menhaden) occurred in Greenwich

Bay when excessive nutrients and physical pro-

cesses combined to create an extensive anoxic event

(RIDEM, 2003). However, despite all of these pres-

sures, Narragansett Bay and its habitats continue to

support an abundant and diverse nekton assemblage,

albeit one whose composition appears to be shifting

over time.

Figure 10.6. Using a throw trap to

quantitatively sample nekton from salt

marsh habitats. Photo from NBNERR photo

library.

133

CHAPTER 10. Nekton

Appendix 10.1. Abundance and Biomass of Nekton Species

Abundance and biomass of nekton species collected during the RIDEM seasonal trawl survey. For

each species, mean abundance (catch per unit effort, CPUE) and mean biomass (biomass per unit

effort, BPUE) are provided as averages between 1979 and 2003. Averages for spring, fall, and all data

combined are provided.

134

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Appendix 10.1 Continued

135

CHAPTER 10. Nekton

Appendix 10.2. Species Composition and Abundance of Fishes

Species composition and abundance of ﬁ shes collected between 1990 and 2003 during the RIDEM juvenile

ﬁ nﬁ sh seining survey. For each species, the average number per seine (across all 20 stations and all years) is

shown for each month of the survey and for the entire survey (across all months).

Appendix 10.1. Continued

136

An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

Appendix 10.2. Continued

137

CHAPTER 10. Nekton

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2

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An Ecological Profile of the Narragansett Bay National Estuarine Research Reserve

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