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Aquatic Ecology 36: 411424, 2002.

2002 Kluwer Academic Publishers. Printed in the Netherlands.

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Patterns of prey selectivity in the cyclopoid copepod Mesocyclops


thermocyclopoides
T. Ramakrishna Rao and Ram Kumar
Department of Zoology, University of Delhi, Delhi 110 007, India (E-mail: raotr@bol.net.in)
Accepted 25 May 2001

Key words: Copepoda, food, Mesocyclops thermocyclopoides, prey handling time, prey selection, predation, prey
switching

Abstract
In the shallow and eutrophic subtropical aquatic ecosystems, which it generally inhabits, the omnivorous copepod
Mesocyclops thermocyclopoides encounters a wide variety of animal prey types including ciliates, rotifers, and
cladocerans. We studied prey selectivity in laboratory-reared adult females of this species given a choice of (i) prey
types belonging to different taxa (ciliates, rotifers, cladocerans, and cyclopoid nauplii), and (ii) different prey
species within a taxonomic group differing in body size, morphology or behaviour. We also tested the effect of
different proportions of prey species on its selectivity. Prey type proportion had no significant effect on selectivity
of the copepod, nor was there any evidence of switching based on the relative abundance of prey. Among the ciliate
prey species tested, the largest species, Stylonychia mytilus was positively selected regardless of its relative abundance, while the smallest, S. notophora was selected only when its density was higher. Offered a choice of three
species of a brachionid rotifer differing in size, the copepod selected the largest of them, Brachionus calyciflorus,
and avoided the smallest B. angularis. The evasive rotifer Hexarthra mira was also avoided. When prey choice
included three cladoceran species Daphnia similoides, Moina macrocopa and Ceriodaphnia cornuta, the copepod
selected the intermediate-sized M. macrocopa regardless of the abundance of the other two species. Although it
fed on Mesocyclops nauplii when there was no choice, M. thermocyclopoides avoided them when alternative food
was available. In a multispecies prey choice test, the copepod selected predominantly the rotifer B. calyciflorus and
the cladoceran M. macrocopa. We suggest that the prey selectivity patterns shown by M. thermocyclopoides are
adaptive in that they lead to ingestion of the most profitable prey.

Introduction
In many eutrophic water bodies, a wide spectrum of
potential food items including algae, ciliates, rotifers,
cladocerans, and copepod nauplii is available to cyclopoid copepods. Yet, many cyclopoids are known
to feed selectively (Greene, 1983; Williamson, 1986;
Krylov, 1988; Janicki & DeCosta, 1990), exerting a
significant influence on the structure and dynamics of
the plankton community (Karabin, 1978; Kerfoot &
Sih, 1987; Matsumura-Tundisi et al., 1990; Irvine &
Waya, 1993). Prey selectivity in cyclopoids may result
from differential vulnerability of prey species (passive
selection) or from a choice exercised by the predator
of accepting or rejecting a prey item (active selection)

(Stemberger, 1985; Roche, 1990). Since prey vulnerability is a product of preys encounter rates with,
and ease of capture by, the predator (Pastorok, 1981),
certain morphological and behavioural attributes of
different prey species (which influence capturability)
(Li & Li, 1979; Williamson, 1980, 1983a; Stemberger,
1985; DeMott, 1995; Wickham, 1995a,b), and their
relative proportions (which affect encounter rates) in
the medium, may be able to explain observed selectivity patterns in passive prey selection. Active prey
selection has also been demonstrated in some calanoid
and cyclopoid species (Stemberger, 1985; Kerfoot &
Kirk, 1991; Hartmann et al., 1993; DeMott; 1995),
but the mechanisms leading to active selection are not
necessarily the same. In the calanoid Acartia tonsa, for

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instance, food selectivity is a consequence of two distinctly different feeding mechanisms, ambush feeding
for motile animal prey such as ciliates, and suspension feeding for small immotile food particles such as
diatoms (Kirboe et al., 1996). This suggests that in
omnivorous copepods that regularly include algae in
their diet (Adrian, 1991; Santer, 1993; Santer & Van
den Bosch, 1994; Hopp et al., 1997), their patterns
of animal prey selection might also be related to the
extent of herbivory in them (Adrian & Frost, 1993;
Kumar & Rao, 1999a).
The ability of a predator to switch from one prey
species to another depending on their relative concentrations is an adaptive behaviour that not only allows
the predator to maximize its net energy intake (Landry,
1981) but may also have a stabilizing effect on the prey
population (Oaten & Murdoch, 1975). Prey switching
has been recorded in some calanoid copepods (Landry,
1981; Kirboe et al., 1996) and is implicated in species
that show a Type III functional response (Akre &
Johnson, 1979).
Mesocyclops thermocyclopoides, a common cyclopoid copepod in local ponds and shallow lakes, is
predominantly carnivorous, but with an ability to utilize plant food also (Kumar & Rao, 1999a,b). Our
earlier laboratory studies on food type effects on the
post- embryonic development and reproductive output of this species revealed prey type-related trends
in its survival, longevity and fecundity (Kumar &
Rao, 1998, 1999b). These trends led us to hypothesize
that Mesocyclops thermocyclopoides feeds selectively
based on prey attributes such as body size, shape,
motility, type of movements, and perhaps taste. Further, we asked if selectivity in this cyclopoid is prey
frequency-dependent.
The objective of our present study was to understand the prey selection patterns in the adult female
of M. thermocyclopoides, given a choice of different
prey types including ciliates, rotifers, cladocerans and
copepod nauplii, differing from each other in one or
more of the morphological and behavioural attributes.
Our experiments were also designed to test the effect
of prey proportions in the medium on the selectivity
patterns of the copepod.

Materials and methods


All experiments were conducted with laboratoryreared adult (age: 68 d; size:1050 53.9 m) females of M. thermocyclopoides. The methods used

for culturing the cyclopoid and the prey species under


laboratory conditions were described earlier (Kumar
& Rao, 1998, 1999b). In addition, the rotifer species
Filinia longiseta and Hexarthra mira were cultured
using Chlorella as food, and the predatory rotifer Asplanchna intermedia using Brachionus angularis and
B. calyciflorus as food. Important parameters of the
test prey species are given in Table 1. Body sizes
(length and width) of the organisms were measured
under a microscope using an ocular micrometer. However, the dry biomass values for the prey species used
in this study were taken from published sources (Dumont et al., 1975; Mookerji, 1992; Foissner & Berger,
1996).
A series of short-duration (3 to 12 hours) experiments were conducted to investigate prey selectivity in
M. thermocyclopoides given a choice of (i) different
species within a taxonomic group, differing in body
size, morphology or behaviour, and (ii) prey types
belonging to different higher taxa (ciliates, rotifers,
cladocerans) (Tables 2 and 3). While designing the
selectivity tests, we sought to answer two important
questions: does the preference of the predator change
as the relative proportions of the prey types change
in the medium, and does such a change reflect preyswitching behaviour? To answer these questions, in
nearly all the tests, prey types were offered in different
proportions, keeping the total prey density constant.
The experimental protocols for different selectivity
tests conducted in this study are given in Table 2 and
the prey species used in multispecies choice test in
Table 3.
Further, in two-prey tests, we applied Landrys
(1981) formula to check for prey-switching behaviour: if N1 and N2 represent the densities of two prey
species and P1 and P2 the numbers respectively of the
two species consumed in a given time period, then for
non-switching predator
P1 /P2 = C(N1 /N2 ),
where C is a proportionality constant reflecting the
innate preference of the predator for one or the other
prey species. Increasing value of C with increasing value of N1 /N2 implies prey switching (Landry,
1981).
Adult copepods used in selectivity tests had sufficient exposure to all the test prey species prior to testing, since they were cultured using as food different
combinations of the same prey species. Prey species
were individually counted and introduced at set concentrations and proportions into 150 ml beakers, each

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Table 1. Length, width and dry weight of different prey species used in selection experiments with
M. thermocyclopoides. Values given are Mean SE (n = 20)
Group

Species

Species
abbreviation

Ciliophora

Stylonychia notophora
S. mytilus
Blepharisma musculus

S.n
S.m
B.m

88 10.85
209 20.8
153 28.4

46 7.3
133 15.2
46 15.4

0.008
0.04
0.025

Rotifera

Brachionus angularis
B. rubens
B. calyciflorus
Hexarthra mira
Filinia longiseta
Asplanchna intermedia

B.a
B.r
B.c
H.m
F.l
A.i

88 6.6
149 6.4
197 12.4
160 7.2
98 7.2
745 33.4

77 1.6
135 4.1
155 11.8

51 2.7

0.05
0.16
0.20
0.20
0.20
0.52

Cladocera

Ceriodaphnia cornuta
Moina macrocopa
Daphnia similoides

C.c
M.m
D.s

529 11.0
799 16.4
1446 25.5

363 8.3
456 16.9
804 14.8

3.0
4.60
27.2

Copepoda

Nauplii of
M. thermocyclopoides

98.75 2.5

0.24

containing one individual of Mesocyclops in 100 ml


water. Only non-ovigerous prey were used in every test
to exclude any possible increase in prey density due to
reproduction during the test period. At each treatment
level, four to five individuals not fed for 1216 h prior
to testing were used. The beakers were placed in an
unilluminated BOD incubator maintaining a temperature of 25 1.5 C, and the copepods were allowed
to feed for a fixed period of time, the duration being related to prey size (Table 1). At the end of the
test the copepods were removed from the beaker and
the prey individuals remaining in each beaker were
carefully counted to obtain an estimate of the number
consumed. A random sampling of some beakers did
not reveal any prey losses during the test period due to
death by natural causes. In tests with ciliates as prey,
however, we kept controls (ciliate at the same test densities but without the predator) to account for changes
in density due to reproduction.
Prey selectivity in M. thermocyclopoides was
quantified using Manlys Selectivity Index (Manly,
1974), modified for the situation in which the predator
consumes a substantial portion of the prey available
and hence prey numbers in the medium decline with
time, as in our experiments. The modified formula
(Chesson, 1983) is

Length
(m)

133 4.2

i =

Width
(m)

Dry
weight
(g)

ln(ni0 ri )/ni0
m

ln((nj 0 rj )/nj 0 )

i = 1, 2, 3, . . . , m,

j =1

where i = Manlys alpha (preference index) for prey


type i; ni0 = the number of items of type i present at
the beginning of a foraging bout; ri = number of items
of food type i in the consumers diet; m = number of
prey types.
Manlys selectivity index () values range from 0
to 1, with values for non-selective feeding being 0.5
with two prey types, 0.33 with three prey types, and
so on, in the medium. Index values above these indicate positive selection and values below, negative
selection (avoidance). Differences in the selectivity
values () for test prey species at different proportions
were tested for statistical significance using a two-way
ANOVA test. Deviations in selectivity index values
from the value for non-selective feeding were tested
for statistical significance using Students t-test.
Since the mechanisms of prey selectivity by the
cyclopoid are likely to be explained at least in part by
prey handling times and prey profitability (prey biomass/prey handling time), we measured the handling
times for all the prey species used in this study. One
individual female (age: 68 d) M. thermocyclopoides,
deprived of food during the preceding three hours,

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Table 2. Experimental protocols used for tests of prey selection by M. thermocyclopoides (Species abbreviations are
spelled out in Table 1)
Taxonomic group

Test No.

I. Prey choice within a taxonomic group


Ciliates
1
Rotifers
2
3
4
5
Cladocerans
6
II. Prey choice between different taxa
Ciliates vs. Rortifers
7
8
Ciliates vs. Cladocerans
9
Rortifers vs. Cladocerans
10
11
Nauplii vs. alternate prey
12

Multi-species choice

13

Prey species

S.n
B.a
B.a
B.a
B.c
C..c

B.m
B.r
F.l
B.c
H.m
M.m

B.c
B.r
S.m
B.c
A.i
Nauplii
Nauplii
Nauplii
Nauplii
Nauplii
Nauplii

S.m
B.m
M.m
M.m
M.m
D.s
M.m
C.c
B.c
B.r
S.n

S.n
B.a
C..c

B.m
B.r
M.m

S.m
B.c
B.c

D.s

S.m
B.c
D.s

Combined prey
density

Volume of
container (ml)

Test
duration
(h)

300
200
200
200
200
30

50
100
100
100
100
100

3
6
6
6
3
12

200
200
100
100
40
40
40
40
120
120
220

100
100
100
100
100
100
100
100
100
100
100

6
6
6
12
12
12
12
12
6
6
3

200

See Table 3.

was offered selected prey species at densities similar to those used in the selectivity tests, and was
continuously observed under a stereomicroscope for
1520 min. The prey handling time (time lapsed between an attack of the prey by the copepod and its
resumption of movement after ingestion) was recorded
to the nearest 0.1 s using a stopwatch. In case of cladoceran prey, the consumption was partial; therefore, the
release of the partially consumed prey by the copepod was taken as the endpoint. For each prey species,
510 individual observations were made to obtain an
average handling time.

Table 3. Prey species and densities used in the multispecies prey selectivity (Test 13)
Species

No. 200 ml1

Stylonychia notophora
S. mytilus
Blepharisma musculus
Brachionus angularis
B. rubens
B. calyciflorus
Ceriodaphnia cornuta
Moina macrocopa
Daphnia similoides

500
250
250
75
50
25
10
6
4

Results
In all the prey choice tests, prey types, but not their
relative proportions, had a significant effect on the
value (two-way ANOVA). However, in many tests
the interaction (prey species proportion) term was

statistically significant, indicating that the effect of relative proportions of the prey on selectivity was prey
type-dependent.

415

Figure 1. Selectivity Index () values for M. thermocyclopoides offered a choice of three prey species belonging to one of the three taxaciliates (a), rotifers (b, c) or cladocerans (d) in different proportions. Values shown are Mean SE (n = 45). The dashed line in each indicates
the value for non-selective feeding. Abbreviated species names are spelled out in Table 1.

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Prey choice within a taxonomic group
Ciliates (Test 1)
Given a choice amongst S. notophora, B. musculus and
S. mytilus, Mesocyclops showed a strong preference
for the larger ciliate species S. mytilus ( = 0.44
to 0.59, p < 0.01 one way ANOVA), irrespective
of its relative abundance in the medium (Figure 1a).
The smallest species S. notophora was ingested nonselectively except at high relative abundance, when it
was positively selected ( = 0.41, p < 0.05). There
was a significant (p < 0.01) negative selection (avoidance) ( = 0.120.21) for the intermediate-sized
Blepharisma musculus at any proportion.

less of its proportion in the medium (Figure 1d). While


the copepod strongly avoided D. similoides, the largest
of the three prey species, ( = 0.04, p < 0.01), it
showed no significant selectivity for the smallest of the
three, C. cornuta ( = 0.240.29, p > 0.1).
Prey choice between taxonomic groups
Choice between ciliates and rotifers (Tests 7 and 8)
Offered a mixed diet of the rotifer Brachionus rubens
and the similar-sized ciliate Blepharisma musculus,
the copepod positively selected B. rubens ( = 0.616
0.723; p < 0.05) at all proportions (Figure 3a).
However, given a choice of S. mytilus (ciliate) and
B. calyciflorus (rotifer), the copepod did not exhibit
any statistically significant prey selectivity ( = 0.46
0.54, p > 0.1), except at equal proportions where the
ciliate was selected positively ( = 0.66) (Figure 3b).

Rotifers (Tests 25)


When the choice was among three different species of
the rotifer Brachionus (B. angularis, B. rubens and
B. calyciflorus), the copepod strongly avoided ( =
0.160.22; p < 0.01) the smallest species B. angularis at any relative abundance (Figure 1b). The
intermediate-sized B. rubens was positively selected
( = 0.402, p < 0.05) only when its proportion in
the medium was high (0.5). In a separate two-prey
test (Test 4) that offered a choice between two rotifer
prey, B. angularis and B. calyciflorus differing in size,
the copepod positively selected ( = 0.620.78) the
larger B. calyciflorus and avoided ( = 0.220.38) the
smaller B. angularis regardless of its proportion in the
medium (Figure 2a).
Two tests in our experimental study examined
the prey selectivity patterns of M. thermocyclopoides
when the choice included an evasive prey either
Filinia longiseta along with B. angularis and B. calyciflorus (Test 3), or Hexarthra mira with B. calyciflorus
(Test 5). In Test 3, selectivity for the evasive and illoricate F. longiseta was positive and significant only
when the proportion of either one of the non-evasive
rotifers was high (0.67). Here too as in Test 2, the
smallest of the three species, B. angularis, irrespective
of its proportion in the medium, was avoided by the
copepod ( = 0.100.21). In Test 5, H. mira, larger
and more evasive than F. longiseta, was avoided by
the copepod at any proportion, in the presence of the
alternate prey (B. calyciflorus) (Figure 2b).

Choice between nauplii and other prey types (Test 12)


Although Mesocyclops thermocyclopoides is known
to feed on its own nauplii when there is no choice
(Kumar & Rao, 1999a), it strongly avoided them
( = 0.079 0.014, p < 0.001) when six alternate prey types, M. macrocopa, C. cornuta, D. similoides, S. notophora, B. rubens, or B. calyciflorus
were simultaneously present in the medium (Figure 4).

Cladocerans (Test 6)
A combination of three cladoceran species C. cornuta, M. macrocopa and D. similoidis offered as
food, elicited in Mesocyclops a strong preference for
M. macrocopa ( = 0.6020.718, p < 0.001) regard-

Multispecies prey choice (Test 13)


In a selectivity test including prey species belonging
to different taxa (see Table 3), the copepod showed
no significant selection (p > 0.1) for any taxonomic
group of prey (Figure 5a); however, in terms of species

Choice between a ciliate and a cladoceran (Test 9)


Given a choice of the cladoceran M. macrocopa and
ciliate S. mytilus, the copepod showed a strong ( =
0.130.17) and significant (p < 0.01) negative selection for the cladoceran (Figure 3c).
Preference between a rotifer and a cladoceran prey
(Tests 10 and 11)
Offered a combination of prey types - the cladoceran
M. macrocopa and the rotifer B. calyciflorus, the copepod positively selected ( = 0.740.91) the rotifer,
and avoided the cladoceran irrespective of its relative
abundance (Figure 3d). However, when the rotifer in
the medium was the predatory A. intermedia, the copepods selectivity was strongly positive for it ( = 0.76,
p < 0.01) at equal proportions, but for the cladoceran
when the rotifer proportion was high (Figure 3e).

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Figure 2. Selectivity Index () values for M. thermocyclopoides offered a choice of two rotifer species B. calyciflorus with B. angularis (a)
or with H. mira (b) in different proportions. Values shown are Mean SE (n = 45). The dashed line indicates the value for non-selective
feeding. Abbreviated species names are spelled out in Table 1.

of the prey, the following trends were evident (Figure 5b): (i) the ciliate Blepharisma musculus was
strongly avoided, as could be expected from results of
Tests Ia and IIa; (ii) the cladocerans Ceriodaphnia cornuta and Daphnia similoides were also avoided; and
(iii) the larger rotifer B. calyciflorus and the cladoceran
M. macrocopa were selected positively. The selectivity
values in all three cases were statistically significant
(p < 0.05) against the null hypothesis of random
feeding.

Prey handling times


Prey handling time in M. thermocyclopoides was a
direct function of prey size (Figure 6); prey profitability however, did not show significant correlation with
prey size (r = 0.41, p > 0.1).
Finally, analysis of our data from all the tests with
two prey types, using Landrys (1981) formula, did
not show significant evidence of prey switching in any
test. Although we were unable to analyze the data
from our three prey-choice tests for switching, the lack
of statistical significance for the proportions factor in
two-way ANOVA analyses indicates that probably the

418

Figure 3. Prey selectivity in M. thermocyclopoides offered the choice of a ciliate and a rotifer (a, b), a ciliate and a cladoceran (c), a cladoceran
and a rotifer (d, e) in three different proportions. Values shown are Mean SE (n = 45). The Selectivity Index () value for non-selective
feeding is shown in each case by a dashed line.

419

Figure 4. Selectivity Index () values for M. thermocyclopoides offered Mesocyclops nauplii and an alternative prey type (Abbreviated species
names are spelled out in Table 1). Values shown are Mean SE (n = 45). The dashed line indicates the value for non-selective feeding.

trends in the three-prey tests were similar to those in


the two prey-type tests.

Discussion
Our study demonstrated that Mesocyclops thermocyclopoides is a selective predator. Its selectivity is
probably influenced by important prey attributes such
as size, body shape, taste and behaviour. Although
we did not test it, hunger level of the copepod is
also known to influence prey selectivity (Stemberger,
1985).
The relationship between prey size and prey handling time is helpful in explaining many of the trends
we observed in this study on prey selectivity. For
M. thermocyclopoides prey-handling times increased
exponentially with prey size (Figure 6). Handling
times for ciliates being negligible (<1.3 s), the largest
of them, Stylonychia mytilus was positively selected
regardless of its proportion in the medium, but the
smallest, S. notophora was selected only when its
relative abundance was higher. The higher selectivity of the copepod for S. mytilus was probably due
to its higher encounter frequency and capture probability, which are generally a function of prey size
(Pastorok, 1981; Kerfoot & Sih, 1987). In nature,
size-selective predation by the cyclopoid copepod can
have a marked impact on the ciliate prey community
(Wickham, 1995a,b). The significantly higher mortal-

ities suffered by larger ciliates due to predation by


Diacyclops (Wiackowski et al., 1994) and Cyclops
abyssorum (Wickham, 1998) clearly suggest that cyclopoid predators tend to attack and capture larger
ciliates. The avoidance by M. thermocyclopoides of
the intermediate-sized heterotrich Blepharisma musculus irrespective of its relative abundance, leads us
to speculate that it might possess some chemical defense against the predator, since it does not possess
any other special, predator-deterrent morphological
or behavioural adaptations. Blepharisma was avoided
by the copepod in another two-prey choice test also
with the similar-sized rotifer Brachionus rubens (Figure 3). Distastefulness to predators is a chemical
defense already reported in some ciliate species (Jack
& Gilbert, 1997). Campanella umbellaria was avoided
by Mesocyclops edax because of its distastefulness
(Williamson, 1980; Wickham, 1995a). Reduced cyclopoid predation due to avoidance may be one of the
reasons why Blepharisma is frequently abundant in
local ponds.
Rotifers are often the most common prey for cyclopoids (Plamann et al., 1997). We may invoke body
size again to explain the selectivity patterns shown
by M. thermocyclopoides with rotifer prey. The three
brachionid species (Brachionus angularis, B. rubens,
B. calyciflorus) offered as prey differ from each other
essentially in size only. The copepod showed a positive preference for the largest B. calyciflorus, while

420

Figure 5. Selectivity Index () values for M. thermocyclopoides offered a combination of different ciliate, rotifer and cladoceran prey species
(a). Manlys Selectivity Index () is also shown separately for the three taxonomic groups (b). Abbreviated species names are spelled out in
Table 1. The dashed line indicates the value for non-selective feeding.

avoiding the smallest B. angularis. Larger prey are


encountered more frequently (Kerfoot, 1978; Kerfoot
& Sih, 1987) and yield higher net energy intake per
unit handling cost. Here, the larger B. calyciflorus
turns out to be nearly four times more profitable than
B. angularis to the copepod, since the handling times
for the two species (6.43 0.5 and 5.8 0.58 s) are
not significantly different (p > 0.10, MannWhitney
U -test).
The evasive, darting movements of Hexarthra are
known to reduce risk of predation by the predatory rotifer Asplanchna (Gilbert, 1980; Gilbert &
Williamson, 1978; Iyer & Rao, 1996) and the
predatory copepod Mesocyclops edax (Gilbert &
Williamson, 1978; Williamson, 1983b). The avoid-

ance of Hexarthra by M. thermocyclopoides suggests that its darting movements offer some protection
against predation, but not those of the other evasive
rotifer Filinia, which was selected for by the copepod
when its proportion was relatively high in the medium.
Contributing to its vulnerability further is the fact that
Filinia, which lacks lorica, requires copepod less handling time (1.83 s) than the loricate B. calyciflorus
(6.43 s). The relatively greater vulnerability of illoricate rotifers to copepod predation has been observed
in many studies (Williamson, 1983b). For Filinia, the
risks of an illoricate body apparently far outweigh any
benefits accruing from its evasive movements, which
may offer some protection only against Asplanchna
(Iyer & Rao, 1996) but not Mesocyclops.

421

Figure 6. Prey handling time and prey profitability (calculated using consumed biomass) in relation to prey body size in M. thermocyclopoides
(all parameters on log-scale). Prey species: 1. Stylonychia mytilus, 2. Filinia longiseta, 3. Brachionus angularis, 4. B. calyciflorus 5. Asplanchna
intermedia, 6. Ceriodaphnia cornuta, 7. Moina macrocopa, 8. Daphnia similoides.

Given a choice of three cladoceran prey species


Ceriodaphnia cornuta, Moina macrocopa and Daphnia similoides, Mesocyclops in the present study selected the intermediate-sized Moina regardless of the
abundance of the other two species. Relative to Daphnia and Ceriodaphnia, Moina has a thin carapace
and may therefore be more vulnerable to Mesocyclops
predation. Ceriodaphnia, which otherwise should be
more vulnerable because of its small size, faces less
predation risk from the cyclopoid because it has a
thicker cuticle (Matsumura-Tundisi et al., 1990) and
hence requires more handling time (282 45.6 s)
than Moina (197 23.4 s). Similarly, Mesocyclops
edax was found to feed selectively on the soft-bodied
Diaphanosoma, avoiding the hard-carapaced Bosmina
(Williamson, 1980).
Avoidance of Daphnia similoides by M. thermocyclopoides in the presence of Moina and Ceriodaphnia,
was recorded in this study. Larger cyclopoids such
as Megacyclops gigas (which is >3 times larger than
M. thermocyclopoides) have been observed to select
Daphnia over Ceriodaphnia (Krylov, 1988). In the
present study, the observed selectivity against Daphnia, needs to be interpreted cautiously. Williamson

(1983a) observed that prey size per se is not a dependable determinant of copepods preference when
its diet choice includes prey species differing in other
attributes. The main reasons for our caution in interpreting the selectivity index values are: (1) The
selectivity index was calculated using prey numbers,
but when considered in terms of biomass ingested, it
presents an entirely different pattern. (2) M. thermocyclopoides, like other cyclopoids indulges in what is
known as wasteful killing when dealing with largesized cladoceran prey (Monakov & Sorokin, 1959;
Brandl & Fernando, 1975; Krylov, 1988). Daphnia
similoides, by dry weight, is nearly 6 times larger than
Moina. Although M. thermocyclopoides killed more
than 80% of the Daphnia in the container, it utilized
only a few, that too partially. Our gross estimate of the
average percentage of individual biomass left unconsumed was >70% with Daphnia, but less than 40%
with Moina. Thus, for each individual killed, the average dry biomass ingested was 8.2 g for Daphnia,
and 3.2 g for Moina. Further, we get a more realistic
picture of prey profitability if the actual amounts of
the two cladoceran prey ingested were examined in
relation to the respective handling times, another fac-

422
tor influencing prey selectivity (Roche, 1990; Gliwicz
& Umana, 1994). M. thermocyclopoides spent more
than twice the amount of time handling Daphnia than
Moina, making net profitability to the copepod nearly
the same (0.018 and 0.016) for both the prey species
(Figure 6).
When the diet choice included prey types of different taxa, the selectivity patterns in Mesocyclops were
rather different. Given a choice of the ciliate Stylonychia mytilus and the rotifer Brachionus calyciflorus in
different proportions, the cyclopoid selected the more
abundant of the two prey types. This need not necessarily indicate prey switching since, as we argued
in the case of cladoceran prey, biomasses of the two
prey types are not comparable. Indeed, analysis of our
data (using Landrys formula) gives no evidence of
prey switching. However, field studies are necessary to
find out if in local ponds high brachionid rotifer densities tend to relieve Mesocyclops predation pressure on
hypotrich ciliates such as Stylonychia.
Our preceding arguments on Daphnia selectivity
help also to explain the negative selection (avoidance)
that Mesocyclops exhibited for Moina in the Moinaciliate (Figure 3c) and Moina-rotifer (B. calyciflorus)
(Figure 3d) choice tests. Since the weight of one adult
Moina is equal to the combined weight of 115 individuals of S. mytilus, it is evident that the copepod
actually ingested more biomass of the cladoceran than
of the ciliate prey.
The cooccurrence of M. thermocyclopoides with
the predatory rotifer Asplanchna intermedia is not
uncommon in local ponds and small lakes. As both
regularly prey on brachionid rotifers, the observed
efficiency of predation on Asplanchna by M. thermocyclopoides has implications for the population
dynamics of brachionid rotifers in nature. Williamson
& Gilbert (1980) suggested that Mesocylops edax, by
feeding on Asplanchna girodi, could reduce its effect on the rotifer (Keratella cochlearis) prey. We
have no field data to assess the relative importance
of Mesocyclops and Asplanchna predation on Brachionus calyciflorus; however, in a laboratory study
of the dynamics of a two predator (Asplanchna intermmedia and M. thermocyclopoides)-one prey (B. calyciflorus) system, we found that B. calyciflorus persisted
longer in it than in controls with Asplanchna alone
(Kumar & Rao, 2001).
Cannibalism in copepods has been well documented (Landry, 1981; Krylov, 1988; Van den Bosch
& Gabriel, 1991). Although Mesocyclops feeds on
nauplii (of the same species, not necessarily of its own

progeny) in the absence of a choice, it avoided them


when alternate prey (ciliates, rotifers or cladocerans)
were available in the medium (Figure 4). In an earlier
study (Kumar & Rao, 1999a), we had demonstrated
that in the presence of algae in the medium, the ingestion rates of this copepod on nauplii were 3690% less
than in the medium without algae.
In multispecies prey choice tests (Figure 5), the
most important trends exhibited by M. thermocyclopoides were: (1) The ciliate Blepharisma and the
cladocerans Daphnia and Ceriodaphnia were negatively selected. (2) The rotifer B. calyciflorus and
the cladoceran Moina macrocopa were positively selected. (3) Other prey species were ingested in a nonselective fashion. The first two trends are consistent
with the results we obtained in the two- and threespecies selectivity tests. In an environment where it is
not uncommon for the copepod to encounter a multispecies prey situation, it is likely that its optimal
diet breadth is governed by more than one criterion
of selection. Although, M. thermocyclopoides, like
other cyclopoid species studied (Krylov, 1988; Janicki
& DeCosta, 1990; Matsumura-Tundisi et al., 1990),
shows strong selection for cladoceran prey species, in
nature it may feed in a more opportunistic way, taking advantage of the availability of a wide spectrum
of prey species of ciliates, rotifers and cladocerans.
Our earlier laboratory study (Kumar & Rao, 1999b)
showed that M. thermocyclopoides achieved its highest reproductive output on a mixed diet of algae,
ciliates, rotifers and cladocerans.
Our present study did not provide any evidence
for prey switching by M. thermocyclopoides, but it
does not necessarily suggest that the copepod lacks
such ability. Prey switching has been reported in two
calanoid species Calanus pacificus (Landry, 1981) and
Acartia tonsa (Kirboe et al., 1996); the prey choice
offered in both cases was between algae and animal
prey. That two entirely different methods are used
by these copepods for capturing algae (suspension
feeding) and animal prey (raptorial feeding) offers a
plausible mechanism by which prey switching could
occur. In the present study however, the prey choice
did not include algae, and therefore we cannot speculate whether M. thermocyclopoides would have shown
switching if the alternate prey had been an alga.

423
Acknowledgements
We are grateful to William DeMott and two anonymous reviewers for their valuable comments and suggestions on an earlier draft of this paper. We thank
Anupama Kak, J. Vanisree and Anita Nair for technical assistance, and gratefully acknowledge the financial support provided to one of us (R.K.) by the
Council of Scientific and Industrial Research.
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