NOAA Technical Memorandum NMFS NE 199
Distribution
and Abundance of Fish Eggs
Collected
during the GLOBEC Broadscale
Georges Bank Surveys, 1995-1999
by John D. Sibunka, Donna
L. Johnson, and Peter
L. Berrien
National Marine Fisheries Serv., 74 Magruder Rd., Sandy Hook, NJ 07732
Print
publication date August 2006;
web version posted January 24, 2007
Citation: Sibunka JD, Johnson DL, Berrien PL. 2006. Distribution and abundance of fish eggs collected during the GLOBEC broadscale Georges Bank surveys, 1995-1999. NOAA Tech Memo NMFS NE 199; 72 p.
Download complete PDF/print version
Abstract:
This regional atlas summarizes and illustrates data on fish eggs
collected from the U.S. GLOBEC Georges Bank broadscale
surveys from 1995 through 1999. Distributions and abundances are
presented for 26 ichthyoplankton taxa from egg
density maps by survey, with accompanying discussion of seasonal
patterns and spawning variation.
INTRODUCTION
The U.S. GLOBEC (United States GLOBal ocean ECosystems dynamics) Georges
Bank program was designed to address the influence of climate
change on productivity, abundance, and distribution of marine organisms
on Georges Bank (also referred to within this paper as “the Bank”). Appendix
Figure 1 shows the geographic study area. The broad-scale surveys were
a principal component of the GLOBEC program, in which bank-wide survey
coverage was conducted to monitor changes in both the physical and biological
processes that influence the Georges Bank ecosystem. This atlas depicts
the distribution and abundance trends of fish eggs collected with a 61-cm
bongo plankton sampler from 30 broad-scale surveys during six months
each year over five years of study. The abundance and distribution of
fish eggs provide valuable information on spawning times, areas, and
intensities. The correlation of this information to environmental parameters
will help in understanding the factors which influence fluctuations in
year-class strength and survival in the early life stages of fish. Pictorial
maps displaying abundance and distribution of fish eggs are presented
for 26 taxa. This report is the first comprehensive fish egg study to
be documented on the Bank since the Marine Resource Monitoring Assessment
and Prediction (MARMAP) 1977 to 1987 program.
METHODS
Broad-scale surveys were conducted monthly, surveying the Georges Bank
region during the winter through late spring-early summer from 1995 through
1999. Individual survey information along with the cruise (survey) report
can be obtained from the U.S. GLOBEC Georges Bank website (http://globec.whoi.edu).
The 1995 monthly surveys began in February and ended in July with 38
stations selected to cover Georges Bank, its slope waters, and adjacent
Gulf of Maine region. Beginning in 1996, the sampling period began in
January and ended in June, and stations were added, resulting in 39 stations
for 1996, 40 stations for 1997, and 41 stations for 1998 and 1999. In
1996 (all months but June), the ichthyoplankton sampling density was
supplemented with extra bongo tows at new locations within the overall
survey scheme. The distribution pattern for supplementary bongo tows
was changed again in 1997 to include a bongo tow made halfway between
stations on all monthly surveys from January to May. This intermediate
bongo tow scheme was retained in 1998 and 1999.
Ichthyoplankton samples were collected with a 0.61-m bongo frame fitted
with 333-mm mesh nets (Posgay and Marak 1980). Digital flow meters were
suspended in the aperture of each net to determine the volume of water
filtered. A 45-kg ball was attached beneath the bongo frame to depress
the sampler. A real-time CTD (conductivity, temperature, depth) profiler
was attached to the towing wire above the frame to measure temperature
and salinity, and to monitor sampling depth in real-time
mode. Egg catches were standardized by multiplying the
number of eggs collected by a derived haul factor for each station tow
to obtain densities per 10 m2 of sea surface area. The haul
factor was derived by:
| Haul factor = |
maximum tow depth
(flowmeter calibration)x(flowmeter revolutions)x(area of net
mouth) |
Standard plankton sampling procedures
derived from the MARMAP program were used (Jossi and Marak 1983). Tows
were smooth oblique from the surface to within a few meters of the
bottom or to a maximum tow depth of 200 m while maintaining an approximate
45° wire angle. Vessel
speed varied from 1½ to 2 knots. Wire payout was 50 m/min and
retrieval was 20 m/min. These rates were adjusted at station depths <60
m to ensure a minimum of 100 m3 of water filtered (Houde 1977).
Payout rates were also adjusted during adverse weather conditions to
minimize surging of the tow wire. The sample from one net retained for
ichthyoplankton analysis was preserved in 5% formalin. The contents from
the other net were preserved in 95% ethanol. All fish eggs were removed
and staged according to their development. The following egg-stage classifications
were used:
- “Early” represented the period from spawned to just before
blastopore closure.
- “Middle” represented the period from blastopore closure
to tail bud almost free (i.e., slightly undercut).
- “Late” represented the period from tail bud free to
just before hatching.
Late stage Gadus morhua, Melanogrammus aeglefinus,
and Pollachius virens eggs were further staged:
a. from tail bud free to tail tip twisted and
flexed >45°,
b. from tail tip twisted and flexed >45° to embryo encircled >3/4
of egg circumference to < embryo full circle of egg,
c. from > full circle of egg to just before hatching.
“Dead before capture” and “abnormal” eggs were
also classified. Dead eggs were distinguished by having their yolk material
and embryo (if present) degraded and concentrated to a side of the egg
chorion. Abnormal eggs were those with an obvious embryonic aberration
such as stunted development or two tail buds. Dead and abnormal eggs
are not included in this atlas.
The identification of some early-stage eggs could not be established
and were categorized as “unidentified.” Other eggs included
as “unidentified” were of various developmental stages, and
their taxonomic identity has yet to be established. These eggs were assigned a type designation. Unidentified
eggs are not presented in this atlas but are listed along with eggs staged
as early, middle, late, dead, and abnormal in the fish egg data base
located on the U.S. GLOBEC Georges Bank website.
Several species share indistinguishable common early-stage characteristics
and cannot be separated. Such features include eggs that overlap in size
(diameter), the presence or absence of an oil globule, eggs that overlap
in size of egg and oil globule, and eggs with similar pigmentation patterns
on the embryo, yolk, or both (refer to Berrien and Sibunka 1999). The
groupings of species whose eggs co-occur but could not be distinguished
consisted of the following: (1) Urophycis spp. and Peprilus
triacanthus, (2) Urophycis spp. and Enchelyopus cimbrius,
(3) Limanda ferruginea and Tautogolabrus adspersus,
(4) Scophthalmus aquosus and Paralichthys oblongus,
(5) Gadus morhua and Melanogrammus aeglefinus,
and (6) Gadus morhua, Melanogrammus aeglefinus,
and Glyptocephalus cynoglossus. The eggs in each
grouping were assigned and tabulated to taxa in a proportion based on
the proportion of identifiable later stage eggs at the same or from a
nearby station.
RESULTS
A total of 26 taxa were identified and are presented in Appendix
Table 1. Also included in Appendix
Table 1 for each taxon are the total number
of tows, the number of positive tows per month(s) when the taxon was
collected, and the corresponding monthly density (number per 10 m2). Maps
showing the standard station locations for each of the five survey years
are given in Appendix
Figure 2. Individual maps are presented for all
positive occurrence surveys for the 26 taxa. These maps show the distribution
of egg densities for the combined early, middle, and late stage eggs
(Appendix
Figures 3-28).
Eggs from fishes in the family Pleuronectidae, which included Limanda
ferruginea, Glyptocephalus cynoglossus, Hippoglossoides
platessoides, and Pseudopleuronectes americanus,
were the most abundant and comprised 68% of identified eggs collected
in five years of survey effort. Limanda ferruginea eggs were
the most abundant taxon collected (66% of the identified eggs). The
order Gadiformes, which included Gadus morhua, Melanogrammus
aeglefinus, Pollachius virens, Brosme brosme, Enchelyopus cimbrius,
and the hakes (Urophycis spp., Merluccius
bilinearis and Merluccius albidus), were the second
most abundant (28% of identified eggs). Eggs of Gadus morhua and Melanogrammus
aeglefinus comprised 10% and 9% of the total egg catch, respectively.
TAXON ACCOUNTS
(Click on headings to see associated figures)
In the following species accounts, taxa are ordered phylogenetically
by convention established in Nelson et al. (2004). For each taxon, information
is provided on: (1) the general habitat and geographic range within the
Northwest Atlantic, (2) results from the 11-year (1977 through 1987)
MARMAP ichthyoplankton surveys (for survey method details see Sibunka
and Silverman 1984, 1989), and (3) new results from the GLOBEC ichthyoplankton
surveys. Identification notes are also provided when warranted.
Margined snake eel, Ophichthus cruentifer (Figure 3)
Adult Ophichthus cruentifer range from the Gulf of Maine to Florida,
but are uncommon north of Cape Cod, MA (Smith and Tighe 2002). Eggs of O.
cruentifer were collected during the MARMAP surveys off southern
Georges Bank from June to October (Berrien and Sibunka 1999).
Only small numbers of O. cruentifer eggs were collected, and
these came from three stations located in the southwest portion of the
survey area during the June 1999 GLOBEC survey. One early stage egg was
taken on the southwest area of Georges Bank, and a second early egg was
collected on an adjacent station in the southern portion of the Great
South Channel. A single middle stage egg was collected on the southwest
shelf slope of the Bank.
Atlantic argentine, Argentina silus (Figure 4)
Adult Argentina silus are found in deep waters off the continental
shelf in southern Labrador to off the southern slope portion of Georges
Bank (Klein-MacPhee 2002a). Catch occurrences of A. silus eggs
during the MARMAP surveys indicate spawning in the eastern portion of
the Gulf of Maine during April and May (Berrien and Sibunka 1999).
Small catches of A. silus eggs were collected during the April
GLOBEC surveys in 1998 and 1999. The catch occurrences for both years
were from deep waters in the Gulf of Maine off the northeast portion
of Georges Bank.
Atlantic pearlside, Maurolicus weitzmani (Figure 5)
Adult Maurolicus weitzmani range from southern Newfoundland to
the Gulf of Mexico and the Caribbean Sea (Harold et al. 2002). The eggs
of M. weitzmani were collected during the MARMAP surveys
on slope regions of Georges Bank from late summer to early winter (Berrien
and Sibunka 1999; taxon referred to as M. muelleri).
A single M. weitzmani egg was collected in deep water during
the March 1995 GLOBEC survey on the southwest shelf-slope region of the
Bank.
Fawn cusk-eel, Lepophidium profundorum [tentative identification] (Figure 6)
Adult Lepophidium profundorum inhabit the outer portion of the
continental shelf and are common along the shelf-slope of Georges Bank
(cf. map 12 in Collette and Klein-MacPhee 2002). The larvae of L.
profundorum are prevalent during the summer on the southern
part of the Bank (Fahay 1992).
L. profundorum eggs were collected in small numbers on the July
1995 and the June 1999 GLOBEC surveys. These egg catches were made in
or near shelf-slope waters along the southern flank of Georges Bank.
Identification Notes
The eggs tentatively identified as L. profundorum have characteristics
that resemble those described for Ophidion marginatum by Fahay
(1992). These characteristics include a smooth and slightly-to-noticeably
oval chorion (diameter 0.91-0.95 x 0.85-0.92 mm), the absence of an oil
globule, a homogenous yolk, and a narrow perivitelline space. In addition,
the high myomere counts (50+) made on late-stage embryos from eggs held
at the Northeast Fisheries Science Center (NEFSC), along with the locations
where these eggs were collected (i.e., Southern New England and Georges
Bank), also indicate these eggs are most likely L.
profundorum.
Fourbeard rockling, Enchelyopus cimbrius (Figure 7)
Adult Enchelyopus cimbrius range along the eastern coast of
North America from Newfoundland to the northern portion of the Gulf of
Mexico. Their distribution on Georges Bank is along the northern edge,
with the largest concentration toward the western portion and into the
Gulf of Maine (cf. Map 15 in Klein-MacPhee 2002b). E. cimbrius eggs were collected
on the MARMAP surveys along the northern edge of Georges Bank from March
to January, with the highest densities occurring in May (Berrien and
Sibunka 1999).
E. cimbrius eggs were collected on GLOBEC surveys from February
to July. They were distributed on the northern portion of Georges Bank,
with small and isolated egg catches made along the southern flank. During
the winter and early spring surveys, E. cimbrius eggs were caught
in small numbers along the northern and southern periphery of Georges
Bank. The exception to this trend was the March 1998 survey results,
which indicated an increase in spawning activity along the north central
and northwest edge area. Both April-May in 1996 to 1999 and May-June
in 1995 were peak spawning months during the five-year GLOBEC sampling program. E.
cimbrius egg catch results for the April and May surveys in 1998
and 1999 show an almost continuous distribution pattern along the entire
northern edge region of the Bank. The smallest egg catches during April
and May were made in 1995. Spawning effort remained high in June 1995
but had begun to decrease during the other four June surveys, with most
eggs collected again on the Bank’s northern edge. Although the
catch results for the entire 1995 survey season were low, the July catches
were smaller than those collected in June. The small egg density values
in July further substantiate that the peak spawning period for E.
cimbrius on Georges Bank had passed. Egg collections
were limited in July to only a few stations located on the north central Bank area.
Hake species, Urophycis spp. (Figure 8)
Adult Urophycis spp. inhabit the North American continental shelf
and slope waters from Labrador to North Carolina and stray as far south
as Florida (Klein-MacPhee 2002b). Urophycis spp. eggs were collected on Georges
Bank during the MARMAP surveys in all months except February. The highest
egg densities occurred during the summer months on the Bank (Berrien
and Sibunka 1999). Urophycis spp. eggs were collected in high
abundance during the GLOBEC surveys in June and July (except June 1996).
Small and isolated egg catches occurred during all other months surveyed.
Their distribution during years of high abundance extended across Georges
Bank. Urophycis spp. eggs were collected during the 1995
winter and early spring (February to April) surveys in small and isolated
catches along the Bank’s shelf-slope area. Most of the May and
June catch results for all survey years show either the onset of spawning
or an increase in spawning intensity on the Bank. Eggs collected on the
May surveys were concentrated along the Bank’s southern area. Catch
results for all June surveys, except for 1996, indicate an increased
level of spawning across the entire Bank, with substantial egg catches
along the entire southern and northern region. The large egg catches made in the northern
Bank region were concentrated on the upper portion of
Cultivator and Georges Shoals. Urophycis spp. eggs were also
collected both in the Great South Channel and on
Browns Bank. The June 1996 survey results indicated limited spawning
taking place on the Bank with two large egg catches made just west of
Cultivator and Georges Shoals. Urophycis spp. egg distribution
and abundance for July 1995 was greater than for June of that year and
resembled the results of the June 1997 to 1999 surveys.
Identification Notes
Hake eggs were not identified to species but were categorized as Urophycis spp.
Two adult species of hake, U. chuss (red hake) and U.
tenuis (white hake), are reported to inhabit Georges Bank and the
adjacent waters. Both hakes begin spawning in the spring and continue
through the summer months (Able and Fahay 1998; Klein-MacPhee 2002b). U.
tenuis are known to spawn in the continental slope waters off
Georges Bank but may not spawn on the Bank or in the Gulf of Maine. Another
adult hake, U. regia (spotted hake), inhabits the waters from
Southern New England through the Middle Atlantic Bight (Klein-MacPhee2002b).
Information from the NEFSC 40-year bottom trawl survey shows that adult U.
regia occur on the southwest portion of Georges Bank and along the
Bank’s southern slope-edge (NEFSC unpubl. data1).
Most U. regia spawning takes place in the central
and southern Middle Atlantic Bight (Able and Fahay 1998). It is possible
that most if not all of the eggs collected on the Bank are U.
chuss and that those eggs collected on the Bank’s
shelf-slope region are U. chuss, U. tenuis, or both.
The larval hakes collected on the broad-scale surveys were identified
only to genus, so larval occurrences did not help clarify the species
composition of the co-occurring hake eggs.
Offshore hake, Merluccius albidus (Figure 9)
Adult Merluccius albidus are found off the U.S. Atlantic coast
in deep basins within the Gulf of Maine and along the southeastern slope
of Georges Bank south to slope waters off Florida (Klein-MacPhee 2002c). M.
albidus eggs were taken during the MARMAP surveys
in all months of the year on Georges Bank. They ranged from the Northeast
Channel to along the shelf slope of Georges Bank (Berrien and Sibunka
1999).
M. albidus eggs were collected during all months of the GLOBEC
broad-scale program but not on all surveys during a given year. The catch
occurrences of M. albidus eggs were small and these
eggs were collected in small numbers along the shelf-slope of Georges
Bank. An increase in egg catch was observed during the late spring, indicating
intensified spawning activity during this time. The only catch occurrence
of this taxon on Browns Bank took place during the May 1997 survey.
Silver hake, Merluccius bilinearis (Figure 10)
Adult Merluccius bilinearis are common along the North American
coast from southern Newfoundland to South Carolina (Klein-MacPhee 2002c). M.
bilinearis eggs were collected during the MARMAP surveys
on Georges Bank all months of the year except February. M. bilinearis egg densities
were the highest during the summer months and were distributed across
the entire Bank (Berrien and Sibunka 1999).
M. bilinearis eggs were collected on the GLOBEC surveys
from January to July, and were distributed over the entire Bank. The
eggs were most abundant in late spring and into the summer on the Bank.
Spawning generally began along the southern portion of the Bank in April
or May. An earlier isolated egg catch occurred on the Bank’s
southeastern shelf-slope in January 1996. In June, except for 1996, egg
densities indicate spawning activity intensified dramatically and spread
across Georges Bank. Catches of M. bilinearis eggs were also made on Browns Bank in late spring
and early summer. In 1996, M. bilinearis spawning was
either late in starting, low in intensity, or both compared to the results
of the other four years. Catch results for the May surveys in 1998 and
1999 show a few eggs were collected at the same station located in the
northwest portion of the Bank. These catches may indicate a small and separate
spawning event by M. bilinearis.
Cusk, Brosme brosme (Figure 11)
Adult Brosme brosme range along the North American coast from
the Newfoundland Banks and the Strait of Belle Isle to New Jersey (Bigelow
and Schroeder 1953). B. brosme were collected on Georges Bank
during the MARMAP surveys from April to October. The largest egg densities
on the Bank occurred from spring through mid-summer (Berrien and Sibunka
1999).
B. brosme eggs were collected on GLOBEC surveys from March to
July and were generally concentrated along the southern flank and on
the Northeast Peak of Georges Bank. Early season B.
brosme eggs were collected as isolated occurrences
along the periphery of the Bank in deep water. Egg distribution and abundance
in April and May indicate a marked increase in spawning activity. Egg
densities were highest along the southeast central portion of Bank, the
Northeast Peak, and Browns Bank. The egg catches during the April 1997
and 1998 surveys were the largest of all five years. By June, catches
had decreased, and the eggs collected were mostly concentrated in deep water in the eastern and southern
shelf-slope portion of the Bank. Small catches of eggs were also made
in the Great South Channel area in June 1997 and 1998. The July survey
in 1995 collected a small amount of B. brosme eggs at one station in deep water in the north
central area of Georges Bank.
Atlantic cod, Gadus morhua (Figure 12)
Adult Gadus morhua are found along the North American continental
shelf from northern Quebec to about Cape Hatteras, NC. They are most
abundant from Labrador to southern Cape Cod (Bigelow and Schroeder 1953). G.
morhua eggs were collected on Georges Bank during the MARMAP surveys
in all months of the year except September. The highest egg densities
on the Bank occurred between December and April (Berrien and Sibunka
1999).
G. morhua eggs were collected on the GLOBEC surveys from January
to June and were sampled over the entire Bank, with the highest densities
on the Northeast Peak area. The wide distribution and large number
of G. morhua eggs in the January broad-scale surveys
indicated that spawning was well underway. The highest January abundances
of eggs were encountered during 1998 and 1999, when the eggs were more widespread than in the other three years.
Egg abundances were lowest in January 1997; these abundances also coincided
with the most limited spatial distribution, but the low catch may be
attributed to reduced sampling effort during this survey. The highest
egg abundances occurred in February and March, indicating that peak spawning
occurs at this time. Eggs were widespread in March, with high egg catches
occurring on the Northeast Peak and the southern flank. G. morhua egg
abundances were lower in April than in February and March, indicating
that peak spawning had passed. These eggs were concentrated along the
southern portion of the Bank. Only three of the five June surveys collected
eggs, and catches were both small in number and isolated in distribution.
For additional information regarding G. morhua egg distribution
and abundance during the GLOBEC program, refer to Mountain et al. (2003;
unpubl. data2).
Haddock, Melanogrammus aeglefinus (Figure 13)
Adult Melanogrammus aeglefinus range along the North American
coast from the Strait of Belle Isle off Labrador to southern New Jersey
(Klein-MacPhee 2002b). They are most abundant from the eastern Nova Scotian
Banks to Cape Cod (Bigelow and Schroeder 1953). M. aeglefinus eggs were
collected from January to July during the MARMAP surveys, and the highest
egg densities occurred in March and April (Berrien and Sibunka 1999).
M. aeglefinus eggs were collected on the GLOBEC surveys
from January to June and were found over the entire Bank. The highest
egg densities occurred on the Northeast Peak and the southern flank of
Georges Bank. The catch results for the January surveys indicate that M.
aeglefinus spawning was either already underway at this time
or had just begun. Most of the eggs collected during January and February
were concentrated on the Northeast Peak and eastern Bank area. By March
and April the distribution of M. aeglefinus eggs progressed from
the Northeast Peak (in February) southwest to the south central Bank
region. Additional egg concentrations also occurred on the northwest
Bank area and in the Great South Channel. March and April were the peak
spawning months for all survey years. The egg catch results for May and
June indicate that spawning had begun to decrease on Georges Bank. The
May results for 1995 to 1997 show spawning activity limited mostly to
the Northeast Peak and eastern Bank. The May results for 1998 and 1999
show that spawning occurred over a broader area than it did during the
previous three years. At this time, the M. aeglefinus eggs
were distributed from the Northeast Peak southwest to about the central
Bank area with additional collections along the Bank’s northern
edge. Egg catches in June show a further decrease
in spawning activity, limited to the eastern Bank region. The year 1995
was the poorest for M. aeglefinus spawning, followed by increased
spawning activity in 1996 and 1997, and with the highest egg catches
made in 1998 and 1999. M. aeglefinus eggs were also
collected on Browns Bank during all survey years. For additional information
regarding M. aeglefinus egg distribution and abundance during
the GLOBEC program refer to Mountain et al. (2003; unpubl. data3).
Pollock, Pollachius virens (Figure 14)
Adult Pollachius virens range from the Hudson and Davis Strait
in northern Canada to South Carolina; however, they are more common in
areas north of New Jersey (Bigelow and Schroeder 1953; Klein-MacPhee
2002b). P. virens eggs were collected on Georges Bank during the
MARMAP surveys from October to May, and the highest egg densities occurred
in December and January (Berrien and Sibunka 1999).
P. virens eggs were collected on all GLOBEC January surveys.
The largest egg concentrations in January were located on the Northeast
Peak region. In addition, smaller egg catches were made on the Bank on
different survey years. P. virens eggs were distributed in January
1996 from the Northeast Peak to the south-central Bank area. Eggs collected
in the January 1998 and 1999 surveys ranged from the Northeast Peak west
along the northern portion of Georges Bank. The January 1997 egg catches,
mainly concentrated in the Northeast Peak area, had the smallest distribution
and lowest abundance compared to the other January surveys. The egg catch
results for all the February surveys except 1997 indicate that spawning
activity may have peaked in January or perhaps earlier and was subsiding
by February. The 1997 February P. virens egg abundance and
distribution were greater than that of the January survey. Egg abundances
were the highest in February of all surveys made in that year. P.
virens eggs collected in February were concentrated in
the Northeast Peak area, with smaller egg catches made in the northwest
Bank area. The results for the other February surveys also showed that
the largest egg concentration occurred on the Northeast Peak region,
with smaller collections of egg catches made on the Bank’s
northwest region. Egg collections further declined to
small and isolated during the March and May surveys on the Bank. P.
virens eggs were also collected
on Browns Bank on several of the January to March surveys. No P. virens eggs
were collected in June or during the one July survey.
Goosefish, Lophius americanus (Figure 15)
Adult Lophius americanus range off the Canadian and United States
coast from the southern and eastern Grand Banks area and the north side
of Gulf of St. Lawrence south to Florida, but they are more common north
of Cape Hatteras (Caruso 2002). Spawning L. americanus shed their eggs in a
large (1x10 m) buoyant veil. Individual eggs may become separated and
float free in the water column if the integrity of the veil is compromised
(Bigelow and Schroeder 1953). L. americanus eggs and larvae were
rarely collected on the Bank during the MARMAP program sampling (Steimle
et al. 1999).
L. americanus eggs were collected on two June surveys during
the GLOBEC program. Both occurrences were single late-stage eggs. The
egg collected in June of 1995 was from the northwest area of the Bank.
The egg collected in 1997 was from the southwest flank region of the
Bank.
Northern searobin, Prionotus carolinus [tentative identification]
(Figure 16)
Adult Prionotus carolinus range from the Bay of Fundy to Florida.
They are common south of Cape Cod on the continental shelf, but also
occur in Massachusetts Bay and on Georges Bank (Klein-MacPhee and McBride
2002; refer to Map 23). Searobin eggs were collected on Georges Bank
during the MARMAP surveys from July to August. The catch occurrences
on the Bank were sporadic and the corresponding eggs densities small
(Berrien and Sibunka 1999; taxa referred to as Prionotus spp.).
P. carolinus eggs were first collected on the June 1998 and 1999
GLOBEC surveys. These P. carolinus eggs were concentrated in the shoal areas on the north central portion of
Georges Bank. The results of the one July survey show that spawning intensity had increased and was again mainly located
on the northern shoal area. A small catch of eggs was also taken on the
Bank’s southwest region during this time.
Identification Notes
Along the United States east coast searobins of the genus Prionotus comprise
two species: the northern searobin (P. carolinus) and the striped sea robin (P.
evolans). Adult P. evolans are common south of Cape Cod from
estuaries seaward to the edge of the continental shelf (Able and Fahay
1998; Klein-MacPhee and McBride 2002). North of Cape Cod their occurrence
is incidental, and they are found inshore close to the coast. Reported
evidence of striped sea robins on Georges Bank is rare; Bigelow and Schroeder
(1953) noted a specimen collected from the eastern portion of the Bank
and stated that they had not seen a specimen east of Cape Cod.
The frequency of occurrence of larval Prionotus spp. on Georges
Bank from the MARMAP surveys is low, and those specimens identified to
species were all P. carolinus (unpubl. data4).
No larval Prionotus spp. were collected on the
GLOBEC broad-scale surveys. It is most likely that the Prionotus spp.
eggs collected on the broad-scale surveys are all P. carolinus.
Tilefish, Lopholatilus chamaeleonticeps (Figure 17)
Adult Lopholatilus chamaeleonticeps are most plentiful from southern
Georges Bank to the Gulf of Mexico, and they inhabit the seaward portion
of the continental shelf region (Able 2002). L. chamaeleonticeps eggs were collected on Georges
Bank during the MARMAP cruises from late spring through early fall (Berrien
and Sibunka 1999).
A single L. chamaeleonticeps egg was collected during
the April 1996 GLOBEC survey on the southwest shelf-slope of Georges
Bank. This early-stage egg was possibly spawned in deep water near or
off the Bank.
Cunner, Tautogolabrus adspersus (Figure 18)
Adult Tautogolabrus adspersus range along the North American
east coast and offshore banks from Newfoundland to the entrance of Chesapeake
Bay, MD (Bigelow and Schroeder 1953). T. adspersus eggs were collected on Georges
Bank during the MARMAP surveys in the summer months with the highest
densities occurring in July (Berrien and Sibunka 1999).
T. adspersus eggs were first collected during June on the GLOBEC
broad-scale surveys, and distributions and abundance varied over the
five survey years. Egg densities were low, and distributions were limited
in 1995 and 1996. The 1996 egg catch results were the smallest for all
five June surveys. In 1995 and 1996 T. adspersus eggs were collected
on the north central shoal area of the Bank. The June 1997 to 1999 egg
catch results showed both a dramatic increase in spawning intensity and
in area. Catches of T. adspersus eggs occurred across the Bank’s
northern half and the entire eastern region including the Northeast Peak.
There were only small isolated occurrences of T. adspersus egg
catches in the southwest Bank area during most of the June surveys. T.
adspersus egg catch results from the only July survey show an increase
in both spawning abundance and distribution for that year. The July catch
resembles the June 1997 to 1999 surveys in terms of egg density and distribution.
Atlantic mackerel, Scomber scombrus (Figure 19)
The range of adult Scomber scombrus is from southern Labrador
to Cape Lookout, NC (Collette 2002). Spawning occurs in June off Cape
Cod and north in United States waters and is usually confined near the
coast (Bigelow and Schroeder 1953). S. scombrus eggs were collected on Georges
Bank during the MARMAP surveys from May to August. The highest egg densities
occurred in May and June on the Bank (Berrien and Sibunka 1999).
S. scombrus eggs were most abundant during the late spring on
the GLOBEC surveys, and they were distributed across the Bank usually
in small numbers. Spawning generally began in May with small and sporadic
egg catches either along the southern edge of Georges Bank in 1995 or
along the northern Bank area and the Northeast Peak in 1997-1999. S.
scombrus eggs were also taken on Browns Bank in 1995. The May 1996
survey did not collect S. scombrus eggs. Egg catches in June
varied between survey years as to area of capture and abundance. The
results of 1995 and 1998 showed that S. scombrus eggs were mainly concentrated along
the north and across the south central Bank region. S. scombrus eggs
were also collected on Browns Bank in 1996. Egg catches for the years
1996 to 1999 indicated spawning occurred mostly in the northern and the
north central Bank region, with small and intermittent egg occurrences
along the southern Bank area. The July 1995 survey showed that S.
scombrus spawning had decreased, with only two small catches of eggs
taken near the edge of the northwest Bank.
Butterfish, Peprilus triacanthus (Figure 20)
The range of adult Peprilus triacanthus is from Nova Scotia to
the Carolinas. They are summer residents and spawners in the New England
offshore waters (Bigelow and Schroeder 1953). P. triacanthus eggs
were collected during the MARMAP surveys on Georges Bank from May to
September. The highest abundances occurred in July and August (Berrien
and Sibunka 1999).
The GLOBEC broad-scale program first collected P. triacanthus eggs
in the spring during the May 1995 and 1999 surveys and in June 1998.
No P. triacanthus eggs were collected in 1996 or
in 1997. The few isolated egg catches from the May and June 1995 and
1999 surveys were taken from slope waters along the southern edge of
Georges Bank. The July 1995 survey results are similar to those from
the previous June, with P. triacanthus eggs again concentrated on
the Bank’s shoals. Isolated egg catches were also made during this
July survey on the Bank’s southeastern flank region and on the
Northeast Peak. In contrast, P. triacanthus eggs collected
in June 1998 were from the north central area of the shoals.
Windowpane flounder, Scophthalmus aquosus (Figure 21)
Adult Scophthalmus aquosus range from the Gulf of St. Lawrence
to Florida. Their areas of abundance are on Georges Bank and in the New
York Bight (Klein-MacPhee 2002d). S. aquosus eggs were collected
during the MARMAP surveys on Georges Bank from April to October. The
highest egg densities occurred during the summer months (Berrien and
Sibunka 1999).
S. aquosus eggs were most abundant during the GLOBEC surveys
in early summer on Georges Bank. Spawning began as early as April in
1997 to 1999, with small isolated catches occurring in the northern and
shallower area. By May and June spawning intensity had increased, extending
across the north, and north central region of the Bank to the Northeast
Peak area, with isolated catches of eggs along the Bank’s southern
edge. The May 1999 survey indicated S. aquosus eggs distributed
from the central Bank area to the southwest in a narrow belt toward the
Great South Channel. The July 1995 survey results indicated intense spawning
activity still occurring across most of the Bank. The distribution and
abundance of eggs for the July 1995 survey were similar to those for
June 1995.
Gulf Stream flounder and smallmouth flounder, Citharichthys arctifrons and Etropus
microstomus (Figure 22)
Adult Citharichthys arctifrons (Gulf Stream flounder) and Etropus
microstomus (smallmouth flounder) are most abundant from Cape Cod
to Cape Hatteras. C. arctifrons inhabit the outer portion of
the continental shelf whereas E. microstomus occur on the inner
shelf region (Able and Fahay 1998; Klein-MacPhee 2002e). The eggs of
these two species are difficult to differentiate (see Identification
Notes below), so are here referred to as C. arctifrons/E. microstomus eggs. C.
arctifrons/E. microstomus eggs were collected on
Georges Bank during the MARMAP surveys from April to October. The highest
egg densities occurred in August; those eggs were distributed across
most of the Bank (Berrien and Sibunka 1999; taxa referred to as Citharichthys/Etropus spp.).
C. arctifrons/E. microstomus eggs were caught as early as May
during the 1995, 1997, and 1999 GLOBEC broad-scale surveys. The distribution
of these eggs was concentrated along the Bank’s shelf-slope. Small
numbers of eggs were also collected from the Northeast Peak in 1997 and
in the Great South Channel in 1999. Catch results for the June 1995,
1997, and 1999 surveys indicate an increase in spawning intensity over
the previous month. C. arctifrons/E. microstomus eggs were collected
in June 1996 on the south-central Bank and in June 1998 along the southern
flank region and above the shoals. In 1996, C. arctifrons/E. microstomus egg
distribution and abundance were the lowest of all the five survey years.
The catch results for the other four June surveys indicate an increase
in egg distribution to the Great South Channel and the Bank’s southern
flank with the largest egg collections made on the south central and
southern shelf-slope to the southwest center area. The single July survey
catch results in 1995 show a further increase in spawning intensity on
the Bank, from the entire south-central and west Bank areas to the northern
Bank above the shoals and in the Great South Channel. The only occurrence
of C. arctifrons/E. microstomus eggs was taken from Browns Bank
during the May to July 1995 surveys.
Identification Notes
The eggs classified as C. arctifrons/E. microstomus are
difficult to differentiate. Adult C. arctifrons spawn mainly
during the summer and early fall, but may spawn intermittently throughout
the year. They range along the U.S. northeast coast from the southwest
portion of Georges Bank to Florida’s Gulf coast at water depths
greater than 37 m (Klein-MacPhee 2002e; refer to Map 35). Scott and Scott
(1988) reported that adult C. arctifrons are found further north
along the outer continental shelf waters to the Scotian Shelf. E.
microstomus also spawn in summer to early fall. They range from Cape
Cod southward in depths generally less than about 37 m (Klein-MacPhee
2002e). The larval distribution maps based on the MARMAP surveys do not
show any E. microstomus occurrences on Georges Bank
(Able and Fahay 1998). Scott and Scott (1988) reported small catches
of smallmouth flounder larvae on the southeast portion of the Bank in
the spring. It has been stated that larval E. microstomus occur
as a stray on the Bank. Two E. microstomus larvae were collected
by the 1 m2 MOCNESS (Multiple Opening Closing Net Environmental
Sampling System) net on the south central Bank area during the April
1999 broad-scale survey. There were several larval occurrences of C.
arctifrons larvae and a single catch of E. microstomus from
the broad-scale surveys (U.S. GLOBEC 2005). It is plausible that a limited
amount of mus spawning does occur on Georges Bank, possibly on
the shoals from a local population. C. arctifrons are assumed
to spawn on the shelf-slope of the Bank. It is very likely that the majority
of the C. arctifrons/E. microstomus eggs collected
during the broad-scale program are C. arctifrons, particularly
those along the shelf-slope of Georges Bank.
Summer flounder, Paralichthys dentatus (Figure 23)
Adult Paralichthys dentatus range from Nova Scotia to South Carolina,
but are most common south of Cape Cod to North Carolina (Able and Fahay
1998). P. dentatus eggs were collected
during the MARMAP surveys on Georges Bank in October and November, with
a small catch in January along the southwestern shelf slope (Berrien and
Sibunka 1999).
P. dentatus egg occurrences on the GLOBEC broad-scale program were
sporadic, and the number of eggs collected was very small. No eggs were collected
on the 1995 and 1997 surveys. One late-stage P. dentatus egg was
taken from the south central slope-edge of the Bank in May 1996. A single
late-stage egg was collected in April 1999 at one station in the extreme
southwestern Bank area. During the subsequent May survey, small catches of
eggs were taken at two stations located in the southwest portion of the Bank.
The June 1998 survey collected a small number of eggs at one station located
in the shoal region on the center of the Bank. P. dentatus eggs in
June 1999 were again collected at one station on the northern
shoals and at two stations in the southwest portion of the Bank.
Identification Notes
The identification of all P. dentatus eggs except for one early stage
was based on late stage development. A few small larvae were also collected
on the Bank during late spring in the same regions where eggs were caught
(see information regarding fish larval collections on the U.S. GLOBEC Georges
Bank website). Prior evidence of P. dentatus spawning in the spring on Georges
Bank has not been documented. Berrien and Sibunka (1999) reported P.
dentatus spawning during April and May in the southern portion of the
Middle Atlantic Bight, but no P. dentatus eggs were collected
north of this area. Their findings were based on the MARMAP study of fish
eggs.
Fourspot flounder, Paralichtys oblongus (Figure 24)
Adult Paralichthys oblongus range from Georges Bank to Tortugas,
Florida (Gutherz 1967). They are abundant from southern New England to Delaware
Bay (Bigelow and Schroeder 1953). P. oblongus eggs were collected
during MARMAP surveys on Georges Bank from May to October, with the greatest
egg densities occurring between July and August (Berrien and Sibunka 1999).
P. oblongus eggs were caught on GLOBEC surveys from May to July and
were collected over the entire Bank. The earliest occurrences of eggs were
during the May 1995 and 1999 surveys. These catches occurred along the southern
shelf-slope of Georges Bank at only one station in 1995 and several stations
in 1999. The catch results of the five June surveys varied from small numbers
of eggs collected at a few stations in 1995 and 1996 to a substantial increase
in egg catches in 1997, 1998, and 1999, indicating a rise in spawning activity
on the Bank. The three latter June surveys had similar egg distribution patterns.
Egg catches occurred along the southern edge, and north across the Bank’s
west central area extending onto the shoals. P.
oblongus eggs were also collected during the June 1995, 1997,
1998, and 1999 surveys in the southern part of the Great South Channel and
on Browns Bank in June 1996. The July 1995 results showed a marked increase
in egg distribution and abundance from the preceding June survey. Spawning
during this time extended across the western Bank and along the Bank’s northern
shelf and southern shelf-slope area.
Witch flounder, Glyptocephalus cynoglossus (Figure 25)
Adult Glyptocephalus cynoglossus range from Labrador to Cape Lookout
(Scott and Scott 1988). These fish are late spring and summer spawners in
the Gulf of Maine and New England offshore waters (Bigelow and Schroeder
1953). G. cynoglossus eggs were collected on Georges Bank during the
MARMAP surveys from April to October; however, the highest egg densities
occurred in May and June (Berrien and Sibunka 1999).
The GLOBEC broad-scale program collected G. cynoglossus eggs
on Georges Bank as early as March in 1998, but most eggs were taken later
during the spring in May and June. Eggs collected in May and June indicate
that most G. cynoglossus spawning occurred along the south-central
to southwest area of Georges Bank and in the Great South Channel. As the
spawning season progressed in 1998, eggs were collected on the northern Bank
area above the shoals during the June survey. The only occurrence of G.
cynoglossus eggs on Browns Bank was during the June 1999 survey. Both
the occurrences and the number of eggs collected were highest in 1997 to
1999 and lowest in 1995 and 1996. In 1996 G. cynoglossus eggs were
first taken during the June survey, which indicates that spawning on the
Bank started late that year.
American plaice, Hippoglossoides platessoides (Figure 26)
Adult Hippoglossoides platessoides range along North America from
near the Arctic Circle south into the Gulf of Maine and as an incidental
occurrence off Montauk Point on Long Island (Klein-MacPhee 2002f). H.
platessoides eggs were collected on Georges Bank during the MARMAP
surveys from January to June, with the highest egg densities in April (Berrien
and Sibunka 1999).
H. platessoides eggs were collected from January to June during the
GLOBEC surveys. The highest egg densities were along the southern half of
the Bank during March and April. Spawning of H. platessoides was already
underway on Georges Bank during the January surveys. The egg catches during
the January and February surveys show most spawning occurred in the Northeast
Peak area and on the eastern portion of the Bank. Small numbers of eggs were
also collected intermittently during this time at stations along the northwest
Bank area. The H. platessoides egg catches for March and April
indicate spawning intensity had increased on the Bank. During this
time spawning progressed in a westward direction from the Northeast Peak
and was mostly concentrated along the central and southern portion of the
Bank. A few small catches of eggs were also taken on the northwest
Bank. May and June data indicate that spawning had peaked and
was now subsiding, with most catches occurring in the east-central Bank
area. There were no H. platessoides eggs collected during the one July survey.
Yellowtail flounder, Limanda ferruginea (Figure 27)
Adult Limanda ferruginea range from the strait of Belle Isle to the
lower Chesapeake Bay. They are most abundant from the western areas
of the Gulf of Maine and Georges Bank to Southern New England (Bigelow and
Schroeder 1953). L. ferruginea eggs were collected on Georges
Bank during the MARMAP surveys from March to September. The highest
egg densities occurred during May and June. (Berrien and Sibunka 1999;
taxon referred to as Pleuronectes ferrugineus).
L. ferruginea eggs were collected on GLOBEC surveys from March to
July and were caught over the entire Bank. The greatest egg distribution
occurred in April to June, and the highest densities were on the Bank’s
eastern portion. L. ferruginea eggs were first collected in
February for all years of the GLOBEC surveys, except in February 1995 when
no eggs were caught. The egg catch results for February indicate that
spawning occurred in the Bank’s eastern portion. Eggs were
also collected on the northwest area in 1997 and 1999. Spawning intensity
increased during the months of March to May with egg catches now made on
most of the Bank and also in the Great South Channel. The largest concentration
of L. ferruginea eggs during this time period was on the eastern Bank
with additional concentrations in May on the northwest Bank. The catch
results for the June surveys indicate that spawning intensity was still high,
particularly during 1995 and 1996. Results for June 1997, 1998, and
1999 surveys indicate that spawning on the Bank had decreased and was not
as widespread as during the May surveys for those years. The abundance-distribution
plots for the June surveys indicate that most spawning is concentrated on
the eastern Bank, with some large egg catches made in the northern portion
of the Great South Channel. The July survey results indicate a dramatic
decrease in L. ferruginea spawning on the Bank, with the largest egg
catches taken on the Northeast Peak. Egg catches were also made on
Browns Bank during late spring and early summer except in 1997, when no eggs
were caught.
The survey results indicate the greatest amount of spawning activity
for L.
ferruginea occurred during 1998 and 1999. These two years appear
to be similar during the spring for egg distribution and abundances.
In these years, intense spawning occurred in May across most of the
Bank (Figure 27). The eastern region was again the primary area for L.
ferruginea spawning,
followed by the northwest area and Great South Channel. The southwest
area had the lowest egg catches at this time. The least amount
of spawning took place during 1995.
Winter flounder, Pseudopleuronectes americanus (Figure 28)
Adult Pseudopleuronectes americanus range from northern Labrador
to Georgia, and are common from the Gulf of St. Lawrence to the Chesapeake
Bay. P. americanus are reported to form localized populations
within their distribution range. One population inhabits Georges
Bank; these fish are known in the commercial fishing industry as lemon
sole. The onset of P. americanus spawning on the Bank is unknown,
but they are known to spawn during April and May. P. americanus eggs
are demersal and adhesive, and they form clusters on the sea bed (Bigelow
and Schroeder 1953).
The P. americanus eggs collected on the GLOBEC broad-scale surveys
were incidental to the catch. They were probably the result of such
factors as fishing too close to the sea bed, strong atmospheric storm activity
or strong cycloidal currents that agitated the water column and suspended
eggs off the sea bottom, and disturbance by bottom feeding adult fish which
may disturb the sea bed and suspend the demersal eggs. These data,
therefore, should not be considered as a representative index for spawning
abundance or as a definitive reference to egg distribution on the Bank,
but rather to indicate that Bank-wide spawning occurrences and give limited
information on where P. americanus eggs are found. Results for the five
years of survey data show that while most of the P. americanus eggs
were collected in March and April, small numbers were also caught both earlier
and later in the season. Most eggs were collected on the central shoal
Bank area.
ENDNOTES
- Available from: J. Sibunka, National Marine Fisheries
Service, Highlands NJ.
- Mountain D, Green J, Sibunka J, Johnson D. In
prep. Growth and mortality of Atlantic cod (Gadus morhua)
and haddock (Melanogrammus aeglefinus) eggs and larvae on Georges
Bank, 1995-1999.
- ibid.
- Available from: J. Sibunka, National Marine Fisheries
Service, Highlands NJ.
ACKNOWLEDGMENTS
The authors express a sincere thanks to all the sea-going scientific
personnel from the GLOBEC Georges Bank broad-scale program. It was their
efforts and those of the vessel crews of the R/V Albatross IV, R/V
Endeavor,
and R/V Oceanus, that made this research possible. A large amount
of gratitude is reserved for Tom Finneran who worked tirelessly to develop
the ‘perfect’ basemap.
The authors acknowledge their appreciation to Dr. Jonathan A. Hare for
reviewing an early draft of the atlas and for his suggested improvements
to the manuscript. We thank Robert N. Reid, Dr. R. Christopher Chambers,
and three anonymous reviewers whose comments helped make this atlas possible.
LITERATURE CITED
Able KW. 2002. Tilefishes. Family Malacanthidae. In: Collette BB, Klein-MacPhee
G, editors. Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd
ed. Washington (DC): Smithsonian Institution Press, p. 397-400.
Able KW, Fahay MP. 1998. The first year in the life of estuarine fishes
in the Middle Atlantic Bight. New Brunswick (NJ): Rutgers University Press,
342 p.
Berrien P, Sibunka J. 1999. Distribution patterns of fish eggs in the northeast
continental shelf ecosystem. Seattle (WA): National Oceanic and Atmospheric
Administration Technical Report NMFS-145, 310 p. Available from: NTIS, Springfield
VA; PB83-210203.
Bigelow HB, Schroeder WC. 1953. Fishes of the Gulf of Maine. US Fish Wildl
Serv, Fish Bull. 53, 577 p.
Caruso JH. 2002. Goosefishes or monkfishes. Family Lophiidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes of the
Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution Press, p.
264-270.
Collette BB. 2002. Mackerels. Family Scombridae. In: Collette BB, Klein-MacPhee
G, editors. Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd
ed. Washington (DC): Smithsonian Institution Press, p. 516-536.
Collette BB, Klein-MacPhee G. 2002. Cusk-eels. Family Ophidiidae. In:
Collette BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s
Fishes of the Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution
Press, p. 208-211.
Fahay MP. 1992. Development and distribution of cusk eel eggs and larvae
in the Middle Atlantic Bight with a description of Ophidion robinsi n.sp.
(Teleostei: Ophidiidae). Copeia 1992:799-819.
Gutherz EJ. 1967. Field guide to the flatfishes of the family Bothidae in
the western North Atlantic. US Fish Wildl Serv Circ. 263, 47 p.
Harold AS, Hartel KE, Craddock JE, Moore JA. 2002. Hatchetfishes and relatives.
Family Sternoptychidae. In: Collette BB, Klein-MacPhee G, editors. Bigelow
and Schroeder’s Fishes of the Gulf of Maine. 3rd ed. Washington (DC):
Smithsonian Institution Press, p. 184-190.
Houde ED. 1977. Abundance and potential yield of the round herring, Etrumeus
teres, and aspects of its early life history in the eastern Gulf of
Mexico. Fish Bull (US) 75:61-89.
Jossi JW, Marak RR. 1983. MARMAP plankton survey manual. National Oceanic
and Atmospheric Administration Technical Memorandum NEFC-F/NEC-21, 258 p.
Available from: NTIS, Springfield VA; PB83-210203.
Klein-MacPhee G. 2002a. Argentines. Family Argentinidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes
of the Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution
Press, p. 160-162.
Klein-MacPhee G. 2002b. Cods. Family Gadidae. In: Collette BB, Klein-MacPhee
G, editors. Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd
ed. Washington (DC): Smithsonian Institution Press, p. 223-261.
Klein-MacPhee G. 2002c. Silver hakes. Family Merlucciidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes of the
Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution Press, p
217-222.
Klein-MacPhee G. 2002d. Family Scophthalmidae. In: Collette BB, Klein-MacPhee
G, editors. Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd
ed. Washington (DC): Smithsonian Institution Press, p. 547-550.
Klein-MacPhee G. 2002e. Sand flounders. Family Paralichthyidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes of the
Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution Press, p.
551-560.
Klein-MacPhee G. 2002f. Righteye flounders. Family Pleuronectidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes of the
Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution Press, p.
560-587.
Klein-MacPhee G, McBride RS. 2002. Sea robins. Family Triglidae. In:
Collette BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s
Fishes of the Gulf of Maine. 3rd ed. Washington (DC): Smithsonian
Institution Press, p. 338-345.
Mountain D, Berrien P, Sibunka J. 2003. Distribution, abundance and mortality
of cod and haddock eggs and larvae on Georges Bank in 1995 and 1996. Mar
Ecol Prog Ser. 263:247-260.
Nelson JS, Crossman EJ, Espinosa-Prez H, Findley LT, Gilbert CR, Lea RN,
Williams JD. 2004. Common and scientific names of fishes from the United
States, Canada, and Mexico. 6th ed. Bethesda (MD): American Fisheries Society
Special Pub. 29, 386 p.
Posgay JA, Marak RR. 1980. The MARMAP bongo zooplankton samplers. J Northw
Atl Fish Sci. 1: 91-99.
Scott WB, Scott MC. 1988. Atlantic fishes of Canada. Can Bull Fish Aquat
Sci. 219: 534-544.
Sibunka JD, Silverman MJ. 1984. MARMAP surveys of the continental shelf
from Cape Hatteras, North Carolina to Cape Sable, Nova Scotia (1977-1983).
Atlas No. 1. Summary of operations. Natl Ocean Atmos Adm Tech Mem Natl Mar
Fish Serv.-F/NFC-33, 306 p.
Sibunka JD, Silverman MJ. 1989. MARMAP surveys of the continental shelf
from Cape Hatteras, North Carolina to Cape Sable, Nova Scotia (1984-87).
Atlas No. 3. Summary of operations. National Oceanic and Atmospheric Administration
Technical Memorandum NEFC-F/NEC-68, 197 p. Available from: NTIS, Springfield
VA; PB90160656/AS.
Smith DG, Tighe KA. 2002. Snake eels. Family Ophichthidae. In: Collette
BB, Klein-MacPhee G, editors. Bigelow and Schroeder’s Fishes of the
Gulf of Maine. 3rd ed. Washington (DC): Smithsonian Institution Press, p.
97-98.
Steimle FW, Morse WW, Johnson DL. 1999. Essential fish habitat source
document: goosefish, Lophia americanus, life history and habitat
characteristics. National Oceanic and Atmospheric Administration Technical
Memorandum NEFC-F/NEC-127, 31 p. Available from: NTIS, Springfield VA;
PB2000-103388.
US GLOBEC Georges Bank Program. Fish egg data base [Internet]. [cited
2005 May 20]. Available from: http://globec.whoi.edu.
