BULLETIN OF MARINE SCIENCE, 54(3): 828-842, [994
VERTICAL DISTRIBUTION OF FISH LARVAE
OFF THE FLORIDA KEYS, 26 MAY-5 JUNE 1989
Seong Sig Cha, Michael F. McGowan and William J. Richards
ABSTRACT
A preliminary description of the vertical distribution of fish larvae offshore of the Florida
Keys was derived from quantitative analysis of the fish families represented in the catch from
selected stations on the 26 May to 5 June 1989 cruise of Project SEFCAR (Southeastern
Florida and Caribbean Recruitment). Eight nighttime samples were taken at stations> 150 m
deep by MOCNESS tows with individual nets sampling 25 m vertical depth strata. Larvae
of 65 families of fishes were identified. The depth distribution of the 14 most abundant
families was explored. These families comprised three groups based on the observed depth
of 50% or more of their abundance. The Scombridae, Carangidae, Labridae, and Bothidae
occurred <25 m deep. The Gonostomatidae, Gobiidae, Myctophidae, Serranidae, Paralepididae, Scorpaenidae, and Bregmacerotidae occurred <50 m deep. The Paralichthyidae, Synodontidae, and Phosichthyidae occurred >50 m deep. The larvae of the remaining 51 families
also could be assigned to these three vertical depth categories with approximately equal
numbers of famili,~sin each depth category but greatest total abundance in the second category (0-50 m).
The Florida Current is that portion of the Gulf Stream system, which flows
northward through the Straits of Florida. It connects the Loop Current in the
eastern Gulf of Mexico to the Gulf Stream off the southeastern U.S. coast. The
axis of the current is located about 25 km offshore of Miami (Molinari and
Leaman, 1987) and 80 km offshore of Key West, where a mean westward countercurrent was observed on the shoreward side off Key West (Brooks and Niiler,
1975). The Florida Current and its interactions with coastal water affect the distribution of fish larvae from local and distant sources and their eventual recruitment to the Florida Keys.
The vertical distribution of fish larvae is affected by physical and biological
factors. Physical influences include light intensity (Woodhead and Woodhead,
1955; Blaxter, 1973; Neilson and Perry, 1990) and the hydrographic structure of
the water column (e.g., Lasker 1975; Ropke et aI., 1990, 1993). Biological influences include ontogenetic stage (Loeb, 1979; Fortier and Leggett, 1983; Sogard
et al., 1987; Fortier and Harris, 1989), concentrations of food (Hunter and Thomas, 1974; Lasker, 1975; Coombs et al., 1983; Fortier and Leggett, 1983, 1984;
Fortier and Harris, 1989), and inter- and intra-specific competition for resources
(Fortier and Harris, 1989). There are strong vertical gradients of these factors and
differences between the Florida Current and coastal waters. To reach an understanding of the effects of these factors on recruitment of fishes to the Florida
Keys we must first describe the vertical distribution of the fish larvae,
Ahlstrom (1959) found that the majority of the fish larvae occurred within the
mixed layer and upper thermocline off California and Baja California. Similar
results have been reported in other areas (Loeb, 1979; Kendall and Naplin, 1981;
Southward and Bar)', 1980; Southward and Barrett, 1983; Coombs et al., 1981;
Boehlert et al., 1985). However species have characteristic depth distributions
(Loeb, 1979, 1980; Kendall and Naplin, 1981; Sogard et al., 1987; Ropke, 1989),
The depth distribution often changes with ontogeny and this can affect advective
transport of the larvae.
The purpose of this study is to provide a preliminary description of the vertical
828
CHAETAL.:FLORIDA
KEYSFISHLARVAE
829
26.0
25.5
24.S
24.0
-82.S
-82.0
-11.S
-81.0
-80.S
-80.0
Longitude
Figure I.
Location of stations for the vertical distribution of fish larvae sampled with MOCNESS.
distribution of fish larvae by family near the Florida Keys. Recruitment of fishes
is dependent on advective transport of larvae and co-occurrence of fish larvae
with favorable predator-prey environments. These factors vary with depth, so
knowledge of the depth distribution of fish larvae is essential to understanding
the recruitment process. Understanding recruitment to the Florida Keys is the goal
of the SEFCAR Project, so this study aims to contribute to achieving that goal.
METHODS AND MATERIALS
Sampling was conducted on the RJV CALANUS
at 29 stations in the Florida Keys from 26 May
through 5 June 1989 during the first SEFCAR (Southeast Florida and Caribbean Recruitment Project)
cruise. XBT or CTD casts were done to measure water column temperature. For the sampling of fish
larvae, a I-m2 MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System, Wiebe
et aI., 1976) was employed. Flowmeter, angle sensor, and CTD sensors were attached to the net frame.
The MOCNESS has nine nets of 0.333-mm mesh aperture. The first in the set of nine nets was fished
obliquely from the surface down to 200 m, or to the deepest multiple of 25 m if the water depth at
the station was less than 200 m. Each subsequent net was towed obliquely through a 25 m stratum.
At stations shallower than 200 m, fewer than nine nets were used at a station. Towing speed was
approximately I m·s-'.
The samples were fixed immediately in 5% formalin buffered with marble chips and transferred to
70% ethanol within 48 h. Fish larvae were sorted from the samples in the laboratory and identified
to the lowest taxonomic level possible. Leptocephali were excluded from this study. The catch of each
taxon in a net was standardized to number per 1,000 m3 of seawater and also to number under 100
m2 of sea surface.
For the study of the vertical distribution of fish larvae off the Florida Keys, offshore stations, whose
familial compositions were relatively similar (McGowan et al., in prep.'), were selected from the 29
stations. To reduce biases due to net avoidance by larger individuals and more agile species (Ahlstrom
1959, 1971; Loeb, 1980; Clutter and Anraku, 1986), analyses were done only on samples collected
at night. Therefore, eight night stations whose depth was greater than ISO m were selected for anaI McGowan.
M. F.. S. S. Chao and W. J. Richards. (ms) Horizontal distribution of the fish larvae off the Florida Keys. May 26-June
5. 1989.
830
BULLETIN OF MARINE SCIENCE, VOL. 54, NO.3,
1994
Table 1. Station data for the study of the nighttime vertical distribution of fish larvae near the Florida
Keys
St.tion
Longitude
Latitude
7
15
27
38
39
48
49
50
-80.643
-81.317
-82.200
-81.350
-81.345
-80.962
-80.377
-80.414
24.537
24.360
24.292
24.410
24.407
24.383
24.805
24.817
Transect
Tennessee
Looe Key
Cosgrove
Looe Key
Looe Key
Sombrero
Davis
Davis
Distance
(km)
Depth
(m)
Date
Local time
22.65
21.02
22.21
15.55
15.84
26.39
13.97
11.96
207
213
237
185
185
222
195
166
5-28
5-28
5-30
6-01
6-02
6-03
6-04
6-03
00:37
19:02
22:25
22:03
03:17
20:40
04:33
23:06
Iyzing the depth distributions of fish larvae (Fig. 1). These stations ranged from 12.0-26.4 Ian distant
from the shoreline (Table 1). Table 1 shows the longitude and latitude, distance from the reefs, depth,
date, and time for these stations.
Mean abundance of families by depth stratum was used to further elucidate depth distributions of
individual families and patterns of co-occurrence at depth by groups of families.
RESULTS
Physical conditions were similar at the eight stations, e.g., sea surface temperatures were between 26.9 and 28.3°C. The mixed surface layer (MSL) was very
o
-
E 100
.I:
••Q.
•
C
200
10
15
20
Temperature
Figure 2.
25
(Oel
Vertical temperature profiles with station numbers noted on the chart.
30
831
CHA ET AL: FLORIDA KEYS FISH LARVAE
0-25
25-50
-••
50-75
E
75-100
.,
100-125
.I:
Q.
c
125-150
150-175
175-200
o
100 200 300 400 500 600 700
800
Abundance (No./ 1,000m3)
Figure 3.
Vertical distribution
of fish larvae; mean abundance of fish larvae at each stratum.
shallow at stations 7, 15, and 27 sampled during the earlier part of the cruise
(Fig. 2). The MSL was somewhat deeper, to about 35 m, at stations 38, 39, 48,
49, and 50. The thermocline was very wide and extended to near bottom. Its
gradient was approximately 0.08°C·m-1• Bottom temperatures were less than 15°C.
There were 154 taxa identified below the familial level. There were representatives of 65 families (Appendix 1). The most abundant 14 families in decreasing
order were Myctophidae, Gonostomatidae, Bregmacerotidae, Scombridae, Serranidae, Paralichthyidae, Phosichthyidae, Bothidae, Paralepididae, Gobiidae, Scorpaenidae, Synodontidae, Carangidae, and Labridae (Appendix 2). The most abundant 14 families occupied 86.5% of the total abundance of eight stations. The
remaining 51 families comprised only 6.7% of the total abundance. The remaining
6.8% were unidentified or only identified to order.
Fish larvae were most abundant in the first and second strata (0-25, 25-50 m)
(Fig. 3). In the first stratum 31.5% of the total fish larvae occurred and in the
second stratum 33.1 % occurred for a total of 64.6%. Below 50 m the abundance
decreased with depth. The number of families occurring in each stratum showed
the same trends as the abundance (Fig 4). In the first and second stratum 40 and
38 families occurred, respectively. From the third to sixth stratum 27, 26, 22, and
21 families occurred, respectively. In the seventh stratum 14 families occurred.
At the eighth stratum two families occurred: Myctophidae and Callionymidae.
Figure 5 shows the relative abundance of the 14 most abundant families in
each stratum. The families can be separated into three groups by the criterion at
which strata more than half of them were distributed. The first group had >50%
of larvae at the first stratum (0-25 m), the second group had >50% of the larvae
832
BULLETIN OF MARINE SCIENCE, VOL. 54, NO.3, 1994
0-25
25-~)0
50-~'5
--.•..
E
~
75-1()0
0-
100-1 :2!5
Gl
Q
125-15.0
150-175
175-200
0
10
20
30
40
50
No. of Families
Figure 4.
The number of families of fish larvae at each stratum.
100
...X
....•
t:0
",. I>
80
'I
u
c
II
'Q
I';'
I>
G)
60
C
DO-25m
25-50m
:lI
.a
oC
e
>
~
II
'i
IE:
o
40
ILl 50-75m
~ 75-100m
WI 100-125m
20
•
125-150m
150-175m
•
175-200m
IIIlIII
0
Is
~
Sem
c.,
Lob Bat
Gon Gob MJe S.,
PI.
Ser
Sr.
PII
SJn Pho
Families
Figure 5, Relative abundance of the 14 most abundant families by depth stratum. The legends are
as follows. Scm: Scombridae, Car: Carangidae, Lab: Labridae, Bot: Bothidae, Gon: Gonostomatidae.
Gob: Gobiidae, Myc: Myctophidae, Ser: Serranidae, PIe: Paralepididae. Scr: Scorpaenidae, Bre: Bregmacerotidae, PH: Paralichthyidae, Syn: Synodontidae, Pho: Phosichthyidae.
CHA ET AL.: FLORIDA
KEYS
FISH LARVAE
833
shallower than 50 m, the third group had >50% of the larvae deeper than 50 m.
The first group is Scombridae, Carangidae, Labridae, and Bothidae. The second
group is Gonostomatidae, Gobiidae, Myctophidae, Serranidae, Para]epididae,
Scorpaenidae, and Bregmacerotidae. The third group is Paralichthyidae, Synodontidae and Phosichthyidae.
The remaining 51 families also can be separated into three groups by the same
criteria as above. All 65 families can be separated into 20 families in the first
group, 20 families in the second group, and 25 families in the third group (Tab]e
2). The second group occupied 75.0% of the total abundance. The first and the
third occupied 13.2% and 11.8%, respectively. Although the number of the families of each group is similar, the second group is by far more abundant than the
first or the third group.
DISCUSSION
Larvae were generally most abundant in the first and second strata (0-25, 2550 m) which were in the mixed layer and upper thermocline. This trend for the
peak abundance of fish larvae to be centered near the thermocline is similar to
that found in other regions. Ah]strom (1959) found that 12 out of 15 species of
larvae lived in the upper mixed layer of California and Baja California. Similar
results were obtained for larvae off the U.S. east coast (Kendall and Nap]in, 1981)
and off the Oregon coast (Boeh]ert et aI., 1985). Loeb (1979) found the highest
abundance and diversity of fish larvae at the bottom of the seasonal mixed layer
in the North Pacific centra] gyre region. The majority of mackerel (Scomber scombrus) larvae in the eastern North Atlantic were taken in the upper 50 m, and
above the thermocline when one was present. In the North Sea more than 91 %
of larvae were taken above the thermocline (Coombs et aI., 1981). Mackere]
(Scomber scombrus) larvae occurred near the surface, in association with copepod
eggs, nauplii, and copepodites (Coombs et aI., 1983).
Food may be the primary attraction for fish larvae in different depths in the
water column. Most families of the first group we discovered have larvae which
are very effective visual predators growing very fast in the warm, near surface
water. Larvae in our third group probably grow slowly in the colder water at their
depth (>50 m).
Ah]strom (1959) and Loeb (1979) described the larvae distributed in deeper
water as being adapted early to their later mesopelagic life. In our study the two
subfamilies of Myctophidae occurred at different depths. Subfamily Myctophinae
was in the third group with 66.1 % deeper than 50 m. Subfamily Lampanyctinae
was in the second group with 75.4% shallower than 50 m. At the generic level,
all genera belonging to Myctophinae (Myctophum, Benthosema, Diogenichthys,
Hygophum, Gonichthys, and Centrobranchus) were in the third group. Among
Lampanyctinae, Lampadena and Taaningichthys were in the first group, Lampanyctus, Ceratoscopelus, and Diaphus were in the second group, and Notoscopelus
and Notolychnus were in the third group. This phenomenon that Myctophinae
larvae were more deeply distributed than Lampanyctinae larvae was also observed
in the North Pacific central gyre (Loeb, 1979) and in the tropical Pacific (Boehlert
et aI., 1992).
Although Gonostomatidae and Phosichthyidae are very close systematically,
they differ ecologically. The primitive Phosichthyidae has non-protracted metamorphosis and "white" photophore development (i.e., all photophores laid down
at once). The specialized Gonostomatidae have gradual, protracted metamorphosis
and gradual photophore development (Ah]strom et aI., 1984). Phosichthyidae were
834
BULLETIN OF MARINE SCIENCE, VOL. 54, NO.3,
1994
Table 2. Relative mean abundances of fish larvae by family in the first stratum (0-25 m), first and
second strata (0-50 m), and in the third through eighth strata (50-200 m)
Mean
abundance
(N·100 m-')
(}-25 m
(}-50 m
5(}-200 m
(%)
(%)
(%)
Istiophoridae
Clupeidae
Sphyraenidae
Dactylo]lteridae
Notosudidae
Exocoetidae
Aulopididae
Chaetodl)ntidae
Echeneididae
Trichiuridae
Scombridae
Carangidae
Coryphaenidae
Labridae
Acanthuridae
BOthidae
Nomeidae
Scaridae
Lutjanidae
Balistidae
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
94.7
92.4
78.4
77.8
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
84.9
93.0
0.0
0.0
0.0
0.0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
15,1
7.0
68.8
100.0
0.0
3.4
68.4
58.0
57.4
56.6
56.4
90.3
78.1
57.4
76.0
75.2
9.7
21.9
42.6
24.0
24.8
74.8
12.7
7.8
6.1
6.3
II
Cynoglossidae
Apogoniclae
Bramidae
Malacosteidae
Holocentridae
Gobiesocidae
Gonostomatidae
Serranidae
Ophidiidae
Pomacentridae
Triglidae
Acropomaltidae
Gobiidae
Ogcoceph,alidae
Paralepididae
Bregmacerotidae
Myctophiclae
Melanostomiidae
Ceratiidae
Scorpaenidae
40.6
0.0
0.0
0.0
0.0
0.0
43.9
22.1
0.0
46.7
0.0
0.0
38.2
0.0
12.8
4.5
21.9
33.3
0.0
10.8
100.0
100.0
100.0
100.0
100.0
100.0
82.6
81.7
77.3
75.0
74.1
74.1
69.0
63.0
59.6
58.5
55.9
54.7
54.0
53.6
0.0
0.0
0.0
0.0
0.0
0.0
17.4
18.3
22.7
25.0
25.9
25.9
31.0
37,0
40.4
41.5
44.1
45.3
46.0
46.4
2.0
6.0
2.1
1.4
1.2
1.1
642.7
199.5
5.1
3.8
10.8
4.9
54.4
2.3
55.1
351.8
1,763.0
6.0
2.0
52.7
III
Stomiidae
Gempylidae
Tetraodontidae
Paralichthyidae
Phosichthyidae
Malacanthidae
Caproidae
Synodontid.ae
Priacanthidae
Antennariidae
Callionymidae
Melamphaidae
Argentinidae
Sternoptychidae
Percophidae
Bathylagidae
46.4
10.0
44.2
11.3
2.4
11.2
35.9
7.0
0.0
0.0
20.7
0.0
4.9
0.0
0.0
0.0
46.4
44.8
44.2
41.4
41.3
38.7
35.9
32.9
30.6
29.5
28.5
23.5
4.9
0.0
0.0
0.0
53.6
55.2
55.8
58.6
58.7
61.3
64.1
67.1
69.4
70.5
71.5
76.5
95.1
100.0
100.0
100.0
1.8
19.7
3.8
157.3
105.6
20.9
5.3
48.3
4.5
11.4
17.8
10.1
17.1
22.5
16.0
8.5
Group
Family
2.5
2.3
1.8
1.7
1.1
0.8
0.8
0.8
0.8
0.8
332.0
43.8
18.1
41.3
835
eRA ET AL.: FLORIDA KEYS FISH LARVAE
Table 2. Continued
Mean
Group
Family
Chaliodontidae
Carapidae
Macrouridae
Scopelarchidae
Chiasmodontidae
Caulophrynidae
Uranoscopidae
Epigonidae
Chlorophthalmidae
0-25 m
0-50 m
50-200 m
abundance
(%)
(%)
(%)
(N·JOO m-')
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
100.0
8.4
8.1
4.6
1.7
1.4
1.2
1.2
0.9
0.8
distributed deeper, in general, than Gonostomatidae in our samples. The few ex:;
ceptions were not abundant. For example, Cyclothone, comprising 95.4% of Gonostomatidae was in the second group and Gonostoma was in the third group.
We observed several other differences in depth distribution of families. Bothidae and Paralichthyidae are both sinistral flatfishes (eyes on the left side of the
head). Paralichthyidae were distributed deeper than Bothidae. Bothidae were in
the first group with 68.4% shallower than 25 m. ParaIichthyidae were in the third
group with 58.6% deeper than 50 m. Among Bregmacerotidae (which as a family
was in the second group) Bregmaceros houdei was in the second group but the
rare B. atlanticus and B. cantori were in the third group. Among Gempylidae (the
third group) Diplospinus and Nesiarchus were in the third group; Gempylus was
in the second group.
Among the third group, the relative abundances of Stomiidae, Tetraodontidae,
Caproidae, and Callionymidae were relatively high at the first stratum. Although
the observed pattern may reflect real differences in depth distributions, the abundances of some families were extremely low in our collections. More data are
needed to see if the observed patterns generally hold near the Florida Keys. In a
2.5-year study of the Flower Gardens in the northwest Gulf of Mexico, McGowan
(1985) found seasonally averaged vertical distributions of larvae by family similar
to those reported here, although relative abundance of families changed with season. The larvae in the Florida Keys are expected to show some seasonal differences in species composition and may show ontogenetic seasonal changes in
vertical distribution for species with highly seasonal spawning.
Ahlstrom (1959) and Badcock and Merrett (1976) showed that the larvae of
some midwater fishes often undergo limited diurnal vertical migrations. Most
species of fish larvae are able to cover distances of the order of 100 m within
hours and in several studies, fish larvae have been shown to migrate vertically
(Smith et aI., 1978; Boehlert et aI., 1985; Brewer and Kleppel, 1986; Fortier and
Leggett, 1983; Sogard et aI., 1987). Day patterns may, therefore, be altered somewhat from those presented here.
ACKNOWLEDGMENTS
We thank the members of the SEFCAR project (c. Paris, D. Goldman, and C. Yeung) for collecting,
sorting, and aiding in the identification of the material. Dr. A. Ropke kindly reviewed the manuscript
and offered valuable advice. The funding for this project was provided through NOAA Cooperative
Agreement No. NA90RAH00075 with the University of Miami.
LITERATURE
CITED
Ahlstrom, E. H. 1959. Vertical distribution of pelagic fish eggs and larvae off California and Baja
California. U.S. Fish Wildl. Serv., Fish. Bull. 60: 107-146.
836
---.
BULLETIN OF MARINE SCIENCE. VOL. 54. NO.3, 1994
1971. Kinds and abundance of fish larvae in the eastern tropical Pacific, based on collections
made on EASTROPAC I. Fish. Bull., U.S. 69: 33-77.
---,
W. J. Richards and S. H. Weitzman. 1984. Families Gonostomatidae, Sternoptychidae and
associated stomiiform groups: development and relationships. Pages 184-198 in Ontogeny and
systematics of fishes. The American Society of Ichthyologists and Herpetologists Spec. Publ. No.
1.760 pp.
Badcock, J. and N. R. Merrett. 1976. Midwater fishes in the eastern North Atlantic. I. Vertical
distribution and associated biology in 30oN, 23°W, with developmental notes on certain myctophids. Prog. in Oceanogr. 7: 3-58.
Blaxter, J. H. S. 1973. Monitoring the vertical movements and light responses of herring and plaice
larvae. J. BioI. A:5s.U.K. 53: 635-647.
Boehlert, G. W., D. M. Gadomski and B. C. Mundy. 1985. Vertical distribution of ichthyoplankton
off the Oregon coast in spring and summer months. Fish. Bull. U.S. 83: 611-621.
---,
W. Watson and L. C. Sun. 1992. Horizontal and vertical distributions of larval fishes around
an isolated oceanic island in the tropical Pacific. Deep Sea Res. 39: 439-466.
Brewer, G. D. and G. S. Kleppel. 1986. Diel vertical distribution of fish larvae and their prey in
nearshore waters of southern California. Mar. Ecol. Prog. Ser. 27: 217-226.
Brooks, I. and P. P. Niiler. 1975. The Florida Current at Key West. J. Mar. Res. 33: 83-92.
Clutter, R. I. and M. Anraku. 1986. Avoidance of samplers. Pages 57-76 in D. J. Tranter and J. J.
Fraser, eds. Zoopl.ankton sampling. UNESCO Paris. Monogr. Methodol. 2.
Coombs, S. H., R. K. Pipe and C. E. Mitchell. 1981. The vertical distribution of eggs and larvae of
blue whiting (Micromesistius poutassou) and mackerel (Scomber scombrus) in the eastern North
Atlantic and North Sea. Rapp. P.-v. Reun. Cons. Tnt. Explor. Mer 178: 188-195.
Coombs, S. H., J. A. Lindley and C. A. Fosh. 1983. Vertical distribution of larvae of mackerel
(Scomher scombrus) and microplankton, with some conclusions on feeding conditions and survey
methods. FAO Fish. Rep. 291: 939-954.
Fortier, L. and R. P. Harris. 1989. Optimal foraging and density-dependent competition in marine fish
larvae. Mar. Ecol. Prog. Ser. 51: 19-33.
--and W. C. Leggett. 1983. Vertical migrations and transport of larval fish in a partially mixed
estuary. Can. J. Fish. Aquati. Sci. 40: 1543-1555.
--and ---.
1984. Small-scale covariability in the abundance of fish larvae and their prey.
Can. J. Fish. Aquati. Sci. 41: 502-512.
Hunter, J. R. and G. L. Thomas. 1974. Effect of prey distribution and density on the searching and
feeding behavior of larval anchovy Engraulis mordax Girard. Pages 559-754 in J. H. S. Blaxter,
ed. The early life history of fish. Springer-Verlag, Berlin. 765 pp.
Kendall, A. W., Jr. and :"1. A. Naplin. 1981. Diel-depth distribution of summer ichthyoplankton in the
Middle Atlantic Bi;ght. Fish. Bull., U.S. 79: 705-726.
Lasker, R. 1975. Field criteria for the survival of anchovy larvae: the relation between inshore chlorophyll maximum layers and successful first feeding. Fish. Bull., U.S. 73: 453-462.
Loeb, V. J. 1979. Vertical distribution and development of larval fishes in the North Pacific central
gyre during summer. Fish. Bull. U.S. 77: 777-793.
---.
1980. Patterns of spatial and species abundance within the larval fish assemblage of the
North Pacific central gyre during late summer. Mar. BioI. 60: 189-200.
McGowan, M. F. 1985. Ichthyoplankton of the Flower Garden Banks, Northwest Gulf of Mexico.
Ph.D. Dissertation, University of Miami, Coral Gables, Florida. 376 pp.
Molinari, R. and K. Leaman. 1987. Surface currents in the Straits of Florida. Mariners Weather Log
31: 11-13.
Neilson, J. D. and R. I. Perry. 1990. Diel vertical migrations of juvenile fish: an obligate or facultative
precess? Adv. Mar. BioI. 26: 115-168.
Ropke, A. 1989. Small-scale vertical distribution of ichthyoplankton in the Celtic Sea in April 1986.
Meeresforsh.32: 192-203.
---,
W. Nellen and U. Piatkowski. 1993. A comparative study on the influence of the pycnocline
on the vertical distribution of fish larvae and cephalopod paralarvae in three ecologically different
areas of the Arabian Sea. Deep-Sea Res. 40: 801-819.
---,
U. Piatkowski and W. Welsch. 1990. A comparative study on the vertical distribution of fish
and cephalopod larvae in three hydrographically and ecologically different areas of the Arabian
Sea (preliminary results). ICES.C.M. I990/L: 89.
Sogard, S. M., D. E. Hoss and J. J. Govoni. 1987. Density and depth distribution of larval Gulf
menhaden, Brevoortla patronus, Atlantic croaker, Micropogonias undulatus, and spot, Leiostomus
xanthurus, in the nOl1hernGulf of Mexico. Fish. Bull., U.S. 85: 601-609.
Smith, W. G., J. D. Sibunka and A. Wells. 1978. Diel movements of larval yellowtail flounder
Limandafenuginea,
determined from discrete depth sampling. Fish. Bull., U.S. 76: 167-178.
Southward, A. J. and R. L. Barrett. 1983. Observations on the vertical distribution of zooplankton,
CHA ET AL.: FLORIDAKEYSFISH LARVAE
837
including postIarvai teleosts, off Plymouth in the presence of a thermocline
and a chlorophylldense layer. J. Plankton Res. 5: 599-618.
--and B. M. Bary. 1980. Observations
on the vertical distribution of eggs and larvae of mackerel
and other teleosts in the Celtic Sea and on the sampling performance
of different nets in relation
to stock evaluation. J. Mar. BioI. Assoc. U.K. 60: 295-311.
Woodhead, P. M. J. and A. D. Woodhead.
1955. Reactions of herring larvae to light: a mechanism
of vertical migration. Nature 176(4477): 349-350.
DATE ACCEPTED:
January
12, 1994.
(S.S.c.) Department of Oceanography, College of Natural Sciences, Chonnam National
University, Kwangju, 500-757, Korea; (M.F.M.) Division of Marine Biology and Fisheries, Rosenstiel
School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149-1098. PRESENT ADDRESS: 1423 Scenic Avenue, Berkeley, California 94708;
(W.J.R.) Southeast Fisheries Science Center, National Marine Fisheries Service, NOAA, 75 Virginia
Beach Dr., Miami, Florida 33149.
ADDRESSES:
838
BULLETIN OF MARINE SCIENCE, VOL 54, NO.3,
1994
Appendix I. List of the fish larvae which occurred at the eight night offshore stations
Order
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Beryciformes
Beryciformes
Beryciformes
Clupeiformes
Cyprinodontiformes
Dactylopteriformes
Gadiformes
Gadiformes
Gadiformes
Gadiformes
Gobiesociformes
Lophiiformes
Lophiiformes
Lophiiformes
Lophiiformes
Lophiiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
M yctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
M yctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Myctophiformes
Ophidiiformes
Ophidiiformes
Ophidiiformes
Ophidiiformes
Ophidiiformes
Ophidiiformes
Family
Aulopidae
Chlorophthalmidae
Notosudidae
Paralepididae
Paralepididae
Scopelarchidae
Synodontidae
Holocentridae
Melamphaidae
Melamphaidae
Clupeidae
Exocoetidae
Dactylopteridae
Bregmacerotidae
Bregmacerotidac
Bregmacerotidae
Macrouridae
Gobiesocidae
Antennariidae
Caulophrynidae
Ceratiidae
Ogcocephalidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Myctophidae
Carapidae
Carapidae
Carapidae
Ophidiidae
Ophidiidae
Ophidiidae
Genus and species
Aulopus
Chlorophthalmus
Scopelosaurus smithii
Sudis hyalina
Melamphaes
Etrumeus teres
Parexocoetus
Dactylopterus
Bregmaceros atlanticus
Bregmaceros cantori
Bregmaceros houdei
Cryptosaras couesi
Benthosema suborbitale
Centrobranchus nigroocellatus
Ceratoscopelus
Ceratoscopelus maderensis
Ceratoscopelus warmingi
Diaphus
Diaphus dumerilii
Diogenichthys atlanticus
Gonichthys cocco
Hygophum
Hygophum reinhardtii
Lampadena
Lampadena luminosa
Lampanyctus
Lampanyctus crocodilus
Lampanyctus cuprarius
Lampanyctus nobilis
Lepidophanes
Myctophum
Myctophum affine
Myctophum asperum
Myctophum nitidulum
Myctophum obtusirostre
Myctophum selenops
Notolychnus valdiviae
Notoscopelus resplendens
Taaningichthys
Carapus
Echiodon
Echiodon dawsoni
Lepophidium
Ophidion
839
CHA ET AL.: FLORIDA KEYS FISH LARVAE
Appendix I. Continued
Order
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Genus and species
Family
Acanthuridae
Acanthuridae
Acropomatidae
Apogonidae
Bramidae
Callionymidae
Carangidae
Carangidae
Carangidae
Carangidae
Chaetodontidae
Chiasmodontidae
Coryphaenidae
Coryphaenidae
Echeneididae
Epigonidae
Gempylidae
Gempylidae
Gempylidae
Gobiidae
Serranidae
Serranidae
Istiophoridae
Labridae
Labridae
Labridae
Labridae
Lutjanidae
Lutjanidae
Lutjanidae
Lutjanidae
Malacanthidae
Nomeidae
Nomeidae
Nomeidae
Percophidae
Percophidae
Pomacanthidae
Pomacentridae
Priacanthidae
Priacanthidae
Scaridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Acanthurus
Howella brodiei
Pterycombus brama
Caranx
Decapterus
Trachurus
Coryphaena
Coryphaena hippurus
Echeneis naucrates
Diplospinus multistriatus
Gempylus serpens
Nesiarchus nasutus
Pseudogramma gregoryi
Rypticus
Bodianus
Thalassoma bifasciatum
Xyrichthys
Lutjanus
Pristipomoides aquilonaris
Rhomboplites aurorubens
Cubiceps baxteri
Cubiceps pauciradiatus
Psenes
Bembrops anatirostris
Eupomacentrus
Pristigenys alta
Sparisoma
Acanthocybium solandri
Auxis
Euthynnus alletteratus
Katsuwonus pelamis
Thunnus
Thunnus thynnus
Anthias
Anthias nicholsi
Diplectrum
Epinephelus
Hemanthias aureorubens
Hemanthias leptus
Hemanthias vivanus
Liopropoma
840
BULLETIN OF MARINE SCIENCE, VOL. 54, NO.3, 1994
Appendix 1. Continued
Order
Perciformes
Perciformes
Perciformes
Perciformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Salmoniformes
Salmonifonnes
Salmoniformes
Salmoniformes
Salmoniformes
Scorpaeniformes
Scorpaenifonnes
Scorpaeniformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Stomiiformes
Tetraodontiformes
Tetraodontiformes
Zeiformes
Zeiformes
Unidentified
Family
Genus and species
Serranidae
Sphyraenidae
Trichiuridae
Uranoscopidae
Serranus
Sphyraena
Bothidae
Bothidae
Bothidae
Paralichthyidae
Paralichthyidae
Paralichthyidae
Cynoglossidae
Argentinidae
Argentinidae
Bathylagidae
Bathylagidae
Bathylagidae
Scorpaenidae
Triglidae
Triglidae
Chauliodontidae
Chauliodontidae
Gonostomatidae
Gonostomatidae
Gonostomatidae
Gonostomatidae
Malacosteidae
Melanostomiidae
Phosichthyidae
Phosichthyidae
Phosichthyidae
Phosichthyidae
Phosichthyidae
Phosichthyidae
Sternoptychidae
Stemoptychidae
Stemoptychidae
Stomiidae
Balistidae
Tetraodontidae
Caproidae
Caproidae
Bothus
Bothus occellatus
Monolene sessi/icauda
Citharichthys
Citharichthys gymnorhinus
Syacium
Symphurus
Argentina
Bathylagus
Bathylagus
longirostris
Peristedion
Chauliodus danae
Chauliodus sloani
Cyciothone
Gonostoma atlanticum
Gonostoma elongatum
Ichthyococcus ovatus
Pollichthys mauli
Vinciguerria
Vinciguerria attenuata
Vinciguerria nimbaria
Vinciguerria poweriae
Stemoptyx diaphana
Stemoptyx pseudobscura
Valenciennellus tripuctulatus
Stomias
Antigonia
Antigonia capros
841
CHA ET AL.: FLORIDA KEYS FISH LARVAE
Appendix 2. Mean abundance of the fish larvae at each stratum which occurred at the eight night
offshore stations
Stratum (Nil ,000 m')
Abundance
(N·IOO
Family
Myctophidae
Gonostomatidae
Bregmacerotidae
Scombridae
Serranidae
Para]ichthyidae
Phosichthyidae
Bothidae
Para]epididae
Gobiidae
Scorpaenidae
Synodontidae
Carangidae
Labridae
Sternoptychidae
Malacanthidae
Gempylidae
Coryphaenidae
Callionymidae
Argentinidae
Percophidae
Nomeidae
Antennariidae
Triglidae
Melamphaidae
Bathy]agidae
Chaliodontidae
Carapidae
Scaridae
Balistidae
Lutjanidae
Melanostomiidae
Apogonidae
Caproidae
Ophidiidae
Acropomatidae
Macrouridae
Priacanthidae
Pomacentridae
Tetraodontidae
Acanthuridae
lstiophoridae
Clupeidae
Ogcocephalidae
Bramidae
Cynoglossidae
Ceratiidae
Sphyraenidae
Stomiidae
Scopelarchidae
Dacty lopteridae
Chiasmodontidae
Malacosteidae
Holocentridae
Caulophrynidae
Uranoscopidae
4
154.8 239.5
112.9
99.6
6.3
76.1
125.8
7.0
]7.7
47.5
]9.0
7.1
]6.4
1.0
20.5
6.6
2.8
10.3
8.3
6.7
2.3
9.0
1.4
5.0
16.2
1.3
12.9
2.5
0.0
0.0
2.3
0.9
0.8
2.7
5.7
0.5
1.5
0.6
0.3
0.0
0.0
0.0
1.0
3.0
1.4
0.0
0.0
3.2
1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.8
0.0
1.4
0.5
1.4
0.5
0.8
0.5
2.4
0.0
0.8
0.0
1.6
0.0
1.5
0.0
0.0
0.0
0.0
0.6
0.4
0.7
0.7
0.0
0,9
0.4
1.0
0.0
0.0
0.9
0.0
0.6
0.8
0.0
0.3
0.5
0.4
0.0
0.7
0.0
0.3
0.0
0.0
0.0
0.7
0.0
0.0
0.0
0.0
0.6
0.0
0.5
0.0
0.0
0.0
0.0
107.0
27.4
30.2
0.0
12.8
27.8
5.1
1.9
6.9
3.8
4.4
7.2
0.0
0.6
0.0
2.4
1.9
0.0
0.0
0.0
0.0
1.1
3.2
0.0
1.5
0.0
0.0
0.4
0.6
0.0
0.0
1.1
0.0
0.6
0.0
0.5
0.4
0.0
0.0
0.5
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.5
99.9
4.6
19.2
0.0
1.8
5.3
7.9
1.0
1.2
0.5
3.8
4.2
0.0
0.0
0,0
2.8
1.0
1.1
0.4
2.1
1.5
0.0
0.0
0.0
1.3
1.]
0.0
1.6
0.0
0,0
0.0
0.0
0.0
0.8
0.5
0.0
0.0
0.0
0,0
0.4
0,0
0.0
0.0
0.0
0.0
0.0
0.4
0,0
0.4
0,0
0,0
0.0
0,0
0.0
0.5
0.0
m-2)
6
66.6
4.0
7.9
0.0
0.0
2.1
5.8
0.0
0.4
0.0
0.4
1.6
0.0
0.6
2.9
0.0
1.2
0,0
1.4
0.2
1.9
0.0
0.0
0.5
0.4
0.0
0.5
0.4
0.0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.4
0.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.2
0.0
0,6
0.0
0.0
0.0
0.0
27.2
4.8
0.9
0.0
0.0
1.0
4.9
0.0
0.5
2.5
1.2
0.0
0.0
0.0
3,8
0,0
0.0
0.0
1.9
0.6
2,5
0.0
0.0
0.6
0.0
1.8
0.0
0.8
0.0
0.6
0.6
0,0
0.0
0.0
0.0
0.0
].4
0.6
0.0
0,0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.5
0.0
0.0
0,0
0,0
0.0
0.0
8.5
3.9
0.3
0.0
0.0
0.7
1.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.4
0.0
0.3
0.0
0.5
3.6
0.5
0.0
0.0
0,0
0.0
0.5
2.9
0.0
0.7
0.0
0.0
0,0
0,0
0.0
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.8 1,763.0
642.7
0.0
351.8
0.0
0.0
332.0
]99.4
0.0
157.3
0.0
]05.6
0.0
0.0
74.8
0,0
55.1
0,0
54.4
52.7
0.0
0.0
48.3
0,0
43.8
0,0
41.3
22,5
0.0
20.9
0.0
]9.7
0.0
18.]
0.0
17.8
0.9
0.0
17.1
16.0
0.0
]2.7
0,0
0.0
11.4
0.0
10.8
10.]
0.0
8.5
0.0
8.4
0.0
8,1
0.0
7,8
0.0
6,3
0.0
6.1
0.0
6.0
0.0
6.0
0.0
%
38.9
14.2
7.8
7.3
4.4
3.5
2.3
1.6
1.2
1.2
1.2
1.1
1.0
0.9
0,5
0.5
0.4
0.4
0.4
0.4
0.4
0.3
0.3
0,2
0.2
0.2
0.2
0.2
0.2
0.]
0.1
0,1
0,]
0.0
5.3
0.1
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
5.1
4.9
4.6
4.5
3,8
3,8
3.4
2.5
2.3
2.3
2.1
2.0
2.0
1.8
1.8
1.7
1.7
1.4
1.4
1.2
1.2
1.2
0.1
0.]
0.1
0.]
0,1
0.1
0.]
0.]
0.1
0.1
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
842
BULLETIN OF MARINE SCIENCE. VOL. 54, NO.3, 1994
Appendix 2. Continued
Stratum (Nil ,000 m')
Abundance
(N·IOO
Family
Gobiesocidae
Notosudidae
Epigonidae
Chlorophthalmidae
Exocoetidae
Aulopididae
Chaetodontidae
Echeneididae
Trichiuridae
Lophiiformes
Perciformes
Pleuronectiforrnes
Myctophiforrnes
Unidentified
Total
%
No. of families
2
0.4
0.0
0.4
0.0
0.0
0.0
0.0
0.0
0.3
0.0
0.3
0.0
0.3
0.0
0.3
0.0
0.3
0.0
0.4
0.5
0.5
0.0
0.0
0.0
0.0
0.6
27.4
53.8
.571.2 599.5
31.5
33.1
40
38
3
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
19.9
270.2
14.9
27
6
4
O.it
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
11.4
176.3
9.7
26
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.4
0.0
0.8
0.0
3.2
104.4
5.8
22
7
0.0
0.0
0.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.8
61.9
0.0
0.0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.7
0.0
0.0
0,0
26.7
3.4
1.5
]4
2]
8
m-2)
0.0
1.1
1.1
0.0
0.0
0.9
0.0
0.8
0.0
0.8
0.0
0.8
0.0
0.8
0.0
0.8
0.0
0.8
0.0
3.3
0.0
3.1
0.0
1.9
0.0
1.6
0.5
297.3
3.1 4,533.2
0.2
100.0
2
65
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.1
0.1
0.0
0.0
6.6
100.0