A Comparative Study of Inshore and Offshore Copepod Populations at Barbados, West

Indies
Author(s): Finn Sander and Euna Moore
Source: Crustaceana, Vol. 35, No. 3 (Nov., 1978), pp. 225-240
Published by: Brill
Stable URL: https://www.jstor.org/stable/20103335
Accessed: 29-04-2020 20:12 UTC

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 Crustaceana 35 (3) 1978, L:. J. Brill, Leiden

A COMPARATIVE STUDY OF INSHORE AND OFFSHORE COPEPOD

POPULATIONS AT BARBADOS, WEST INDIES
BY

FINN SANDER
Bellairs Research Institute of McGill University, Barbados, W.I.

and

EUNA MOORE
Department of Biology, Cave Hill Campus, University of the West Indies, Barbados, W.I.

INTRODUCTION

Quantitative and qualitative studies of copepods are of fundamental importance

inasmuch as these organisms, by virtue of their ubiquitous distribution and relative
abundance everywhere, form a vital role in the economy of the sea. Though
individually small in size, they contribute, as a group, a considerable part of the
animal substance of the sea, and usually constitute about 70% of the Zooplankton
(Sverdrup, Johnson & Fleming, 1942). The tropical and subtropical Western
Atlantic and adjacent regions are not exceptions, and percentage contribution,
biomass and/or number of copepods are generally greatest in proximity of land
or island masses (Johnson, 1949; Grice, 1956; Hopkins, 1966; Moore, 1967;
Herman & Beers, 1969; Sander & Steven, 1973; Nutt & Yeaman, 1975). In this
connection the present study seeks to examine qualitative and quantitative aspects
relating to the importance of copepods to the inshore and offshore Zooplankton
populations on the leeward side of the island of Barbados, West Indies. The
results may be regarded as an indication of the extent biological and physical
factors may serve to influence and maintain relatively high populations of zoo
plankton in the downstream wake of a small isolated oceanic island.
The offshore data form an integral part of a multiparameter study of the biologi
cal production system of the tropical western Atlantic Ocean near Barbados and
complement published data on primary productivity (Steven, 1971; Steven &
Glombitza, 1972).
METHODS

Inshore Zooplankton was collected between 9.00 and 10.00 a.m. at approximately
weekly intervals from September 15, 1971, to September 7, 1973, off the Bellairs
Research Institute of McGill University on the west coast of the island. Thus, 94
samples were obtained from surface tows roughly following the 10 m contour
line for one kilometer. In addition, 50 concurrent vertical hauls were taken from
15

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226 FINN SANDER & EUNA MOORE

September 15, 1971, to September 7, 1972, from surface to approximately one

meter off the bottom at 25 m. These were discontinued after one year when it
became apparent that quantitatively and qualitatively the surface tows appeared to
sample adequately the thermally mixed shallow water inshore. A cursory examina
tion of the Zooplankton samples collected from eight inshore stations by Sander &
Steven (1973) and results from projects on the east coast (Robert Kidd, personal
communication) and south-west coast (Parti?, 1975) of the island indicate that
the above locality provides a valid estimate of the number of coastal-neritic plank
tonic copepod species off Barbados.
An offshore sampling station, situated at 13?15,N 59?43'W, in 460 meters of
water about 10 kilometers west of the island, was visited at about fortnightly inter
vals 52 times from 13/ix/67 to 3/xii/69. On each occasion one oblique tow to a
depth of 400 m and one surface tow of 15 minutes duration were made between
8.00 and 10.00 a.m. A 0.5 m diameter No. 6 net (0.239 mm mesh), fitted with a
calibrated flowmeter which provided an estimate of the total volume of water
filtered, was used for both sites to enable a valid quantitative comparison of the
data. Preliminary results showed that although various organisms such as small
pteropods and copepod nauplii were missed in a No. 6 net but retained in a No. 20
net, these contributed a relatively small part of the biomass of the total Zooplankton.
Each Zooplankton sample was then divided into two equal parts with a Folsom
splitter. One part was fixed with neutralized formaldehyde and stored for sub
sequent identification and enumeration and the other was placed in a pre-weighed
container, transferred to an oven, and dried to a constant weight at 60? C.
The coefficient of variation, determined from 10 subsamples of a single sample,
was 5.63% for copepods and 8.67% for other Zooplankton. Hence, an error of
less than 10% could be expected generally from the method of counting.

RESULTS AND DISCUSSION

The results and discussion are presented in two main sections dealing respectively
with ( 1 ) seasonal variation and ( 2 ) effects of the island mass, as they relate to the
pelagic copepods off Barbados. In the former, an attempt is made to interpret
the significance of peaks in copepod numbers by quantitative and qualitative break
down of the component species and by direct comparison to the results of previous
studies. In the latter, copepod density, composition, diversity and equitability,
inshore and offshore, are considered.

Seasonal Variations
An important consideration regarding treatment of the results concerns the
variation in copepod numbers found during the two periods of observations. It is
necessary to resolve whether such variations were determined mainly by local or
seasonal climatic factors. If the latter were of major significance one might expect
their effects to be manifested by regular periodicities in the alternations of peaks
and troughs. The results, presented graphically in figure la (offshore) and lb

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 COPEPOD POPULATIONS AT BARBADOS 227

15 - t-1-1-1-r
1971 C.FURCATUS 34?/.]('35%C.ARCUICORNIS 1972 50% C. FUR
_, A.SPINATA
10-= 7i?*48%
A.SPINATA

Fig. 1. Abundance of copepods in offs

broken lines represent surface and

(inshore), do not demonstrate suc

be treated as though varying ra
means, in turn, used for compara
Seasonal variation in the quantity
is well established (Menzel & Ry
1971), but supposed evidence of
proximity of Barbados has also be
the occurrence of minimum numb
maximum concentrations from N
based on monthly surface hauls t

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228 FINN SANDER & EUNA MOORE

the island. Lewis, Brundritt & Fish (1962) reported that the largest number of
copepods, based on monthly samples collected 8 km west of the island, were taken
during the winter and summer of 1959 and during the summer of I960. By con
trast, unpublished data taken from Sander & Steven (1973), did not demonstrate
seasonally in copepod abundance. The failure to demonstrate consistent patterns
of seasonally, or lack thereof, may in some instances reflect too short a collection
period or too few observations in a given time during which the infrequent
monitoring might not be sufficiently sensitive to record adequately the true annual
distribution of abundance of the various zooplankters. The present data on inshore
copepods in particular meet the demand for frequent observations. But considering
the surface collections alone, one might be tempted to accept two maxima per year,
one in the winter and one in the summer (with about a one-month lag in 1972-73),
more or less in conformity with a number of previous studies.
However, a qualitative analysis of the inshore data shows that the peaks result
from high incidence of one or two species at a time, rather than a concerted
response from most or many of the species making up the total copepod population
(fig. lb). This finding is reinforced by the data from vertical hauls. The occasional
abundance of one or two species of copepods could possibly be traced ultimately
to the occurrence of some highly "favourable" condition(s) in the environment.
Such occurrence must necessarily be short-lived if the essential characteristic of
stability in the environment is to obtain. If such a "favourable" circumstance is the
input of nutrient material then its effect would be most dramatically expressed
at the secondary production level by those organisms which, at the critical time,
were most physiologically and reproductively advantaged to exploit and translate
it (them) into an increase in numbers. So it is not necessary for all the component
species of a population to show increase in numbers in order to demonstrate seasonal
effect. The essential principle lies ultimately in the periodicity of the "favourable"
circumstance. Again, on the premise of the foregoing arguments it would be
expected that the bonus effects of any such "favourable" change would be expressed
in those species which extensive and intensive research has shown to be constant
contributors to the copepod population. In this locality Oithona plum?fera and
Calocalanus pavo dite two such species yet their population numbers show a much
more even temporal distribution. On the other hand some of the species such as
Acartia spinata, Clausocalanus arcuicornis and Oithona oculata which show these
occasional blooms are also at times completely absent from the sampled area, and
studies on their reproductive habits have, so far, not indicated that they are con
tinuous breeders. These facts would therefore speak to the hypothesis that the
seeming population bursts are governed by those factors specifically affecting the
biology of the species concerned ? in much the same way that some cladoceran
populations are affected by specific physiological factors ? rather than by large
scale seasonal variations whose effects should be reflected in a large number of
species.
Another possible explanation of this peculiar species-abundance phenomenon is

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 COPEPOD POPULATIONS AT BARBADOS 229

that lateral displacement may, at times, introduce into the region limited water
masses dominated by one or two species. This possibility gains plausibility in face
of the fact that some of the peaks (e.g. vertical, 29/ix/71, and surface, 28/ix/72)
appear and disappear between consecutive weekly observations. The lengths of the
maturation periods of the species involved, and the improbable depletion of these
copepods in one short week indicate that there are factors affecting observed
population size, other than simple on-the-spot production.
To sum up, graphical representations of copepod abundance reproduced from

1000

S ' 0 ' N D | J
1971

Fig. 2. Graphical comparison of the data on temporal distribution of copepods as determined by:
A, Fish (1962); B, Lewis, Brundritt & Fish (1962); C, Sander & Moore (offshore data); D, Sander
& Steven (1973); E, Sander & Moore (inshore data).

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230 FINN SANDER & EUNA MOORE

Fish (1962), Lewis, Brundritt & Fish (1962) and Sander & Steven (1973) in
conjunction with present inshore and offshore surface data (fig. 2) constitute a
visual comment on the topic of seasonality and serve to demonstrate that in the
context of a longer time-span these seasonal maxima and minima, which some
times seem quite valid on short-term data, really oscillate around annual means.

The Island Mass Effect

Numerical estimates of copepods inshore and offshore are listed in table I
together with the respective average percentage contributions to the total Zooplank
ton. The most striking feature is the high percentage contributions by copepods
inshore amounting to almost 90% near the surface. The scatter diagram in figure 3
depicts a strong positive correlation between surface copepod numbers and zoo
plankton biomass, expressed as dry weight, which implies that copepods inshore
were chief contributors not only in numbers but also in biomass. Although
absolute numbers and percentage contributions of copepods were higher inshore
than offshore, the data on total Zooplankton do not suggest a decreasing gradient
inshore to offshore indicative of an island mass effect. This can probably be
explained by the fact that since the offshore and inshore collections were not

Table I
Comparative inshore and offshore Zooplankton data expressed as numbers per
cubic meter
INSHORE OFFSHORE
Total Copepods % Copepods Zooplan
Zooplankton Total
Surface 327 292 89.30% 405 250 61.73%
Vertical 373 292 78.28% 185 121 65.41%

concurrent, pronounced annual variations in average Zooplankton numbers might

prevent a manifestation of an island mass effect based on the present data alone.
Such differences, in fact, represent another notable feature of the results. Average
total Zooplankton counts for both oblique and surface hauls offshore were high
during most of 1968 compared to the rest of the period studied. Mean values for
oblique and surface hauls respectively were 268/m3 and 669/m3 in 1968 versus
102/m3 and 226/m3 in 1969. The yearly differences were apparent for copepods
also and are clearly demonstrated in figure la. Although no explanation of this
can be offered at present it is worth noting that the period of high copepod and
total Zooplankton counts corresponds with the time when marked enrichment of
phosphate and nitrate was observed on four occasions at the base of the euphotic
zone (unpublished data). Such nutrient enrichment would be expected to increase
the level of primary production which in turn would sustain greater quantities of
Zooplankton.
An indication of the island mass effect becomes apparent when the present

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 COPEPOD POPULATIONS AT BARBADOS 231

100.00

10.00-H

g
LU
.? *. *
. .*
>
ce
?
z
o 1.00
Z
<
CL
O
o
N
O)
E
O 10H
O

.01-H1
i?i?i?i?i?i?i?i?i?i?i?i?r
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14
COPEPODS/M3*102
Fig. 3. The relationship between inshore copepod numbers and

data are compared to the literature. Inshore average c

ticular were high compared to the average of 83/m3
Sea stations occupied by Calef & Grice (1967) and t
Grice & Hart (1962) for the northwestern Sargasso
the comparative abundance of copepods in close proxim
from generally increased availability of food close
postulated that internal waves, which were shown to
island shelf in conjunction with high nutrient concent
of the shallow water thermocline, may act as a princ
which inshore island waters are enriched. This sou
operates to support increased plankton populations and

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 232 FINN SANDER & EUNA MOORE

ever, the ultimate factor which serves to maintain numerically high average popu
lations of Zooplankton and phytoplankton must necessarily be the same, namely a
mechanism for retaining the plankton in close proximity of the island. This is
important not only for meroplanktonic organisms, such as pelagic larvae of littoral
benthic invertebrates which seek to settle over the narrow shelf of the island, but
also for neritic holoplanktonic types, including some species of copepods. Emery
(1972) has proposed such a mechanism in the form of an eddy system in the
downstream wake of Barbados inhibiting a rapid dispersal of the neritic plankton
in a westerly direction by the prevailing equatorial current. Similarly, Boden
(1953) has suggested that the circulation around Bermuda also produces a system
of retention for that island, and McGary (1955) pointed out that Zooplankton
maxima in Hawaiian inshore waters were associated with cyclonic eddies in the lee
of these islands.

Total numbers of 57 and 134 species of copepods were identified for the inshore
and offshore sites respectively and are listed in table II. All the species have been
reported previously in the North Sargasso Sea (Grice & Hart, 1962), the Caribbean
Sea (Owre & Foyo, 1964, 1972), the Florida Straits (Owre & Foyo, 1967), or the
area of the equatorial Atlantic influenced by the Amazon (Calef & Grice, 1967).
None are therefore new to science. The lesser numbers of species reported for
offshore by Wickstead (1956), Fish (1962), and Lewis & Fish (1969) can
probably be explained by the fact that Wickstead obtained all his samples from the
months of March and April only, and that the coarse nets employed by Fish
( stramin and No. 1) and Lewis & Fish (stramin) no doubt allowed numerous
small species to escape.

Table II
Copepod species collected in inshore (I) and offshore (O) at Barbados
CALANOIDS
lO Acartia negligens Dana, 1852
IO Acartia spinata Esterly, 1911
I Acartia sp.
IO Acrocalanus longicornis Giesbrecht, 1888
O Aegisthus mucronatus Giesbrecht, 1891
O Arietellus seto sus Giesbrecht, 1892
O Augaptilus longicaudatus (Claus, 1863)
O Bathycalanus richardi G. O. Sars, 1905
O Bradycalanus typicus A. Scott, 1909
IO Calanus tenuicornis Dana, 1852
IO Calanopia americana F. Dahl, 1894
IO Calocalanus pavo (Dana, 1852)
IO Candada pachydactyla (Dana, 1852)
O Candada paenelongimana Fleminger &
Bowman, 1956
IO Centropages fur cat us (Dana, 1852)
O Centropages violac? us (Claus, 1863)
IO Claus o calan us arcuicornis (Dana, 1852)
IO Clausocalanus fur cat us (Brady, 1883)
O Euaetideus gi es bree ht i (Cleve, 1904)
IO Eucalanus attenuatus (Dana, 1852)
IO Eucalanus crassus Giesbrecht, 1888
O Eucalanus elongatus (Dana, 1852)
O Eucalanus monachus (Giesbrecht, 1888)
O Eucalanus ?nucronatus Giesbrecht, 1888
O Eucalanus pileatus Giesbrecht, 1888
O Eucalanus subcrassus Giesbrecht, 1888
O Euchaeta malayensis (Sewell, 1929)
IO Euchaeta marina (Prestandrea, 1833)
O Euaugaptilus hecticus (Giesbrecht, 1889)
O Euchaeta wit hi (Sewell, 1947)
O Euehirella amoena Giesbrecht, 1888
O Eue h ir ella pulchra (Lubbock, 1856)
O Euchirella venusta Giesbrecht, 1888

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 COPEPOD POPULATIONS AT BARBADOS 2 33
O Euaugaptilus bullifer (Giesbrecht, 1889)
O Euaugaptilus hectic us (Giesbrecht, 1889)
0 Euaugaptilus magnus (Wolfenden, 1904)
O Ga?tan us miles Giesbrecht, 1888
O Ga?tan us pileatus Farran, 1903
O Haloptilus acutifrons (Giesbrecht, 1892)
O Haloptilus longicornis (Claus, 1863)
O Haloptilus ornatus (Giesbrecht, 1892)
O Haloptilus spiniceps (Giesbrecht, 1892)
O Heterorhabdus papilliger (Claus, 1863)
O Heterorhabdus s p inifron s (Claus, 1863)
O H et er o Stylit es longicornis (Giesbrecht, 1889)
O Labidocera acutifrons (Dana, 1852)
O Labidocera aestiva Wheeler, 1901
IO Labidocera nerii (Kroyer, 1849)
O Lucicutia flavicornis (Claus, 1863)
O Lucicutia magna Wolf enden, 1903
IO Mecynocera clausii I. C. Thompson, 1888
O Mecynocera sp.
O Megacalanus sp.
IO Metridia princeps Giesbrecht, 1889
O Metridia venusta Giesbrecht, 1889
O Monacilla t y pica G. O. Sars, 1905
O Mormonilla minor Giesbrecht, 1891
O Nannocalanus minor (Claus, 1863)
?O Neo calan us gracilis (Dana, 1852)
IO Neocalanus robustior (Giesbrecht, 1888)
IO Paracalanus aculeatus Giesbrecht, 1888
IO Paracalanus par vus Claus, 1863
O Paracandacia simplex (Giesbrecht, 1889)
O Phaenna spinifera Claus, 1863
1 Phyllopsus bidentatus G. Brady, 1883
O Pleuromamma abdominalis (Lubbock, 1856)
O Pleuromamma gracilis (Claus, 1863)
O Pleuromamma piseki Farran, 1929
O Pleuromamma quadrangulata (F. Dahl,
1893)
O Pleuromamma xiphias (Giesbrecht, 1889)
O Pont ella atl?ntica (Milne-Edwards, 1840)
IO Pontellina plum?t a (Dana, 1852)
O Pontellopsis perspicax (Dana, 1852)
O Pontellopsis regalis (Dana, 1852)
O Pseudochirella obesa G. O. Sars, 1920
IO Rhin calan us cornutus (Dana, 1852)
O Rhincalanus nasutus Giesbrecht, 1888
O Scolecithricella tenuiserrata (Giesbrecht,
1892)
O Scolecithricella vittata (Giesbrecht, 1892)
O Scolecithrix bradyi Giesbrecht, 1888
IO Scolecithrix danae (Lubbock, 1856)
O Scolecithrix minor (Brady, 1883)
O Scottocalanus per secan s (Giesbrecht, 1895)
IO Temor a stylifera (Dana, 1852)
IO Temor a turbinata (Dana, 1852)
O Temoropia mayumbaensis T. Scott, 1894
O Undeuchaeta minor Giesbrecht, 1888
IO Undinula vulgar is (Dana, 1852)
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0 Valdiviella brevicornis Sars, 1905
IO Valdiviella in s ignis Farran, 1908
CYCLOPOIDS
O Caligulus sp.
IO Clytemnestra scutellata Dana, 1848
O Conaea gracilis (Dana, 1853)
O Corycaeus amazonicus Dahl, 1894
O Corycaeus americanus M. S. Wilson, 1949
IO Corycaeus cat us F. Dahl, 1894
O Corycaeus clausi F. Dahl, 1894
IO Corycaeus flace us Giesbrecht, 1891
IO Corycaeus latus Dana, 1852
O Corycaeus laut us Dana, 1852
IO Corycaeus limbatus Brady, 1883
IO Corycaeus s pecios us Dana, 1852
O Corycaeus t y pic us (Kroyer, 1849)
IO Cop ilia mirabilis Dana, 1852
O Cop Hi a mediterr?nea (Claus, 1863)
IO Copilia quadrata Dana, 1852
O Copilia vitrea (Haeckel, 1864)
IO Fart'anula gracilis (Dana, 1853)
I Farranula car ?nata (Giesbrecht, 1891)
O Lepeoptheirus sp.
I Lubbockia aculeatus (Giesbrecht, 1891)
IO Lubbockia squillimana Claus, 1863
O Oithona fr?gida Giesbrecht, 1902
O Oithona helgolandica Claus, 1863
IO Oithona nana Giesbrecht, 1892
IO Oithona oculata Farran, 1913
?O Oithona plum?fera W. Baird, 1843
I Oithona robusta Giesbrecht, 1892
IO Oithona setigera (Dana, 1852)
IO On cae a mediterr?nea Claus, 1863
IO Oncaea not opus Giesbrecht, 1891
IO Oncaea venusta Philippi, 1843
O Pachos punctatum (Claus, 1863)
O Ratania flava Giesbrecht, 1892
1 Sapphirella tr?pica Wolf enden, 1906
O Sapphirina auronitens Claus, 1863
O Sapphirina intestinata Giesbrecht, 1891
IO Sapphirina nigromaculata Claus, 1863
IO Sapphirina opalinata Dana, 1852
IO Sapphirina ovatolanceolata Dana, 1852
O Sapphirina stellata Giesbrecht, 1891
O Unidentified parasitic spp. (2)
O Vettoria granulosa (Giesbrecht, 1891)
HARPACTICOIDS
O Eut er pina acutifrons (Dana, 1847)
O H ar pact i cus gurneyi Jakubisiak, 1933
IO Macro s et ella gracilis (Dana, 1848)
IO Microsetella norvegica Boeck, 1864
IO Mi ero s et ella rosea (Dana, 1848)
IO Mirada efferata Dana, 1852
O Oculosetella gracilis (Dana, 1852)
 234 FINN SANDER & EUNA MOORE

The results conform to standard concepts of diversity in tropical marine environ

ments. In the same way it is generally accepted that in northern (and harsher)
regions there are fewer Zooplankton species with greater numbers of individuals
than in more tropical areas (Sutcliffe, I960), tropical inshore waters are considered
characterized by comparatively high concentrations of plankton composed of few
species, whereas oceanic waters support a more sparse but widely diversified zoo
plankton population (Deevey, 1964; Bj?rnberg, 1971). The literature bears this
out. For example, Grice & Hart (1962) identified nearly one hundred species of
copepods in the Sargasso Sea, and Herman & Beers (1969), in comparing Bermuda
inshore Zooplankton (including copepods) with offshore sites, concluded that the
inshore regions have relatively larger numbers of individuals per given volume
but a lower diversity of forms. The same pattern is repeated at Puerto Rico (Coker
& Gonzalez, I960), Jamaica (Moore, 1967), and off the south east coast of the
United States (Bowman, 1971).
Implicit in the popular explanation of why tropical oceans support a greater
diversity of Zooplankton than inshore waters is the same idea most frequently
offered to account for the differences in diversity between the plankton of tropical
and temperate or polar oceans ? namely, that the greater environmental stability
of tropical oceans has allowed the Zooplankton there to evolve and diversify fur
ther than in areas with less stable conditions, such as temperate or polar seas which
experience pronounced periodic climatic fluctuations (Fischer, I960; Dunbar,
I960). In the same context, tropical coastal waters may also be considered regions
of oscillation of environmental factors, including, but not necessarily limited to,
salinity and occasionally temperature. Bowman (1971) has applied Sanders' (1968)
stability-time hypothesis, proposed to account for variations in coastal and oceanic
benthic diversity, to the pelagic realm, and suggested that coastal and estuarine
copepods belong to physically controlled communities, "where physical conditions
fluctuate widely and the animals are exposed to severe physiological stress. Adap
tations are primarily to the physical environment. The species forming such com
munities are few in number and eurytopic". In contrast, oceanic copepods are part
of biologically controlled communities, "where physical conditions are stable for
long periods of time. Stress is from biological rather than physical forces. Such
communities are composed of many species, which are stenotopic". However,
neither type of community exists in pure form, and, although the inverse relation
ship between absolute and species numbers of copepods inshore and offshore
at Barbados lends support to the generally accepted concepts of diversity in tropical
oceanic and neritic Zooplankton, this appears to be true of the present investigation
which does not provide the best example of low diversity in tropical inshore waters.
Differences between temperature and salinity (excepting rare instances of flood
rains) inshore and offshore are minimal (Sander & Steven, 1973), and the narrow
island shelf necessarily results in considerable intrusion of oceanic water. A better
example is given by Bacon (1971) who collected only 12 species of copepods in
estuaries of the Caroni mangrove swamps, Trinidad, subject to marked variation

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 COPEPOD POPULATIONS AT BARBADOS 235

30

20H

ioH
LU
?
LU
a.

u.
o B
CL
Ui
CD 30H
S
D
Z

20

ioH

0 10
10 20
20 30 40
% SPECIES MAKING UP
Fig. 4. Relationship between equitability and species diversity offshore (A) and ins
Barbados.

in salinity. Similarly, Grice (1956) identified only 20 species (and two additional
genera) in his investigation of the copepods of the Alligator Harbour estuary which
is subject to pronounced variation in temperature and salinity.
Another expression of diversity is the degree to which the species of a population
are equally abundant. This can be calculated as an equitability component (Lloyd
& Ghelardi, 1964) but a rough estimate of equitability is the percent of all cope

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 236 FINN SANDER & EUNA MOORE

pod species in a sample that together comprises half the total number of copepods
in that sample (Bowman, 1971). In figures 4a and 4b this measurement is plotted
against the number of species for offshore and inshore surface samples respectively.
It can be seen that in the offshore samples 10-30 percent of the species generally
constitute half of a given sample compared to 5-20 percent in the inshore ones.
The copepods in the offshore collections were thus more evenly divided among the
component species than the inshore specimens, indicating that the offshore locality
is more oceanic and influenced less by the island mass than the near-shore site. The
results in table III also suggest a greater degree of equitability offshore than inshore
where the bulk of the copepod population was comprised of relatively few species.
The latter are typically grazers of small size which conform to Margalef's (1971)
conception of copepods which are common in places of pulsating productivity and
are of wide geographical distribution, including coastal waters of the Caribbean

Table III

Percentage contributions to total copepods by the twelve most common species

inshore and offshore at Barbados
Surface Vertical
Copepod species Max. Mean Max. Mean
INSHORE
Acartia s pin at a 93.11 18.33 77.54 9.90
Clausocalanus furcatus 62.61 16.54 43.23 17.17
Calo calan us pavo 66.78 8.11 23.18 7.11
Clausocalanus arcuicornis 39.47 7.87 31.62 8.58
Oithona plum?fera 69.42 6.91 34.84 6.36
On cae a mediterr?nea 31.19 6.87 22.38 5.37
Far r?nula gracilis 19.25 5.45 13.48 4.11
Macro s et ella gracilis 73.27 5.12 29.95 3.65
Und i nula vulgar i s 39.35 4.10 10.50 2.76
Oithona oculata 94.68 3.63 66.67 10.37
Oithona setigera 41.41 2.44 65.17 11.32
Temor a tur bin at a 66.90 1.54 70.73 4.09
Total 86.91 Total 90.79
OFFSHORE
Far r?nula gracilis 62.93 21.32 38.71 9.85
Oncaea venusta 76.06 15.79 31.65 8.81
Undinula vulgar i s 40.11 15.25 26.22 11.83
Oncaea mediterr?nea 34.00 5.88 24.83 10.15
Euchaeta marina 20.32 5.59 18.17 10.15
Calo calan us pavo 12.64 4.05 10.00 2.58
Corycaeus s pecios us 10.34 2.31 11.98 2.61
Oithona setigera 12.11 2.21 13.20 4.20
Oithona plum?fera 9.09 2.03 14.28 3.93
Paracalanus aculeatus 10.70 2.02 22.91 3.47
Oithona nana 7.04 1.99 18.92 7.09
Macro s et ella gracilis 11.11 1.98 14.17 3.85
Total 78.23 Total 77.64

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 COPEPOD POPULATIONS AT BARBADOS 237

area. Margalef's examples of copepods belonging to this group, and listed in

table II, are Clausocalanus arcuicornis and species of the genera Acartia and
Oithona. Going offshore to areas of lower productivity, there is an increase of large
species often belonging to higher trophic levels (Margalef, 1971). The species of
comparatively large individual size, and which were more abundant at the oceanic
station than inshore, were Undinula vulgaris, Euchaeta marina, Candada pachy
dactyla and Scolecithrix danae. It is likely that by virtue of their size these species
contributed more to the biomass than their numerical contribution would indicate.
Farranula gracilis, U. vulgaris, E. marina, Paracalanus aculeatus, Corycaeus
speciosus, Macro s et ella gracilis were amongst the chief contributors offshore, and
are well-known oceanic copepods which Bj?rnberg (1971) cited in her group of
"most frequent and/or numerous" oceanic copepods in the Caribbean area. The
authors recognize that opinions sometimes differ in the literature as to the type of
certain copepods such as widely ranging U. vulgaris considered by Johnson (1949)
and Jones (1962) as a neritic form. Inshore, the island has a very narrow coral
ledge and undoubtedly the occurrence of some of the above-mentioned oceanic
species as chief numerical contributors in coastal waters of the island reflects the
inevitable intrusion of oceanic water. In fact, Bj?rnberg reported that species such
as U. vulgaris and those of the genera Farranula and Oncaea may be numerous in
the Caribbean islands with narrow coral ledges, inasmuch as over the narrow shelves
of these islands "the coastal-neritic, the neritic-slope and the oceanic shelf faci?s
of Fleminger (1959) mix and form one faci?s with the characteristics of all three".
Bj?rnberg also mentioned Clausocalanus furcatus and Calocalanus pavo in this
connection, and it is interesting, and a little puzzling, to note that the former was
not prevalent at the deep water station as well. At present no reasonable explanation
can be offered for this anomaly, unless it be the long-term quantitative variations
of copepods, previously noted, accompanied by significant qualitative changes.
C. furcatus, particularly, is a frequent and abundant contributor close to shore at
Barbados. It occurs world-wide in warm water (Gonzalez & Bowman, 1965), and
has been reported to occur in abundance in the tropical and subtropical Western
Atlantic and adjacent regions (Moore, 1949; Grice & Hart, 1962; Calef & Grice,
1967; Bowman, 1971; Deevey, 1971). Specifically, Grice & Hart described C.
furcatus as the most abundant copepod in their Gulf Stream and Sargasso Sea
samples, and Calef & Grice reported that it was one of the two most numerous
species in the area of the equatorial Atlantic influenced by the Amazon ? which
includes Barbados (Steven & Brooks, 1972). Significantly, the other prominent
species was Oithona plum?fera, also an important contributor inshore and offshore
during the present investigation.
In contrast to C. furcatus, which was a consistent contributor to the shallow water
Zooplankton, the numerical contribution of Acartia spinata inshore was highly
irregular, and the elevated average of the latter resulted from high percentage
contributions when total copepod counts were high also. As previously mentioned,
these peaks may be caused by lateral displacement, or they may represent direct

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 238 FINN SANDER & EUNA MOORE

responses to most favourable feeding conditions which were not a consistent feature
of the inshore waters at Barbados. There is evidence that Acartia may be a relatively
inefficient feeder compared to more oceanic species (Conover, 1956), which
could suggest that A. spinata may not prosper under true oceanic conditions where
standing crop of phytoplankton may be considerably and consistently lower than
near shore. A. spinata has also been described as abundant in proximity of land
or island masses in other parts of the Caribbean and adjacent areas where phyto
plankton concentrations might be expected to be high. Herman & Beers (1969)
recorded its distinct presence over the North Lagoon off Bermuda, and Emery
(1968) observed swarms of this species at Alligator Reef, Florida. Also "Smith
sonian-Bredin" Expeditions have collected it in Fort de France Harbour, Mar
tinique, near Pigeon Point, Tobago, and off Cocoa Point, Barbuda. It is abundant
in Lameshur Bay, St. John, and occurred both in the bays and over the shelf at
Puerto Rico (Gonzalez & Bowman, 1971).
In conclusion, it is apparent that, although the island mass affects the copepod
density, diversity and equitability in the downstream wake of Barbados, the species
composition of copepods reflects the influence of the surrounding oceanic water
masses. It is suggested here that a causal relationship exists between the successful
invasion of the inshore waters by various oceanic copepods and the fact that salinity
and temperature over the narrow island shelf do not differ markedly from offshore
conditions (Sander & Steven, 1973). It is not necessary, therefore, for such organ
isms to exhibit a wide tolerance to changes in these physiologically important
environmental parameters. Further studies are necessary to determine with certainty
whether the general increase in proximity of the island mass is due to reproductive
recruitment rather than to reduced mortality, or to an accumulating phenomenon
resulting from persistent advection of offshore stock. It may, of course, be a com
bination of all three, since none of them are mutually exclusive. Implicit in the
above is the probability that a system of retention operates to maintain the abun
dance of copepods, which may vary from year to year.

ACKNOWLEDGEMENTS

Many thanks are due to Mr. Keith Alleyne for his technical assistance, and to
Dr. Abraham Fleminger and Dr. Harry Yeatman for identification of copepods.
We are also indebted to Dr. Carol Lalli, professor Delphine McLellan and
Dr. Harry Yeatman for reading the original manuscript and offering constructive
criticism. The work was supported by a grant in aid of research from the National
Research Council of Canada.

R?SUM?
Ce travail est une ?tude dans le temps de l'abondance et des changements qualitatifs des popu
lations de Cop?podes dans les eaux c?ti?res et dans les eaux du large, ? la Barbade, Antilles.
La question des variations saisonni?res, exprim?es pour l'ensemble ou pour une population indivi
duelle, est consid?r?e dans le contexte d'un long intervalle de temps. Les r?sultats ont montr? qu'il

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 COPEPOD POPULATIONS AT BARBADOS 239

existait des variations significatives non-saisonni?res dans les quantit?s de Cop?podes, qui varient
de fa?on al?atoire autour des moyennes annuelles. Ce sont 57 esp?ces pr?s de la c?te et 134 esp?ces
au large qui ont ?t? respectivement identifi?es pendant les p?riodes d'?tude. L'effet de masse insulaire
est quelque peu amorti dans la mesure o? le gradient eaux c?ti?res/eaux du large est mod?r?. Cepen
dant une relation inverse entre nombre absolu et nombre d'esp?ces de Cop?podes, pr?s des c?tes
d'une part, au large d'autre part, appuie le concept d'une diversit? croissante avec l'?loignement
des c?tes. Un degr? d'?quitabilit? plus bas pour la population de Cop?podes c?ti?re que pour la
population du large a ?t? observ? et la composition des groupes respectifs a ?t? compar?e. Une
intrusion d'eau oc?anique au voisinage des c?tes appara?t dans la forte contribution num?rique des
esp?ces oc?aniques ? la population c?ti?re.

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Received for publication 9 December 1975.

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