The Impact of Fish Exclusion on the Abundance and Species Composition of Larval Odonates:

Results of Short-Term Experiments in a North Carolina Farm Pond

Author(s): Peter J. Morin
Source: Ecology, Vol. 65, No. 1 (Feb., 1984), pp. 53-60
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1939457
Accessed: 18-03-2015 21:37 UTC

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 colo(i %. 65(1). 1984. pp. 53--60
C 1984 by-the Ecological Society of America

THE IMPACT OF FISH EXCLUSION ON THE

ABUNDANCE AND SPECIES COMPOSITION OF
LARVAL ODONATES: RESULTS OF SHORT-TERM
EXPERIMENTS IN A NORTH CAROLINA
FARM POND'

PETER J. MORIN2
Department >t Zoology, Duke Ulnit'ersity, Durham, North Carolina 27706 USA

Abstract. Under natural conditions in a man-made North Carolina farm pond, a single small
species, Perithelnis tenera, numerically dominates a littoral assemblage of larval dragonflies. Censuses
of exuvia of larvae metamorphosing from the pond in three successive years demonstrated a consistent
negative correlation between numerical dominance in the assemblage and species-specific size in the
final larval instar. This pattern suggested that size-dependent processes, such as vertebrate predation,
might structure the odonate assemblage.
Exclusion of vertebrate predators from patches of emergent vegetation by screen enclosures for
3 mo increased total abundances of larval odonates by an order of magnitude. Dominance shifted
from small species to species of intermediate size, especially Pach'dipl/a lo/ngipenllis. where fish
were excluded. The largest species remained rare and were unaffected by fish exclusion. Four odonate
species were facultatively multivoltine, completing larval development within 1-2 mo after coloni-
zation of the fish enclosures. Fish exclusion also increased the abundance of large microcrustacea,
especially the cladoceran Sitnocephalus serrulatuis. Large microcrustacea increased in abundance
despite a corresponding increase in the abundance of their macroinvertebrate predators, where fish
were excluded.
The usual dominance of small Perithelnis in this assemblage was a probable consequence of
differentially heavy predation by fish on moderate-sized species, and the unexplained failure of large
species to recruit well under experimental or natural conditions. These results support the general
importance of predation in structuring freshwater communities and document an important mode of
population regulation operating during the aquatic larval phase of the odonate complex life cycle.
hey, ittords: Anisoptera: comninunit' structure; Iratgonflst; fish preda tion; Lepomis; Ilnacroen-
thlos; Micropterus; North Carolina; Perithemis: Pachydiplax; ponds; pred(ltion; zooplanktoni.

INTRODUCTION of community structure (e.g., Zaret 1980). Results from

The impact of vertebrate predation on the species
composition of freshwater planktonic macroinverte-
brates is well known (Brooks and Dodson 1965, Zaret
1980) and provides a striking example of the efficacy
of predation in structuring one component of fresh-
water communities. In contrast, the role of vertebrate
predation in structuring assemblages of freshwater
macroinvertebrates remains controversial. Compara-
tive studies suggest that fish predation may regulate
the abundance and species richness of aquatic insects
(e.g.. Macan 1977). One group of experiments indi-
cates that fish influence the biomass and abundance of
benthic macroinvertebrates (Hall et al. 1970, Eriksson
1979. Crowder and Cooper 1982). However, other ex-
periments suggest that exclusion of fish from the lit-
toral zone has little impact on the abundance or species
richness of benthic macroinvertebrates (Thorp and
Bergey 1984I, h). Divergent outcomes of similar ma-
nipulations of predators in different freshwater com-
munities complicate the synthesis of emerging theories
I Manuscript received 28 July 1982; revised and accepted
27 December 1982.
2
Present address: Department of Biological Sciences, Nel-
son Labs, Busch Campus, Rutgers University, Piscataway,
New Jersey 08854 USA.
a larger sample of communities are needed to correlate
disparate responses to fish exclusion with other mea-
sured differences among communities. Important fac-
tors responsible for these different responses may in-
clude variation in habitat complexity or natural levels
of predator abundance among communities (Crowder
and Cooper 1982).
In this paper I show that experimental exclusions of
fish and other aquatic vertebrates dramatically alter
the abundance and dominance of species comprising
one important assemblage of littoral macroinverte-
brates, the aquatic larvae of dragonflies (Odonata: An-
isoptera). These experiments tested a hypothesis in-
spired by the following observation: the odonate
assemblage in my study pond was dominated by a small
late-breeding species, despite the fact that such species
are at a competitive or predatory disadvantage among
larger, early breeding odonates in similar southeastern
ponds (Benke 1978, Benke et al. 1982). I postulated
that dense populations of size-selective fish might foster
the predominance of small species by depressing the
abundance of larger odonates that might otherwise
outcompete or prey on smaller species. These short-
term exclusion experiments confirm the proposed
efficacy of vertebrate predators in controlling this im-

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54 PETER J. MORIN
portant assemblage of freshwater invertebrate preda-
tors (Benke 1978, Johnson and Crowley 1980), illus-
trate one mode of population regulation in species with
complex life cycles (Wilbur 1980), and provide new
insight into the duration of larval development of sev-
eral odonate species.
MATERIALS AND METHODS
I studied the larval odonate assemblage of a 0.44-ha
man-made farm pond (Carroll's Pond) located in Or-
ange County, North Carolina. Carroll's Pond is :28
yr old, has a maximum depth of ;3 m, and provides
water for a small herd of cattle. The pond's narrow
zone of emergent vegetation (1-2 m wide) is dominat-
ed by an aquatic knotgrass, Paspalurn distichum. Beds
of emergent Paspalum are the principal oviposition
sites patrolled by adult dragonflies, and most larval
dragonflies metamorphose in or near these beds. Elev-
en species of dragonflies successfully metamorphose
from the pond in most years. Fourteen additional
species patrol and oviposit, but their larvae fail to
metamorphose from Carroll's Pond in most years (see
Appendix). Damselflies (Odonata: Zygoptera) are nei-
ther conspicuous nor abundant in this community. The
most abundant vertebrate predators in Carroll's Pond
are the fish Lepomnisinacrochirus (bluegill sunfish) and
Micropteiuis sa/hnoides (large mouth bass). Lepomnisis
known to prey heavily on larval odonates (Sadzi-
kowski and Wallace 1976). Other potential predators
include the turtles Chrysemnys scripta, Sternother-tis
odora tius, and Chelydra s e}p en tina.
I measured natural patterns of larval odonate abun-
dance in the entire pond by collecting cast exoskele-
tons (exuvia) left around the margin of the pond by
larvae as they metamorphosed into teneral dragonflies.
Exuvial census is a standard technique for estimating
the abundance of many aquatic insects (Benke and
Benke 1975, Southwood 1978). Exuvia were collected
daily from the entire perimeter of the pond during each
annual period of metamorphosis (usually from early
April through late September) in 1979, 1980, and 1981.
The shoreline was bordered by closely cropped pas-
ture and mown lawns, restricting suitable perches for
metamorphosing odonates to the immediate margin of
the pond, where exuvia were conspicuous and readily
collected. Most exuvia were concentrated in a narrow
band within 1 m of the shoreline (Jacobs 1955). Al-
though exuvia were occasionally lost during periods
of heavy rain, total counts of exuvia collected from
the entire perimeter of the pond provided the best
available estimate of the abundance of each odonate
species metamorphosing from the pond in a given year.
I experimentally examined the short-term impact of
vertebrate predators on odonate species composition
by measuring larval recruitment and metamorph pro-
duction in screen pens that either excluded large ver-
tebrate predators or remained permeable to fish and
turtles. I constructed eight open-topped pens (1.22 m
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Ecology, Vol. 65, No. 1
long x 1.22 m wide x 0.61 m high) of wooden frames
completely covered by bottom and side panels of Fi-
berglas screen (7 meshes/cm). I made four pens
permeable to large predators by cutting a 0.45 x 0.61
m window in the screen at the center of one side panel.
Hereafter, these are termed open pens. The four re-
maining pens (hereafter termed enclosures) lacked such
openings and excluded fish and turtles. The pens were
placed in a discrete 63 m long Paspal/um bed along the
north shore of Carroll's Pond on 22 June 1981. I added
natural densities of well-washed Paspalum to each pen
to provide cover. Pen walls projected at least 0. 1-0.2
m above the waterline. Female dragonflies began ovi-
positing in the pens within several hours after pen
placement. I augmented natural recruitment by adding
first-instar hatchlings of several species to all pens,
before it became obvious that larvae would naturally
colonize the pens. On 16 and 20 July 1981, I added the
following numbers of first-instar odonates to each pen:
55 Tram ea lacerata, 55 Celithemis fasciata, 100 Li-
bellula incest, 100 Erythemnis sitpli/icollis, and 50
Plathemis lydia. In addition, small instars of odonates
in the Pa spa/lum bed could colonize the pens by pass-
ing directly through the screen mesh. Of course, open
pens were permeable to odonates of all sizes.
Paspaluin grew rapidly and provided dense cover
and spatial heterogeneity for weed-dwelling odonates.
After 1-2 wk, habitat complexity within the pens re-
sembled that found in the unmanipulated Paspalum
bed. Several millimetres of silt and detritus accumu-
lated on the pen bottoms, providing ample habitat for
burrowing and bottom-sprawling species. Grazing by
tadpoles of Rana catesbeiana prevented fouling of pen
walls by periphyton. Tadpoles entered the open pens
naturally, and each exclusion pen was stocked with 6-
10 tadpoles.
I assessed the effects of fish exclusion, by measuring
the abundance of naiads either metamorphosing from
pens or remaining in the pens at the termination of the
experiment in late September 1981. Larvae remaining
in the pens in late September were diapausing individ-
uals that presumably would have continued develop-
ment during the following spring (Paulson and Jenner
1971). I collected exuvia daily within each pen, until
metamorphosis ceased in enclosures and the pond in
mid-September. Direct observation of nocturnally
eclosing naiads indicated that exuvia collected from
the interior walls and vegetation of each pen originated
from larvae living in the pens. Overhanging wooden
frames (;3 cm wide) on each pen's top prevented me-
tamorphosing naiads from wandering outside the pens
to eclose.
I made comparable daily collections of exuvia from
the 63-m Paspalurn bed surrounding the pens, and from
the remaining pond shoreline. I standardized counts
of exuvia from the pens and unmanipulated portions
of the pond to numbers of metamorphosing odonates
per metre of shoreline per census interval, to facilitate
February 1984 FISH PREDATION ON LARVAL ODONATES
comparison of emergence rates among natural and ex-
perimental habitats. I assumed that each pen corre-
sponded to 1.22 m of shoreline. Emergence rates were
calculated only over the census interval in which odo-
nates metamorphosed from both the pens and the pond,
23 July through 18 September 1981.
The experiment was terminated after =3 mo, on 22,
23, and 24 September, when I carefully lifted the pens
and their contents from the pond. Overwintering odo-
nate larvae retained by the screen mesh were sorted
from debris by hand and preserved. The contents of
open pens provided a baseline estimate of the abun-
dance of odonates recruiting in the presence of pred-
ators, via a sampling protocol that was directly com-
parable to the census of the contents of predator
exclosures. All sampling biases were assumed to be
similar between treatments, and differences in the
contents of open and predator-exclusion pens were
attributed to direct or indirect consequences of the
presence or absence of large vertebrate predators. Be-
cause these experiments were conducted for less than
a complete yearly cycle of odonate oviposition, de-
velopment, and metamorphosis, they cannot speak to
entire patterns of odonate abundance described by
collections of metamorphosing naiads over a full year.
However, short-term predator exclusions effectively
document the consequences of predation in compa-
rable experimental communities, and can identify
species that are susceptible to predation.
I used three statistics to describe differences in pop-
ulations of each odonate species between open pens
and fish enclosures: (1) the number of metamorphosing
naiads per pen, (2) the number of overwintering naiads
per pen, and (3) the relative abundance of each species
in each pen. Relative abundance was defined by the
fraction of the total number of metamorphosing and
overwintering naiads in each pen belonging to each
species. Relative abundance is a measure of domi-
nance that is statistically independent of differences in
total odonate abundance. Species can conceivably dif-
fer in total abundance between treatments, while not
differing in relative abundance, if dominance is inde-
pendent of variation in total odonate carrying capacity
among pens.
I also sampled fish densities at the termination of
the experiment by rapidly tilting the open side of each
open pen out of the water, trapping foraging fish. Pens
were then completely lifted from the water, and trapped
fish were collected, anesthetized, and preserved.
Densities of fish in open pens were calibrated against
natural densities during a similar experiment in 1982
using a catch-per-unit-effort technique. Plastic min-
now traps were placed in each of four open pens and
four haphazardly selected sites in the natural weed
bed. Traps were left in place for 24-48 h, and then
trapped fish were censused. Large Lepounis could not
enter the traps, but fish in the small size-classes pre-
dominating in the pens in 1981 were readily captured.
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55
I assumed that if open pens concentrated fish above
natural densities, more fish would be caught per min-
now trap per day in the pens than in the weed bed.
Low visibility and thick vegetation precluded visual
methods of estimating fish densities.
I measured one other aspect of pen community
structure to compare the resemblance of open pens to
the natural weed bed. At the termination of the ex-
periment, microcrustacea and other small arthropods
were sampled in each pen and at four haphazardly
chosen sites in the weed bed. Samples were taken by
plunging an inverted l-L polyethylene bottle into the
weeds to a depth of 0. 15 m and then allowing the
bottle to fill with pond water and organisms. Samples
were filtered through an 88-,um mesh Nitex filter, and
filtered organisms were preserved in a solution of su-
crose and formaldehyde (Haney and Hall 1973). All
preserved microinvertebrates in each sample were
identified and counted with the aid of a dissecting mi-
croscope. Counts of each taxon from the two kinds of
pens and the unmanipulated weed bed were compared
(via ANOVA and Duncan's multiple range test) to de-
tect differences in the availability of these odonate re-
sources (Johnson 1973, Benke 1978) between natural
and experimental habitats. These samples also provid-
ed an indirect measure of the intensity of fish predation
in open pens and the natural weed bed. I assumed that
if the open pens trapped fish and concentrated verte-
brate predators above natural densities, abundances
of larger microcrustacea would be significantly lower
in the open pens than in the weed bed, due to corre-
sponding differences in the intensity of predation by
size-selective fish (Zaret 1980).
RESULTS
Censuses of metamorphosing naiads revealed rela-
tively similar patterns of dominance and abundance
over three successive years of emergence (Table 1).
The smallest species, Perithemis tenera, was an order
of magnitude more abundant than any other anisop-
teran in each year. In general, numerical dominance
and species-specific body size were negatively corre-
lated. Over the entire odonate assemblage, there was
a significant negative correlation between species-spe-
cific body length in the final larval instar and the log-
arithm of the abundance of final-instar naiads meta-
morphosing from the pond within each year (Table 1).
Species often varied considerably in abundance among
years, but this variation had little impact on the broad
patterns of dominance described by the ranked abun-
dances of species within years.
Exclusion of vertebrate predators by screen exclo-
sures dramatically altered both the absolute abun-
dance (number per pen) and relative abundance (per-
cent per pen) of metamorphosing and overwintering
odonates (Table 2, Fig. 1). Few odonates metamor-
phosed from the open pens containing fish. However,
three species, Pachydiplax longipennis, Erythemis
56 PETER J. MORIN Ecology, Vol. 65, No. 1

TABLE 1. Numbers of odonate exuvia collected from the entire perimeter of CalToll s Pond in each of three successive
years of daily collections. Species-specific mean body lengths in the final instar are based on 5-10 individuals of each
species collected in 1979 and 1980, except for DronogioIg phus. Numbers in parentheses are the rank abundance of each
species within each year. Correlations at the bottom of the table describe the relation between the logarithm of abundance
and mean body length over all species in each year.

Body length Abundance (no./yr)

Species (mm) 1979 1980 1981
Perithelnis tenera 13. 1 2432 (1) 3766 ( 1) 5324 ( 1)
Ervthemlios simplicicollis 15.2 329 (3) 181 (3) 61 (4)
PachXvdiplax iongipenniis 16.9 491 (2) 589 (2) 234 (2)
Epitheca cVnosural 18.0) 57 (5) 137 (4) 21 (6)
Platheinis lvdia 19.7 68 (4) 99 (5) 104 (3)
Libellula luctuosa 21.7 17 (7) 25 (7) 32 (5)
Gniphus brisnle vi 22.2 11 (8) 26 (6) 6 (7)
Latdona deplanata 22.4 2 (10) 10 (8) 0 (10)
Epicordulia regina 23.6 4 (9) 0 (10) 1 (9)
Trainea lacerat(I 27.1 43 (6) 7 (9) 3 (8)
Dromogoinphus spinosus 30.9 1 (II) 0 (10) () (10)
Correlation between body length
and log, abundance -0.83 --0.90 -0.84
Correlation significance level .01 .01 .01

Sim/plic(ollis, and Perithemis teliera, metamorphosed
in abundance from fish exclusion pens. Small numbers
of the large species Trameia l/(cer(at(a also metamor-
phosed from the fish exlcusion pens. The first meta-
morphs were collected from fish exclusion pens on 23
July 1981, and metamorphs continued to appear through
18 September 1981. Because all odonates entered fish
exclusion pens as eggs or very small larvae, these four
species clearly completed larval development in only
1-2 mo.
Perithemis tencra, the smallest species, was the nu-
merically dominant species metamorphosing from both
the open pens and the natural weed bed (Tables I and
2). In contrast, predator exclusion generated a striking
increase in the abundance of metamorphosing Pachy-
diplax longipeunis, a species of intermediate size. More
Paochvtiplax metamorphosed from the four predator
enclosures combined (the equivalent of 4.88 m or 2. 1%
of the pond's shoreline) than were collected from the
entire perimeter of the pond (229.8 m) in most years
(Tables I and 2).
Low metamorph production in the open pens was
similar to rates of metamorph production in the PucN-
pa/lumi bed. Numbers of exuvia per metre of shoreline
collected between 23 July and 18 September for the
three most abundant species were: 6.5 Perithemiis/m,
0.2 Ervthemiislm, and 0.3 Pach vdipl/a/m in the Pas-
p(/uclu bed; 1.8 Perithemnislm, 0.2 Er'theniislm, and
0.0 Pachx'dipl/x/m in the open pens; 18.4 Perithe-
inisIm, 13.7 Ervthemnislm,and 108.8 Pachyvdipllx/m in
the predator exclosures. Differences in average meta-
morph production between the Paspalunm bed and the
open pens were trivial compared to observed differ-
ences between open and fish exclusion pens.
Predator exclusion also greatly enhanced the abun-
dance of overwintering naiads of moderate-sized
species (Table 2). Species with significantly enhanced
larval abundance in the absence of fish included Erx'-

TABLE 2. Absolute abundances (mean ? SD) of metamorphosing and overwintering larvae in open and fish exclusion pens.
Values in columns headed P indicate the level of significance (from ANOVA) of differences in the abundances of each
species attributed to the presence or absence of fish. Dashes indicate species that did not metamorphose from either
treatment.

Metamorphosing larvae Overwintering larvae

(no./pen) (no./pen)
Species Fish present Fish absent P Fish present Fish absent P
Perithemiis tenera 2.3 ? 2.6 22.5 ? 24.9 .150 22.3 ? 14.3 55.0 ? 64.2 .358
Er 'themiis simnplicicollis 0.3 ? 0.5 17.3 ? 4.6 .001 11.8 ? 5.5 184.0 ? 99.9 .014
Pachvdiplax longipennis 0.0 + 0.0 143.0 ? 62.3 .004 26.0 ? 15.3 423.3 + 113.6 .001
Epitheca c-ynosur- - 8.3 ? 8.5 53.0 ? 25.1 .015
Platheinis lvdia - 0.8 ? 0.5 20.0 ? 8.3 .004
Libell/lu ineesta - - - 0.8 ? 1.0 43.0 ? 12.8 .001
Libel/u/l luctuosa - - - 0.3 ? 0.5 48.0 ? 24. 5 .008
Epicorululia region - - - 0.5 ? 0.6 0.8 ? 1.0 .670
Triniea lacerata 0.0 ? 0.0 2.0 ? 2.2 .114 0.0 ? 0.0 0.3 ? 0.5 .355

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February 1984 FISH PREDATION ON LARVAL ODONATES 57

60 fish
0 p~~~~~~~~~~~resent
50 LIZ......
-o
M
40 *t absent
~~~~~~~~~~~~fish

<30

20

Perithemis Erythemis PKchydipKax Epitheca Plathemis L.incests L. luctuosa

small * - Final Instar Body Size - N large

FIG. I. Mean relative abundance calculated from combined abundances of metamorphs and overwintering larvae of each
odonate species collected from the experimental pens. Relative abundances describe the percentage of the total number of
odonates in each treatment belonging to each species and describe changes in numerical dominance that are independent of
absolute total odonate density. Species of consistently low absolute and relative abundance are not shown. When a species
exhibited significant differences in relative abundance between treatments (via ANOVA), those differences are indicated by
P < .05 or ** P < .01 above the species bars.

tilheniis simplici(/5llis, P(l(hw(/ipl(ax longipeUnnis, Epi- vae of L. ilcesta were easily distinguished from L.
thieca cviosura, Pl(1the mis lNdia, Libell/lw infcest(a, and lucktuosa by counts of setae on the lateral lobes of the
Libell/lu /uctuos(a. In contrast, the abundance of over- labium (Needham and Westfall 1955).
wintering instars of small Perithe mis tenera did not Fish were abundant in all open pens at the termi-
increase significantly under fish exclusion, while its nation of the experiment, while only one small Le-
relative abundance (dominance) actually declined be- poinis was found in one of the four predator exclusion
cause of the increased abundance of other species (Fig. pens. Means and ranges of fish densities in the open
1). Relative abundances of Pach'(lipl/-x lonlipennis pens were 13.8 Lepomnis mal(Ilrofhirus/pen (range 12-
and Libellula spp. increased concomitantly with their 18) and 1.8 Micropterus s(/lhnoides/pen (range 0-4).
increased absolute abundance in fish exclosures. Most fish collected from the pens were small individ-
However, absolute abundances of Ervthetmis simpli- uals with mean standard lengths (snout to caudal pe-
collis, Plathemiis ltdia, and Epithecac(lonosur(l in- duncle; ranges in parentheses) of 43 mm (24-129 mm)
creased in proportion to the total abundance of the for Lepomiis and 50 mm (40-62 mm) for Mic ropterus.
entire odonate assemblage, resulting in unchanged rel- Comparison of the abundance of small Lepomiis
ative abundances for these species. The two largest captured in minnow traps placed in either open pens
species collected from the pens, Epicordulia regina or the weed bed in 1982 indicated that open pens did
and Traenca /acer(t(a, were rare in both treatments not concentrate fish above natural densities (Table 3).
and were apparently unaffected by fish exclusion. Ex- In fact, results from four successive trapping intervals
clusion of vertebrate predators produced a clear shift in July 1982 indicated that the number of fish per trap
in absolute and relative abundance from assemblages per sampling interval was slightly lower in the open
dominated by a small species to assemblages domi- pens than in the weed bed.
nated by species of intermediate size, especially Abundances of microinvertebrates sampled from the
Pac hx'diplax longipennis.
Three relatively uncommon species known to meta-
morphose from the pond (Ladiona deplanata, Gomn- TABLE 3. Means and ranges of the number of Lepoinis intic-
phuis brien/evi, and Dromnogoinphuis spinosus) failed to rochirus captured in plastic minnow traps placed in open
recruit in the pens. Ladtona (lepl/anat and Gomnphus pens or the unmanipulated weed bed on four sampling dates
in 1982. Open pens were identical to the pens used in 1981,
briieN/'1i are spring-breeding species that had com- and were located in similar positions in the weed bed. Sam-
pleted reproduction long before the pens were placed pling dates and sampling intervals indicate the timing and
in Carroll's Pond. Neither of these species was suffi- duration of each trapping effort. All means are for four
replicates in each habitat on each date.
ciently abundant under natural conditions to contrib-
ute conceivably to the fish exclusion effect. Droano-
Sampling Lepomis per minnow trap
gouiphus spinosus is quite rare at Carroll's Pond, and interval
no ovipositing adults were observed in 1981. Interest- Sampling date (h) Weed bed Open pens
ingly, Libellula incest, a species that failed to meta- 10 July 1982 24 1.8 (1-2) 1.3(0-3)
morphose from the pond despite the presence of large 11 July 1982 24 1.8 (1-3) 1.0 (0-3)
numbers of ovipositing adults, recruited in predator 12 July 1982 24 1.5 (0-4) 1.5 (1-2)
14 July 1982 48 2.8 (1-4) '.0(0-3)
exclusion pens at moderate levels of abundance. Lar-

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58 PETER J. MORIN Ecology, Vol. 65, No. 1

TABLE4. Comparison of the abundance of various microinvertebrates in experimental pens and the unmanipulated weed
bed in 1981. Abundances are the mean (?SEM) number of animals in four replicate samples from each habitat. Overall
differences among habitats are indicated by significant values of P from an ANOVA for each species over all habitats.
Specific differences among habitats are indicated by results of Duncan's multiple range tests; means that do not differ
significantly are connected by underlining.

Microinvertebrate abundance (no./L)

Taxon Exclusion pens Open pens Weed bed ANOVA P
Sibnocephal/us serrulatus 320.3 ? 114.4 29.8 ? 8.6 17.8 + 3.9 .0004
Ostracods 22.5 ? 6.1 99.8 228.7 82.7 ? 9.3 .0080
Cyclopoid copepods 33.0 ? 4.6 62.5 + 22.9 34.3 ? 7.9 .53
Chvdorus sphaericus 16.3 ? 3.3 33.5 + 11.3 13.8 ? 3.5 .16
Scapholeberis kingi 0.8 + 0.8 4.5 + 1.7 1.0 ? 1.0 .08
Mites 1.0 ? 0.4 2.0 + 0.0 2.8 + 0.5 .03
Total 393.8 ? 118.7 232.0 + 17.3 152.3 ? 19.1 .04

open pens did not differ significantly from abundances in Carroll's Pond and elsewhere (Crowder and Cooper
in the Paspaluin bed (Table 4). In contrast, the aver- 1982) suggest that it may be more profitable to ask why
age abundance of Simocephalus serrulatus, a large lit- the impact of fish exclusion varies among communi-
toral cladoceran (adult body length 1-3 mm), was sig- ties, than to suggest that the species composition of
nificantly enhanced by fish exclusion. This increase in freshwater macroinvertebrates is generally unaffected
large microcrustacea occurred despite an order of by fish predation (Thorp and Bergey 198 la).
magnitude increase in the abundance of odonates which Differences in odonate abundance and species com-
also prey on Cladocera. These results further empha- position among caging treatments in Carroll's Pond
size the close resemblance of open-pen communities cannot be easily dismissed as cage artifacts. Several
to the natural weed bed and illustrate the pleiotropic lines of evidence suggest that the open pens faithfully
effects of fish exclusion on community structure. mimicked the natural community found in the Pas-
p(luln bed. Open pens did not concentrate fish above
DISCUSSION natural densities measured in the surrounding Paspa-
Striking differences in the abundance and domi- lum bed. Abundances of various microcrustacea were
nance of odonate species between open pens and fish similar in both the open pens and the weed bed. Emer-
exclosures clearly implied that fish can influence the gence rates of odonates in open pens and the weed
structure of odonate assemblages. These results sup- bed were similarly low compared to much higher
port the general importance of fish predation in struc- emergence rates in fish exclusion pens. No evidence
turing freshwater communities (e.g., Zaret 1980, suggests that open pens exaggerated the effects of fish.
Crowder and Cooper 1982) and experimentally con- Odonate responses to the presence of fish can be
firm previously untested hypotheses regarding poten- parsimoniously attributed to direct effects of verte-
tial relations between fish predation and odonate brate predation. Lepomlis Inacrochirus is known to prey
species composition (Benke 1978, Johnson and Crow- heavily on larval odonates (e.g., Sadzikowski and
ley 1980). Wallace 1976). Shifts of dominance from smaller to
It is unclear why previous fish exclusions in some larger odonate species were qualitatively similar to well-
communities containing many of the same odonates known responses of zooplankton species composition
failed to produce analogous results (Thorp and Bergey to fish predation; a small species predominated when
198 la). Disparate responses of communities to fish ex- fish were present, but larger species became dominant
clusion may simply reflect variation in the abundance when fish were excluded (e.g., Brooks and Dodson
of predators among ponds and lakes. Exclusions of 1965, Zaret 1980). This result appeared to be a simple
sparse populations of fish may have little apparent ef- consequence of the severe impact of fish on the abun-
fect on community structure when compared to results dance of odonates with moderate body sizes in the
of similar exclusions where fish occur at much greater, final larval instar. Fish did not enhance the abundance
but nonetheless natural, densities. Different responses of any odonate species, offering no evidence for shifts
to predator exclusion may also reflect variation among in dominance caused by predator-mediated competi-
communities in habitat complexity, given the impor- tion among prey (Paine 1966, Morin 1981).
tant role that spatial refuges play in moderating the The strong depression of microcrustacean abun-
intensity of predation in freshwater communities dance by fish suggested an additional mechanism that
(Benke 1978, Crowder and Cooper 1982). Striking re- may have contributed to the effect of fish exclusion on
sponses of the weed bed community to fish exclusion odonate abundance; fish and odonates may conceiv-

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February 1984 FISH PREDATION ON LARVAL ODONATES
ably compete for microcrustacean prey. Fish exclu-
sion dramatically increased the abundance of the large
cladoceran Silnocephalus serraulatus, despite a corre-
sponding increase in the abundance of larval odonates
that also prey on microcrustacea. Similar increases in
the abundance of Silnocephalus follow fish exclusion
in other communities (D. M. Johnson, personal coln-
minnication). Observed effects of fish on odonates may
result both from direct predation on naiads and from
competition between fish and surviving naiads for re-
duced stocks of microcrustacean prey. This latter pos-
sibility is offset by the observation that benthic insect
larvae may contribute more to odonate diets than do
microcrustacea (Benke 1978).
The consistent negative relation between body size
of final-instar larvae and population sizes of various
species of metamorphosing odonates initially suggest-
ed that a simple relation might exist between body size
and susceptibility to predatory exclusion. Although all
odonates pass through a wide range of sizes during
larval development, size in the final instar may be crit-
ical in determining larval success in the face of size-
selective vertebrate predation. If vulnerability to pre-
dation increases with body size, small species may
have an obvious advantage over larger species. Small-
er species can metamorphose at smaller, less vulner-
able body sizes than their larger counterparts. Larger
species are committed to metamorphosis at larger sizes,
and may thereby incur greater intensities of size-de-
pendent vertebrate predation. Interspecific differences
in size among larvae metamorphosing from Carroll's
Pond corresponded to well-known species-specific dif-
ferences (Needham and Westfall 1955), indicating that
the negative relation was not simply a manifestation
of density-dependent growth. However, the large
species Epicordulia retina and Tralnea lalcera ta re-
mained equally rare, regardless of altered fish densi-
ties. The failure of these large species to respond to
fish exclusion suggested that their low abundances in
Carroll's Pond were not explained by size-selective
fish predation.
The negative relation between size and abundance
in this odonate guild clearly has a complex basis that
may be only partially related to predation. Further,
the consistent reproductive failure of some odonates
listed in the Appendix cannot be generally attributed
to fish predation. Only one of the species that regularly
failed to metamorphose from Carroll's Pond, Libellula
incest, increased in abundance in the absence of fish.
The continued failure of larval recruitment of other
abundant ovipositing species (e.g., Celitheinis faiscia-
ta, Libell/la cvanea, and others) suggested that their
exclusion in the larval phase of the life cycle was not
related to fish predation. Adults of these species pre-
sumably maintain populations at Carroll's Pond only
by migration from other ponds where they are repro-
ductively successful.
The potential roles of physiological constraints,
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59
habitat specificity, and competition with predator-re-
sistant species in producing exclusions and patterns of
dominance require further study. For instance, ob-
served patterns of larval dominance may have simply
resulted from variation in the availability of critical
microhabitats (e.g., Crowley and Johnson 1982), and
this pattern may have been only coincidentally cor-
related with body size. A negative relation between
final-instar size and dominance may not generalize to
other communities (Thorp and Bergey 1981l, Crowley
and Johnson 1982), but such comparisons are clouded
by differences in sampling techniques and potential
variation in predation intensity among studies.
In addition to documenting the effects of fish on
odonate species composition, caging experiments can
illuminate aspects of dragonfly natural history that are
difficult to address with purely descriptive studies. For
instance, previous studies of dragonfly assemblages in
the southeastern United States have suggested that
most odonates are univoltine (Paulson and Jenner 1971,
Benke and Benke 1975). In contrast, the metamorpho-
sis of four species (Peritheinis tenera, Pachydiplax
longipen nis, Eryth enis simnplicicollis, and Tran'meala-
cer(Ita) from fish exclosures in Carroll's Pond following
only 1-2 mo of development demonstrated these
species are facultatively multivoltine in this portion of
their geographic range. The great rapidity of odonate
development observed in this study emphasized the
importance of monitoring metamorphosing individu-
als, even during relatively short-term experiments.
Without such monitoring, significant differences among
treatments can effectively metamorphose and fly away,
while remaining undetected.
This experiment demonstrated that predation can be
an important regulatory mechanism during the larval
phase of the odonate complex life cycle. Fish greatly
reduce the number of larvae making the successful
transition between the aquatic larval phase and the
terrestrial/aerial adult phase of the life cycle. Such in-
teractions suggest the potential for novel competitive
interactions between fish and terrestrial species over
aquatic and terrestrial phases of a shared odonate re-
source. This admittedly speculative interaction de-
serves further study and is supported in part by other
observations. Certain birds can depend heavily on te-
neral odonates for food during the breeding season
(Orians 1980). A negative relation between reproduc-
tive success in some waterfowl and the presence of
fish is also correlated with reduced abundances of
shared macroinvertebrate prey where fish are present
(Eriksson 1979). Predators with complete habitat sep-
aration (aquatic vs. terrestrial) may conceivably com-
pete for different life history stages of the same prey
species, especially if prey abundance in both habitats
is tightly linked by metamorphosis and reproduction,
and if prey abundance in one habitat is limited by pre-
dation.
The degree of coupling between the abundance of
60 PETER J. MORIN Ecology. Vol. 65, No. 1

adult and larval odonates may be quite variable. Com- Johnson, D. M. 1973. Predation by damselfly naiads on
position of the aquatic larval phase of odonate assem- cladoceran populations: Fluctuating intensity. Ecology 54:
25 1-268.
blages probably reflects a differential "filtering" of eggs Johnson, D. M., and P. H. Crowley. 1980. Odonate "hide
and larvae of susceptible odonates by vertebrate pre- and seek"': Habitat-specific rules" Pages 569-579 in W.
dation (this study), predatory and competitive inter- Charles Kerfoot. editor. Evolution and ecology of zoo-
actions among larvae (Benke 1978, Benke et al. 1982, plankton communities. University Press of New England,
Crowley and Johnson 1982), and possible physiologi- Hanover, New Hampshire, USA.
Macan, T. T. 1977. The influence of predation on the com-
cal constraints (Corbet 1980). Composition of the adult position of freshwater animal communities. Biological Re-
phase of odonate assemblages probably reflects input views 52:45-70.
from the larval phase at that pond, migration of adults Morin. P. J. 1981. Predatory salamanders reverse the out-
from other breeding sites, and competitive and pred- come of competition among three species of anuran tad-
poles. Science 212:1284-1286.
atory interactions among adults. Integrated studies of Needham, J. G.. and M. J. Westfall, Jr. 1955. A manual of
both aquatic and larval phases of odonate assemblages the dragonflies of North America (Anisoptera). University
are needed to sort out the relative importance of these of California Press. Berkeley. California. USA.
potentially complex interactions in determining the Orians, G. H. 1980. Some adaptations of marsh-nesting
distribution and abundance of species with complex blackbirds. Princeton University Press, Princeton, New
Jersey, USA.
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ACKNOWLEDGMENTS
Paulson, D. R., and C. E. Jenner. 1971. Population struc-
This study would have been impossible without the co- ture in overwintering larval Odonata in North Carolina in
operation of John and Lucille Carroll, who own the pond. relation to adult flight season. Ecology 52:96-107.
Their patient tolerance of backyard ecology is greatly appre- Sadzikowski, M. R., and D. C. Wallace. 1976. A compar-
ciated. M. K. Morin, R. Harris, D. Campbell. A. Gotten, S. ison of the food habits of size classes of three sunfishes
Lewis, H. F. Nijhout, K. Clay. and H. M. Wilbur assisted (Lepomnis Inalrochirius Rafinesque, L. gibbosus (L innaeus)
in the field and/or commented on the manuscript. R. T. Paine and L. c'vaiuel//Is Rafinesque). American Midland Natural-
supplied an obscure reference, and comments by reviewers ist 95:220-225.
D. M. Johnson and A. C. Benke improved the manuscript. Southwood, T. R. E. 1978. Ecological methods. Second
Any errors of interpretation remain my own. edition. J. Wiley and Sons, New York, New York, USA.
During part of this study I was supported by National Sci- Thorp, J. H., and E. A. Bergey. 1981Ia. Field experiments
ence Foundation Grant DFB-791 1539 to H. M. Wilbur, and on responses of a freshwater. benthic macroinvertebrate
by a teaching assistantship in the Department of Zoology, community to vertebrate predators. Ecology 62:365-375.
Duke University. Computing funds were provided by the De- Thorp, J. H., and F. A. Bergey. 1981lb. Field experiments
partment of Zoology, Duke University. on interactions between vertebrate predators and larval
midges (Diptera: Chironomidae) in the littoral zone of a
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