Nektonic macroinvertebrates in a wetland pond (Crescent

Pond, Delta Marsh, Manitoba)

Brenda J. Hann
Department of Zoology, University of Manitoba
Winnipeg, Manitoba R3T 2N2

Introduction strength of invertebrate-waterfowl associations for

breeding and feeding (Murkin and Kadlec 1986) and
Temporal and spatial variability in structure and invertebrate-macrophyte-waterfowl associations (Lillie
composition of the macroinvertebrate community and Evrard 1994).
throughout the open-water season in wetland water Invertebrates constitute vital elements in the marsh
bodies has been investigated in few studies (Laurie 1942; food web in coastal wetlands as has been shown in recent
Macan 1949, 1965, 1966). The paucity of such studies studies of energy and biomass transfer using stable
is attributable at least in part to difficulties associated isotope techniques (Neill and Cornwell 1992; Keough
with sampling in a heterogeneous habitat, ranging from et al. 1996). There is often considerable energy and
open water to dense submersed macrophytes to biomass transfer between the aquatic and terrestrial
flocculent sediments, and in part to difficulties with habitats as many macroinvertebrates spend portions of
taxonomic identifications of an extremely diverse fauna their life cycle in each niche (Merritt and Cummins
as well as lack of information on life histories of many 1984).
species. Thus, priorities in wetland ecology can be The purpose of this study was to describe spatial
clearly delineated: (1) taxonomy–the need to know and temporal patterns of nektonic macroinvertebrate
species of invertebrates, diversity, life histories, abundance and community composition in a northern
generation times, etc. and (2) functional ecology–the prairie marsh pond subject to varying amounts of
need to know diet, nutrient and energy cycling, food predation pressure. In 1986, fish were present whereas
web dynamics, use of fine particulate organic matter in 1988, no fish were present until late summer. This
(FPOM) and coarse particulate organic matter (CPOM), report provides a preliminary examination of the
both occurring as detritus derived from macrophytes, nektonic community that will be followed by a
habitat complexity, and trophic linkages (Murkin and comprehensive, species-specific treatment of abundance
Kadlec 1986; Murkin and Batt 1987; Murkin and patterns.
Wrubleski 1988; Krieger 1992).
Diversity and productivity of aquatic invertebrates Methods
in wetlands is remarkably high, yet little studied (Batzer
and Wissinger 1996). In wetlands, interannual vagaries Crescent Pond is a small littoral pond within the
of water depth (flooding to drought conditions) and Delta Marsh, and a remnant oxbow of the Assiniboine
presence or absence of fish interact with seasonal River which formerly flowed northward into Lake
subtleties of predator-prey relations and population Manitoba from 4,500 to 2,200 years BP (Teller and Last
dynamics (demographic stochastisticity) to accentuate 1981). Although the pond has probably existed for 2,000
the complexity of invertebrate communities. to 2,500 years, Goldsborough (1987) has documented
Macroinvertebrates are important to waterfowl in the progressive in-filling and reduction in open-water
northern prairie marshes (Murkin and Kadlec 1986). The area of the pond over the last 40 years. Water level
densities and biomass of spring aquatic regulation of the adjacent Lake Manitoba which began
macroinvertebrates, in particular, influence waterfowl in 1960-61 may have contributed to the process, but
densities in marsh habitats. Swanson and Meyer (1977) Burchill (1987) suggested that Lake Manitoba water
found that waterfowl feed extensively on invertebrates levels would have been relatively stable as a
during the spring to meet the high protein requirements consequence of a decrease in precipitation variability
of breeding and egg laying. Waterfowl use fish presence/ even in the absence of the Fairford control structure.
absence as cues to estimate invertebrate densities in The dominant submersed vegetation in Crescent
selecting breeding sites (Mallory et al. 1994). Pond includes Ceratophyllum demersum, Myriophyllum
Differential responses among waterfowl species in spicatum, Potamogeton zosteriformis, P. pectinatus and

68 UFS (Delta Marsh) Annual Report, Vol. 30, 1995

Hann Nektonic wetland macroinvertebrates

the macroalga Chara sp. The pond is surrounded by a determined at open water (3, 5, 7, 8, 12, 17, 18) and
marginal band of Typha, with Phragmites encircling this marginal (4, 6, 9, 19) sites by first averaging values for
zone. horizontal and vertical traps at each site, then over all
In 1986, macroinvertebrate sampling took place sites in each group. Water temperatures were recorded
every three to four days (23 May - 5 October), using 1- weekly at deep and shallow sites using maximum-
L glass activity traps (modified from Murkin et al. 1983) minimum thermometers.
at four locations (N, E, S, M) in Crescent Pond (Fig. 1). Macroinvertebrates were identified using Merritt and
A total of six traps were placed per location (3 stakes, 2 Cummins (1984), Pennak (1978), and Edmondson
traps/stake) with stakes located approximately 2 meters (1959).
apart outside the marginal Typha band in the open water
area of the pond. Activity traps were attached just below Results
the water surface by hooks to wooden stakes. All traps
were set with the opening of the funnel facing Water temperature (maximum and minimum)
downwards toward the sediment-water interface in the showed consistent patterns of variation over the summer
evening and retrieved in the morning of the following months and varied little among sites in 1986 (Fig. 2).
day. Contents of the traps at each location were Mean maximum temperature was 22.5°C; mean
concentrated into a 250 mL polyethylene bottle with 10 minimum temperature was 18.6°C. Divergence between
mL of ethanol as a narcotizing agent. Mean abundances maxima and minima increased toward the end of the
per trap were determined at each of the sites, then values summer. Water depths declined substantially over the
for three marginal sites (N, E, S) were averaged and summer months and isopleths are shown for late May
standard errors calculated. As there was only one open (Fig. 3a) and August (Fig. 3b).
water site (M), standard error could not be determined. Water chemistry parameters showed a consistent
Water temperature was recorded twice per week pattern over the season (Table 1, Figs. 4, 5). Specific
using maximum-minimum thermometers at each site. conductance increased from 1,140 to almost 1,800
Water depth was determined weekly at both N and M µS·cm-1, and alkalinity from 4,760 to over 7,000 µeq·L-1
sites using a calibrated depth stake. Water samples were over the open water season. Similarly, calcium,
collected weekly from the M site for analysis by the magnesium, potassium, and sodium ions showed gradual
analytical chemistry laboratory at the Freshwater increases in concentration (mg·L -1) from May to
Institute, Department of Fisheries and Oceans, October. The average pH was 8.34.
Winnipeg, MB. Dytiscid larvae showed peak densities in May at the
In 1988, macroinvertebrate sampling occurred once marginal sites and throughout June at the open water
per week (31 May - 23August). Sampling sites were site (Fig. 6a). These larvae are actively swimming
established at several sites along transects (W-E, N-S) predators that seize their prey effectively with large
in the open water region of Crescent Pond (Fig. 1).
Activity traps were attached to wooden stakes as in 1986,
but pairs of traps were set with funnels directed 24 May 1988
horizontally and vertically. Traps were set in the evening 60
50

for approximately 12 hours. At each site, contents of 60 70

80 90

traps placed horizontally and vertically were kept 100

80
70

separate for analysis. Mean abundances per trap were

19 88
x 19 86
sh allow water site s xxx
op en wat er sites 8 August 1988
4 xxx 20 10
18 19 30
6
5 xxx 40
7 50 60 20
17 12
30

3 50 40
8 xxx
9

Figure 1. Crescent Pond sampling locations in 1986 and Figure 2. Water depth isopleths for Crescent Pond in
1988. 1988.

UFS (Delta Marsh) Annual Report, Vol. 30, 1995 69

Nektonic wetland macroinvertebrates Hann
Table 1. Water chemistry for Crescent Pond, 1986.

20 27 4 12 20 2 7 21 5 14 20 27 2 15 22 7 20
May May June June June July July July Aug Aug Aug Aug Sept Sept Sept Oct Oct

TDN (µg/L) 1180 1410 1300 1210 1330 1520 1630 910 1270 1130 1450 1140 1650 1280 1590 710 1430
Na (mg/L) 130 140 161 170 171 159 165 151 165 168 179 182 191 205 200 198 202
K (mg/L) 14.3 15.0 17.2 17.8 19.1 11.5 22.9 22.9 23.3 22.5 21.7 21.8 23.2 24.0 24.2 24.6 24.6
Ca (mg/L) 42.0 43.0 49.0 48.0 46.0 44.6 48.4 43.7 50.6 52.7 48.7 56.0 59.9 59.9 57.9 52.0 50.5
Mg (mg/L) 45.0 48.3 54.7 55.3 55.3 57.9 60.0 57.9 57.4 61.0 63.8 70.0 72.7 75.9 75.4 72.4 74.6
Fe (mg/L) 0.04 0.04 0.04 0.04 0.04 0.08 0.08 0.04 0.08 0.08 0.04 0.04 0.04 0.04 0.04 0.04 0.04
Mn (mg/L) 0.01 0.01 0.06 0.10 0.01 0.01 0.06 0.06 0.28 0.11 0.03 0.13 0.21 0.03 0.01 0.01 0.01
Cl (mg/L) 188 198 307 257 260 250 260 240 270 300 320 340 320 360 360 300 300
SO4 (mg/L) 79.2 79.2 99 99 110 100 110 100 110 120 140 140 120 160 160 120 120
SuspN (µg/L) 150 209 88 216 307 329 632 292 188 178 111 126 469 461 883 898
SuspC (µg/L) 1040 1610 576 1740 1710 1710 5640 2270 1330 1230 820 1090 3710 2940 5490 6420
pH 8.06 8.53 8.44 8.25 8.32 8.42 8.30 8.35 8.13 8.43 8.31 8.47 8.24 8.19 8.36 8.60 8.44
Cond@25°C (µS/cm)1140 1200 1350 1400 1440 1460 1500 1410 1510 1550 1580 1640 1710 1780 1770 1770 1790
IC (µmole/L) 5200 5800 6400 5250 5300 5100 5600 6450 6600 5350 5500 5600 6400 7530 6660 5730 5820
OC (µmole/L) 1700 3500 2900 2400 2200 2150 2300 2450 2000 3000 1700 1600 2150 2520 2310 2480 2610
Alk (µeq/L) 4760 4890 5460 5452 5526 5770 5874 5676 6108 6404 6528 6810 7052 7122 6994 6984 7000
Org.Acid (µeq/L) 813 777 311 459 1 1 1 1 1 1 630 936
anions (µeq/L 12087 13093 17153 14554 14924 15071 15995 15621 16904 16338 17438 18144 17900 20968 20157 17400 17749
cations (µeq/L) 11817 12591 14386 14792 14769 14197 15112 14096 15018 15529 16017 17025 17869 18761 18408 17790 18070

30
220 80
Date vs Na (mg/L)
28 Crescent Pond (1986) Date vs K (mg/L) 70
Date vs Ca (mg/L)
200
Date vs Mg (mg/L)
26 60

Ca, Mg, K (mg/L)

180
Na (mg/L)

24 50
Temperature ( C)

40
22 160

30
20
140
20
18
120 10
20-May-86
27-May-86

4-Jun-86

12-Jun-86
20-Jun-86

2-Jul-86
7-Jul-86

21-Jul-86
5-Aug-86

14-Aug-86
20-Aug-86

27-Aug-86

2-Sep-86
15-Sep-86

22-Sep-86
7-Oct-86

20-Oct-86
16

14
Sample date

12
Figure 5. Concentrations of major ions in Crescent Pond
10 in 1986.
23/5

30/5

6/6

13/6

20/6

27/6

4/7

11/7

18/7

25/7

1/8

8/8

15/8

Date
mandibles. Dytiscid larvae typically seek moist soil in
Figure 3. Mean maximum and minimum water
which to pupate so it may be that the larvae at the
temperatures in Crescent Pond in 1986.
marginal sites in May were looking for pupation sites
1900 7500
and then appeared again in July as adults. Dytiscid adults
1800
7000 were present in highest numbers in early July in marginal
1700
sites (Fig. 6b). Several other Coleoptera adults were
Conductivity (µS/cm)

6500
present throughout the season (e.g., Acilius, Agabus,
Alkalinity (µeq/L)

1600

1500 Alkalinity 6000 Hydaticus, Haliplus, Hydrobius, Hydrochara) (Fig. 6c).

1400
Ephemeropteran naiads (e.g., Caenis sp.) were most
Conductivity
5500
abundant from May to early July at both marginal and
1300

5000
open water sites (Fig. 6d). Univoltine taxa typically have
1200
a peak in adult emergence in June and early July. Naiads
1100 4500 remained present throughout the season, showing a
20-May-86

27-May-86
4-Jun-86

12-Jun-86
20-Jun-86

2-Jul-86

7-Jul-86
21-Jul-86

5-Aug-86
14-Aug-86

20-Aug-86
27-Aug-86

2-Sep-86
15-Sep-86

22-Sep-86
7-Oct-86

20-Oct-86

slight increase in abundance again (e.g., Cloeon sp.) in

late August and September. Bivoltine taxa have one
Figure 4. Specific conductance and alkalinity in Crescent summer generation, then the naiads that are produced
Pond in 1986. will overwinter. Emergence patterns can vary with

70 UFS (Delta Marsh) Annual Report, Vol. 30, 1995

 Hann Nektonic wetland macroinvertebrates
1.2 120

open water
open water Typha interface
1.0 Typha interface 100

0.8 80
E
Abundance (#/trap)

Abundance (#/trap)
A
0.6 60

0.4 40

0.2 20

0.0 0
May June July Aug. Sept. Oct. May June July Aug. Sept. Oct.

14 40

open water
12 Typha interface open water
Typha interface

30
10
Abundance (#/trap)

Abundance (#/trap)
B F
8

20
6

4
10

0 0
May June July Aug. Sept. Oct. May June July Aug. Sept. Oct.

1.6 1.0

open water open water

Typha interface
Typha interface
0.8
1.2

G
Abundance (#/trap)

Abundance (#/trap)

C 0.6

0.8

0.4

0.4
0.2

0.0 0.0
May June July Aug. Sept. Oct. May June July Aug. Sept. Oct.

2.0 1.6

open water open water

Typha interface Typha interface

1.5 1.2

D H
Abundance (#/trap)

Abundance (#/trap)

1.0 0.8

0.5 0.4

0.0 0.0
May June July Aug. Sept. Oct. May June July Aug. Sept. Oct.

Figure 6. Seasonal abundance (numbers per trap) of macroinvertebrates in Crescent Pond in 1986. A. Dytiscidae
(Coleoptera) larvae. B. Dytiscidae (Coleoptera) adults. C. other Coleoptera adults. D. Ephemeroptera naiads. E.
Corixidae nymphs. F. Corixidae adults. G. Notonectidae nymphs and adults. H. Anisoptera (Odonata) naiads.

UFS (Delta Marsh) Annual Report, Vol. 30, 1995 71

 Nektonic wetland macroinvertebrates Hann
1.6 0.35

open water open water

Typha interface 0.30 Typha interface

1.2
0.25 L
I

Abundance (#/trap)
Abundance (#/trap)

0.20

0.8
0.15

0.10
0.4

0.05

0.0 0.00
May June July Aug. Sept. Oct.
May June July Aug. Sept. Oct.

3.0 10

open water
Typha interface open water
2.5 Typha interface
8

J
2.0 M

Abundance (#/trap)
Abundance (#/trap)

1.5

4
1.0

2
0.5

0.0 0
May June July Aug. Sept. Oct. May June July Aug. Sept. Oct.

12

open water
Typha interface
10

8
K
Figure 6 (continued) I. Zygoptera (Odonata) naiads. J.
Abundance (#/trap)

Hyallela azteca (Amphipoda). K. Water mites (Acari).

6
L. Hirudinea. M. Fish.
4

0
May June July Aug. Sept. Oct.

abundance, locality, and from year to year within the 6h). Adults typically emerge in August and September,
same species (Brittain 1982). lay eggs, then either eggs or naiads overwinter.
Corixid nymphs peaked (e.g., Hesperocorixa) in late Damselfly naiads (Zygoptera) (e.g., Coenagrion,
June to early July, then declined, and showed a second Enallagma, Lestes) peaked in late August to October
more substantial peak in abundance from mid-August (Fig. 6i) and probably overwinter as late instar naiads,
throughout September, particularly in the open water emerge in spring, lay eggs, then mature over the summer
(Fig. 6e). Corixid adults showed the highest abundance and early autumn. Emergence periods differ between
at marginal sites from late August to October (Fig. 6f). species and for the same species between years
Notonectids, which lurk at the water surface awaiting (Kormondy and Gower 1965).
prey, peaked in abundance in late August and September Amphipods (Hyalella azteca), both pre-reproductive
and were more frequently trapped at marginal sites (Fig. and reproductive adults, were present early in the spring
6g). and probably overwintered as adults (Fig. 6j). These
Odonates, voracious “sit-and-wait” predators, were released their young and die, so large individuals are
present throughout the season among vegetation, gone from the population by May. Juveniles produced
although dragonfly naiads (Anisoptera) (e.g., Aeschna) during spring reproduction are steadily recruited into
were most abundant from May to early August (Fig. the population, mature as young adults in July and

72 UFS (Delta Marsh) Annual Report, Vol. 30, 1995

Hann Nektonic wetland macroinvertebrates

7 6

open water open water

margin margin
6
5

A E
5
4

Abundance (#/trap)
Abundance (#/trap)

3
3

2
2

1
1

0 0
June July Aug. June July Aug.

7 2.0

open water open water

6 margin margin

5 B 1.5 F
Abundance (#/trap)

Abundance (#/trap)
4

1.0
3

2
0.5

0 0.0
June July Aug. June July Aug.

0.8 0.8

open water
0.7 open water
margin
margin

0.6 G
0.6
C
Abundance (#/trap)

Abundance (#/trap)

0.5

0.4 0.4

0.3

0.2 0.2

0.1

0.0 0.0
June July Aug.
June July Aug.

open water
5 margin
Figure 7. Seasonal abundance (numbers per trap) of
D macroinvertebrates in Crescent Pond in 1988. A.
4
Dytiscidae (Coleoptera) larvae. B. Dytiscidae
Abundance (#/trap)

(Coleoptera) adults. D. other Coleoptera adults. D.

3
Ephemeroptera naiads. E. Corixidae nymphs. F.
Corixidae adults. G. Notonectidae nymphs and adults.
2

0
June July Aug.

UFS (Delta Marsh) Annual Report, Vol. 30, 1995 73

Nektonic wetland macroinvertebrates Hann

0.4 10

open water
open water margin
margin
8
0.3
H K

Abundance (#/trap)
Abundance (#/trap)

0.2

0.1
2

0.0 0
June July Aug. June July Aug.

0.5 2.5

open water open water

margin margin
0.4 2.0

I L
Abundance (#/trap)

Abundance (#/trap)
0.3 1.5

0.2 1.0

0.1 0.5

0.0 0.0
June July Aug.
June July Aug.

350 250

open water
margin open water
300
margin
200

250 J M
Abundance (#/trap)
Abundance (#/trap)

150
200

150
100

100

50
50

0 0
June July Aug. June July Aug.

Figure 7 (continued) H. Anisoptera (Odonata) naiads. I. Zygoptera (Odonata) naiads. J. Hyallela azteca (Amphipoda).
K. Water mites (Acari). L. Hirudinea. M. Conchostraca.

August, and produce a second generation of young in this study, include Lymnaea stagnalis, Physa gyrina,
which overwinter as juveniles. Similar life history Stagnicola elodes, Gyraulus circumstratus, Helisoma
patterns have been observed in West Blue Lake, MB sp.
(Biette 1969) and in a northern Ontario lake (Lindeman In 1986, Pimephales promelas (fathead minnows)
and Momot 1983). Water mites (Acari) were consistently and Culaea inconstans (5-spined stickleback) were
abundant at all sites throughout the season (Fig. 6k). present throughout the season in the open water areas,
Leeches (Hirudinea) (e.g., Erpobdella punctata, with peak abundance in late June and early July, then
Glossiphonia complanata) were present throughout the again in late September and October (Fig. 6m).
season (Fig. 6l). Mollusca, not quantitatively sampled

74 UFS (Delta Marsh) Annual Report, Vol. 30, 1995

Hann Nektonic wetland macroinvertebrates

great blue heron,

Dytiscid larvae showed the same pattern of night hero n

abundance as in 1986, with peak numbers from May to

br ook sticklebacks)
(fathead minnows,
early July in open water, with no discernible difference

Am bystoma
passerines
waterfowl,
Verteb rate

tigrinum
predators

fish
between vertical and horizontal traps (Fig. 7a). Dytiscid
adults peaked in late May and early June at marginal
sites, but substantial numbers of adults were found

Notonectidae
(water mites)

chironom ids
Mesostoma
(flatworms)

Trichoptera
Ch aoborus

Dytiscidae
Corixidae
Hirudinia

Tanypod
Invertebrate

Hydra

Acar i
predators
throughout the season (Fig. 7b). Very few dytiscid adults
were trapped in the horizontally placed traps. Small
numbers of other coleopterans (larvae and adults) were

Ephemeroptera
Chironomidae /
Conchostraca

Entomostraca
(Gastropoda)

Oligochaetes
Lepidoptera
Primary

Amphipoda
Ostracoda
Copepoda
Cladocera
Mollusca
tadpoles

Rotifera
Diptera
consumers
found throughout the season (Fig. 7c). (herbiv ores /
detritivores)
Corixid nymphs were present in low abundance
throughout the season, showing a late summer peak in Primary
producers macrophytes epiphyton metaphyt on phytoplankton epipelon
marginal areas as seen in 1986 (Fig. 7d). Corixid adults
were rare in 1988 (Fig. 7e) in comparison with 1986.
Figure 8. Trophic level diagram for Crescent Pond.
They peaked in late June and August, especially in
marginal areas. Notonectids (Fig. 7f) and odonates (both
dragonflies and damselflies) (Fig. 7g,h) were relatively herbivore-detritivore) occurred at highest densities,
sparse in 1988. Notonectids showed no clear seasonal biomass and species richness in midsummer in open
or spatial pattern of distribution. Damselflies were water sites with dense submersed vegetation at normal
generally more abundant in June in open water. water depths. However, in a preliminary reconnaissance
Ephemeropteran naiads peaked in mid-June, and of Crescent Pond benthos and phytomacrofauna, Wright
increased again in late August (Fig. 7i), as in 1986. (1969) found higher densities of crustaceans (primarily
Amphipods peaked in August, and virtually all were amphipods) and dipterans (chironomids) at open water
trapped in horizontally placed traps (Fig. 7j). Water mites sites in spring and fall and considerable variability in
(Acari) were abundant from May to August, with invertebrate abundance existed between sites dominated
considerably higher numbers retrieved from the by different macrophytes. Most taxa in Crescent Pond
vertically placed traps in open water (Fig. 7k). Leeches in our study also exhibited either early or late summer
(Hirudinea) were present in numbers comparable to peaks in abundance, rarely peaks at both time periods.
1986 (Fig. 7l) and peaked in early July and mid-August However, densities in the two studies cannot be directly
at marginal sites. Conchostraca were remarkably compared as a consequence of different sampling
abundant in the absence of fish predators, especially methods. Wright (1969) used Ekman dredge samplers,
from May to early July, and most were trapped in the thereby emphasizing the benthic invertebrates, whereas
vertically placed traps (Fig. 7m) in open water. In 1988, Murkin traps which accentuate the water column
no fish were collected until late in August when a few organisms (nekton) were used in our study. In addition,
fathead minnows were found. Water temperatures were the higher degree of taxonomic resolution employed in
similar to those recorded in 1986. our study necessarily leads to finer subdivision of
A preliminary food web diagram has been assembled temporal patterns which can be discerned among taxa.
using available information (Fig. 8) but dietary In an attempt to generalize spatial distribution
interactions have not yet been fully investigated. patterns, Murkin et al. (1992) showed that nekton
densities (total nekton, cladocerans, corixids, ostracods)
Discussion at the emergent macrophyte-open water interface were
consistently lower than densities in open water subjected
The macroinvertebrate taxa found in Crescent Pond to fluctuating water levels. However, when spatial
during this study represent a large subset of the regional patterns of individual taxa were examined in our study,
faunal pool. The macroinvertebrate fauna is dominated certain groups showed consistent preference for
by larvae of insects, whereas the microinvertebrate fauna marginal areas, while others were generally found in
consists overwhelmingly of Entomostrata, including open water, and yet others showed no preference.
Cladocera, Copepoda, and Rotifera (Zrum and Hann Several taxa, for example, dytiscid larvae,
1995). Fish eat many of these before they leave the water ephemeropteran larvae, amphipods, and conchostracans,
or as they are about to do so, and therefore play an occurred in markedly lower abundance in the presence
integral role in influencing the translocation of insect of fish in 1986. Of these, only dytiscid larvae are
biomass from aquatic to terrestrial environment. themselves predators, but all are typically consumed
Murkin et al. (1991) documented that total readily by planktivorous fish. Macan (1966) similarly
invertebrates and functional groups (predator-parasite, found that coleopteran larvae were decimated after the

UFS (Delta Marsh) Annual Report, Vol. 30, 1995 75

Nektonic wetland macroinvertebrates Hann

introduction of fish into a pond. Further detailed Kneib, R.T. 1984. Patterns of invertebrate distribution
examination of effects of fish predation on the and abundance in the intertidal salt marsh: causes
macroinvertebrate fauna awaits their identification to and questions. Estuaries 7: 392-412.
species. Kormondy, E.J. and Gower, J.L. 1965. Life history
Several groups of macroinvertebrates are variations in an association of Odonata. Ecology
underrepresented in this study as a consequence of the 46: 882-886.
sampling methods employed which relied on active Krieger, K.A. 1992. The ecology of invertebrates in
swimming behaviour of the individual Great Lakes coastal wetlands: current knowledge
macroinvertebrate organism. Surface swimmers (e.g., and research needs. J. Great Lakes Res. 18: 634-
Gyrinidae, Velidae) and those taxa in which individuals 650.
are relatively immobile or firmly attached to substrata Laurie, E.M.O. 1942. The fauna of an upland pond and
(e.g., case-building trichopterans, flatworms, its inflowing stream at Ystumtuen, North
phytophilous chironomids, snails, bryozoans, Hydra, Cardiganshire, Wales. J. Anim. Ecol. 11: 165-181.
and leeches) were not accurately quantified. Lillie, R.A. and J.O. Evrard. 1994. Influence of
macroinvertebrates and macrophytes on waterfowl
Acknowledgments utilization of wetlands in the Prairie Pothole Region
of northwestern Wisconsin. Hydrobiologia 279/280:
I thank Alvin Dyck and Brian Grantham for 235-246.
assistance with sampling, field station staff for logistic Lindeman, D.M. and Momot, W.T. 1983. Production of
support, and the Portage Country Club for permission the amphipod Hyalella azteca in a northern Ontario
to conduct research on their property. The study was lake. Can. J. Zool. 61: 2051-2059.
partially funded by an NSERC operating grant to BJH Macan. T.T. 1949. Survey of a moorland fishpond. J.
and CareerStart funding to AD. Anim. Ecol. 18: 160-188.
Macan, T.T. 1965. The fauna in the vegetation of a
References moorland fishpond. Arch. Hydrobiol. 61: 273-310.
Macan, T.T. 1966. The influence of predation on the
Batzer, D.P. and Wissinger, S.A. 1996. Ecology of insect fauna of a moorland fishpond. Arch. Hydrobiol. 61:
communities in nontital wetlands. Ann. Rev. Ecol. 432-452.
Syst. 41: 75-100. Mallory, M.L., Blancher, P.J., Weatherhead, P.J. and
Bennett, D.V. and Streams, F.A. 1986. Effects of McNicol, D.K. 1994. Presence or absence of fish
vegetation on Notonecta (Hemiptera) distribution as a cue to macroinvertebrate abundance in boreal
in ponds with and without fish. Oikos 46: 62-69. wetlands. Hydrobiologia 279/280: 345-351.
Biette, R.M. 1969. Life history and habitat differences Merritt, R.W. and Cummins, K.W. 1984. An introduction
between Gammarus and Hyalella azteca in West to the aquatic insects of North America. Kendall/
Blue Lake, MB. M.Sc. Thesis, University of Hunt Publ. Co., Dubuque, Iowa.
Manitoba. Murkin, H.R. 1989. The basis for food chains in prairie
Brittain, J.E. 1982. The biology of mayflies. Ann. Rev. wetlands. IN Northern Prairie Wetlands, ed. A.G.
Entomol. 27: 119-147. van der Valk, pp. 316-338. Iowa State Univ. Press,
Burchill, C.A. 1987. Would Lake Manitoba have Ames, Iowa.
stabilized naturally? University Field Station (Delta Murkin, H.R. and Batt, B.D.J. 1987. The interactions
Marsh) Annual Report 22: 32-36. of vertebrates and invertebrates in peatlands and
Driver, E.A. 1977. Chironomid communities in small marshes. IN Aquatic Insects of Peatlands and
prairie ponds: some characteristics and controls. Marshes of Canada. pp. 15-30. Mem. Entomol. Soc.
Freshw. Biol. 7: 121-133. Can. 140: 1-174.
Edmondson, W.T. 1959. Freshwater Biology. 2nd Murkin, H.R., Abbott, P.G. and Kadlec, J.A. 1983. A
Edition. John Wiley and Sons, New York, NY. comparison of activity traps and sweep nets for
Goldsborough, L.G. 1987. Ontogeny of a small marsh sampling nektonic invertebrates in wetlands.
pond: revisited. University Field Station (Delta Freshw. Invert. Biol. 2: 99-106.
Marsh) Annual Report 22: 37-39. Murkin, H.R. and Kadlec, J.A. 1986. Responses of
Keough, J.R., Sierszen, M.E. and Hagley, C.A. 1996. benthic macroinvertebrates to prolonged flooding
Analysis of a Lake Superior coastal food web with of marsh habitat. Can. J. Zool. 64: 65-72.
stable isotope techniques. Limnol. Oceanogr. 41: Murkin, H.R. and Wrubleski, D.A. 1988. Aquatic
136-146. invertebrates of freshwater wetlands: function and
ecology. Ch. 20, pp. 239-249. In: Hook, D.D. et al.

76 UFS (Delta Marsh) Annual Report, Vol. 30, 1995

Hann Nektonic wetland macroinvertebrates

(eds.), The Ecology and Management of Wetlands. Reynolds, J.C. and Scudder, G.G.E. 1987. Experimental
Vol. 1. Ecology of Wetlands. Croon Helm, London. evidence of the fundamental feeding niche in
Murkin, H. R., Kadlec, J.A. and Murkin, E.J. 1991. Cenocorixa (Hemiptera:Corixidae). Can. J. Zool.
Effects of prolonged flooding on nektonic 65: 967-973.
invertebrates in small diked marshes. Can. J. Fish. Robert, L.L., Jr. and Matta, J.F. 1984. Aquatic
Aq. Sci. 48: 2355-2364. macroinvertebrates in an irregularly flooded salt
Murkin, E.J., Murkin, H.R. and Titman, R.D. 1992. marsh: diversity and seasonal variation. Env.
Nektonic invertebrate abundance and distribution Entomol. 13: 1097-1104.
at the emergent vegetation-open water interface in Swanson, G.A. and Meyer, M.I. 1977. Impact of
the Delta Marsh, Manitoba, Canada. Wetlands 12: fluctuating water levels on feeding ecology of
45-52. breeding blue-winged teal. J. Wildl. Mgmt. 41: 426-
Neill, C. and Cornwell, J.C. 1992. Stable carbon, 433.
nitrogen, and sulfur isotopes in a prairie marsh food Teller, J.T. and Last, W.M. 1981. Late Quaternary history
web. Wetlands 12: 217-224. of Lake Manitoba, Canada. Quaternary Research
Nelson, J.W. and Kadlec, J.A. 1984. A conceptual 16: 97-116.
approach to relating habitat structure and Wright, J. 1969. A quantitative survey of the benthos
macroinvertebrate production in freshwater and phytomacrofauna of an isolated body of water
wetlands. Trans. North Am. Wildl. Nat. Res. Conf. in the Delta Marsh. University Field Station (Delta
49: 262-270. Marsh) Annual Report 4: 65-70.
Pennak, R.W. 1978. Freshwater Invertebrates of the Zrum, L. and Hann, B.J. 1995. The impact of
United States. John Wiley and Sons, New York, NY. invertebrate and vertebrate predation on littoral
Rabe, F.W. and Gibson, F. 1984. The effect of zooplankton in a wetland ecosystem. University
macrophyte removal on the distribution of selected Field Station (Delta Marsh) Annual Report 29: 132-
invertebrates in a littoral zone. J. Freshw. Ecol. 2: 144.
359-371.

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