2084

NOTES

Control of seedling emergence by ground cover: a potential mechanism

involving seed predation
R. J. READER
Departttiet~tof Botany, Ur~iversit)~
of Guelph, Guelph, Ont., Canacic~NIG 2WI
Received November 12, 1990

READER, R. J. 1991. Control of seedling emergence by ground cover: a potential mechanism involving seed predation. Can.
Can. J. Bot. Downloaded from www.nrcresearchpress.com by MCGILL UNIVERSITY on 04/07/13

J. Bot. 69: 2084-2087.

The presence of ground cover (i.e., plant biomass and litter) could restrict seedling emergence by providing a habitat for
seed predators. To test this hypothesis, a field experiment was conducted with three old-field forbs (Daucus carota, Cerltaureu
tzigm. Tarmacurn offiiciinule). Ground cover was either removed or left in place and sown seeds of the three forbs were either
protected from predators or left unprotected. Where seeds were protected from predators, seedling emergence did not improve
when ground cover was removed. Where seeds were not protected from seed predators, seedling emergence did improve
significantly for all three species when ground cover was removed. Apparently, seed predators removed more unprotected
seeds where ground cover was present than absent, thereby reducing seedling emergence. These results are consistent with
the hypothesis that ground cover could restrict seedling emergence by providing a habitat for seed predators.
Key words: old field, seedling emergence, seed predation.

READER, R. J. 1991. Control of seedling emergence by ground cover: a potential mechanism involving seed predation. Can.
J. Bot. 69 : 2084-2087.
La prCsence de couverture sur le sol (i.e., biornasse vCgCtale et litiere pourrait rCduire I'emergence des plantules en
For personal use only.

fournissant un habitat pour les graminivores. Afin de vCrifier cette hypothese, l'auteur a effectuC une experience au champ
sur trois especes de vieilles prairies (Daucus carota, Centuurea nigru, Taravacrrtn officit~ale).La couverture de sol a CtC soit
CliminCe ou encore IaissCe en place et les graines des trois espkces de plantes ensemencees ont CtC soit protegees des gra-
minivores ou encore IaissCes sans protection. L i ou les graines ont CtC protCgCes des graminivores, I'emergence des plantules
ne s'est pas arnCliorCe lorsque le couvert du sol a CtC enleve. L i oh les graines n'ont pas CtC protCgCes des graminivores,
I'Cmergence des plantules s'est sensiblernent amCliorC chez les trois especes lorsque la couverture du sol a CtC CliminCe.
Apparemment les graminivores Climinent plus de graines non protCgCes 18 ou la couverture du sol est prCsente que la ou elle
est absente, rCduisant ainsi I'Cmergence des plantules. Ces rksultats concordent avec I'hypothese que la couverture du sol
pourrait rCduire 1'Cmergence des plantules en fournissant un habitat aux graminivores.
Mots clts : vieilles prairies, Cmergence des plantules, grarninivores.
[Traduit par la rCdaction]

Introduction
Environmental factors that restrict seedling emergence can
play an important role in determining plant abundance (Harper
1977). In old fields, pastures, and grasslands, dense ground
cover (i.e., plant biomass and litter) often restricts seedling
emergence. This has been demonstrated by removing ground
cover experimentally and showing that more seedlings emerge
in cleared plots than in plots with ground cover left intact (e.g.,
Sagar and Harper 1960; Putwain and Harper 1970; Gross and
Werner 1982; Reader and Buck 1986).
Three mechanisms have been suggested to explain why
ground cover restricts seedling emergence. First, ground cover
may inhibit seed germination by changing microclimatic con-
ditions (e.g., lowering the red to far red ratio of incoming light
or reducing the amplitude of daily temperature fluctuations)
(Rice 1985; Keizer et al. 1985, VanTooren 1990). Second,
ground cover may inhibit seed germination by changing soil
chemistry (e.g., adding allelochemicals to the soil) (McPherson
and Thompson 1972; Werner 1975). Third, ground cover may
act as a physical barrier to shoot extension by germinated seeds
(McPherson and Thompson 1972; Sydes and Grime 1981). I
suggest that there is a fourth potential mechanism by which
I
ground cover may restrict seedling emergence, namely, that
Printed In Canada 1 ImprinlC au Canada
ground
- cover provides a habitat for seed predators who reduce
seedling emergence by removing seeds.
Postdispersal seed predation can reduce both seed supply
and seedling emergence greatly in old fields, pastures, and
grasslands (e.g., Sarukhan 1974; Borchert and Jain 1978;
Grieg-Smith and Sagar 198 1). In one old field, ants and rodents
removed more seeds from experimental dishes set in undis-
turbed vegetation than from dishes set o n bare ground
(Mittelbach and Gross 1984). I tested the hypothesis that
removing ground cover experimentally improves seedling
emergence by reducing seed predation. Seeds of three old-field
forbs were either protected from seed predators or left unpro-
tected and ground cover was either removed o r left intact. T h e
u
number of seedlings emerging was then compared in the four
experimental treatments.
Materials and methods
Stuclj area and species
The experiment was conducted in an abandoned pasture located
aboul8 krn east of Guelph, Ontario (43"33'N, 80°10'W). The pasture
has not been grazed by livestock since 1969 or earlier (Thomas and
Dale 1974). The pasture consisls of a series of rolling hills, each up
to 10 m high. The experiment was set up at the bottom of hills where
ground cover was the densest (i.e., 585 2 49 glm'; Hogenbirk and
 NOTES
TABLE1. F-values and P-values (ANOVA) for the effects of removing ground cover
(RGC) and excluding predators (EP) on the seedling emergence of three forb species
RGC
S~ecies F P
D. carota 145.7 <0.0003
T. officir~ale 66.1 <0.001
C. nigrcl 8.1 <0.04
Reader 1989). Ground cover consisted mainly of forbs and grasses
. .. . that dominate the vegetation in the study area (Reader and Best 1989).
. ..
.....:....
,. Three forb species (Daucus car-ota L., Cer~ta~o-ea nigr-a L., Tllrux-
acurn offieinale Weber) were used in the experiment. All three species
Can. J. Bot. Downloaded from www.nrcresearchpress.com by MCGILL UNIVERSITY on 04/07/13
reproduce only from seed (Alex and Switzer 1976) so seed predation
would limit their potential abundance.
Seedlit~gernergerlce under field conditior~s
The effects of ground cover and seed predation on seedling emer-
gence by the three forbs were measured in an experiment that had
four treatments: (i) ground cover left intact and seed predators not
excluded (control); (ii) ground cover removed but seed predators not
excluded; (iii) ground cover left intact and seed predators excluded;
and (iv) ground cover removed and seed predators excluded. The four
treatments were set up at the bottom of each of five hills. The four
treatments were assigned randomly to four 1 x I m plots that were
set up side by side.
Six 10-cm diameter subplots were set up within each 1 x 1 m
plot. Subplots were arranged in a 3 x 2 grid, with subplots about
25 cm apart. Seeds of the three test species were sown in three of the
For personal use only.
six subplots (one species per subplot) and the other three subplots
were left unseeded. Fifty seeds per subplot were sown for C. nigrcl
and D. carota and 500 seeds per subplot were sown for T. oflcir~ale
because results of a preliminary study indicated that the number of
seedlings emerging from sown seeds was likely to be an order of
magnitude lower for T. oflcir~ule than for either C. nigra or D. cur-
ora. Sown seeds had been collected from plants in the study area and
seeds were dry stored at about 20°C for 1-4 months before they were
sown. Unseeded subplots were used to correct for possible variation
among subplots in the number of seedlings that emerged from the
seed bank. Both unseeded subplots and seeded subplots were assigned
randomly to species.
To exclude seed predators from a subplot, a plastic tube (10 cm
diameter x 8 cm long) was hammered 4 cm into the ground and the
open end and plus sides of the tube were covered with a piece of
. ,
fibreglass screen (1 mm mesh). The screen was held in place with a
rubber band. Ground cover was removed by hand from the entire
1 x 1 m plot. Rootstocks were also removed to minimize regrowth.
Seeds were sown evenly in subplots in mid-September and sub-
sequent seedling emergence (i.e., cotyledons expanded) was recorded
at weekly intervals until the following June. On each sampling date,
newly emerged seedlings were counted and removed using forceps to
avoid double counting. In subplots protected from seed predators,
each screen was removed for about 10 mln to count seedlings.
The number of seedlings that emerged from sown seeds was cal-
culated for each of the three forbs using the following equation:
where SE is the number of seedlings emerging per sown seed, E, is
the number of seedlings emerging in the subplot containing sown
seeds, E,,, is the number of seedlings emerging in the subplot without
sown seeds, and S is the number of seeds sown per subplot. An anal-
ysis of variance (ANOVA)for a randomized complete block design
was used to determine the statistical s~gnificanceof the effects of
excluding predators and removing ground cover on seedling emer-
gence within a block (i.e., hill).
EP RGC x EP
F P F P
821.8 <0.0001 147.7 <0.0003
101.6 <0.0005 32.6 <0.004
50.4 <0.002 12.4 <0.02
Effect of plastic tube and scree11 or1 seedlir~gemergence
The plastic tube and screen used to exclude seed predators may
have changed microclimatic conditions experienced by seeds. To
determine whether the tube and screen affected seedling emergence,
the following experiment was conducted simultaneously in the green-
house, where seed predators were absent. Seeds were added to
10-cm diameter pots (six pots per forb species) that contained a com-
mercially prepared potting medium (Promix BX, Les tourbieres pre-
mier, Quebec). Fifty seeds per pot were sown for C. nigra and for
D. carota and 500 seeds per pot were sown for T. officir7ale. Three
of the pots had plastic tubes and screens and the other three pots had
no tubes or screens. Pots were watered irregularly to simulate natural
wetting and drying of the substrate and seedling emergence was
recorded at weekly intervals for 6 months. The mean number of seed-
lings that emerged per sown seed was calculated for each species. A
t-test (Sokal and Rohlf 1981) was used to determine the statistical
significance of a difference in mean seedling emergence in pots with
tubes and screens versus pots without tubes or screens.
Results
Seedling emergence under field corlditions
More seedlings emerged where ground cover had been
removed than where ground cover was left intact when seeds
were not protected from predators (Fig. 1). However, with
seeds protected from predators, about equal numbers of seed-
lings emerged where ground cover had been removed and
where ground cover was left intact. Each of the three forbs
showed this response.
ANOVA results (Table 1) indicated that both removing ground
cover and excluding predators significantly (P < 0.05)
increased the emergence of seedlings of all three species. There
was also a significant interaction between the effects of remov-
ing ground cover and excluding predators. Removing ground
cover improved seedling emergence when predators were pres-
ent but not when predators were excluded.
Effect of plastic tube and screen on seedlitlg emergence
In the greenhouse, seedling emergence was largely unaf-
fected by the presence of the plastic tube and screen used to.
exclude seed predators in the field (Fig. 2). Only C. nigra
showed a slight increase in seedling emergence with the exclo-
sure present. However, results of the t-test indicated that the
exclosure did not improve seedling emergence significantly
for C. nigra (t = 1.3, df = 2, P > 0.2), D. carota (t = 0.2,
df = 2, P > 0.9), and T. oj'icinale (t = 1.3, df = 2, P > 0.2).
Discussion
for the three forbs the idea that
ground cover can restrict seedling emergence by providing a
habitat for seed predators. When seeds were protected
predators, seedling emergence did not improve with the
removal of ground cover. But when seeds were not protected
from seed predators, removing ground cover did improve seed-
 CAN. J . BOT.

Daucus carota Centaurea nigra

I .o 1 Daucus carota I .o 1 Centaurea *n
1 I I '1

0.01 , Taraxacurn officinale

Can. J. Bot. Downloaded from www.nrcresearchpress.com by MCGILL UNIVERSITY on 04/07/13

0
a,
a,
V) Taraxacurn officinale
.,-
0 0.2 1

FIG. 2. Comparison of mean ( 2 I SE) seedling emergence in the

FIG. 1. Comparison of mean ( ? I SE) seedling emergence with
ground cover left intact or removed when seeds of three forbs
(D. corotLl, T. o#i:citzule, and C . nigra) were and were not protected
from predators. B, ground cover intact, with predators; B, ground
For personal use only.
greenhouse without (0) or with (0)the plastic tube and screen used
to protect seeds of three forbs (D. carota, T. officinale, and C . tzigra)
from predators in the field experiment.
emergence by inhibiting seed germination or by preventing
shoot extension of germinated seeds. Additional studies are

needed to determine whether seed predation interacts with

cover removed, with predators: 0 . ground cover intact, without pre-
ground cover to control seedling emergence in habitats other
dators: 0.ground cover removed. without predators.
than old fields and abandoned pastures.
ling emergence. Apparently. seed predators removed more
unprotected seeds where ground cover was left intact than Acknowledgments
where ground cover was removed. This reduction in seed sup-
ply reduced seedling emergence more with ground cover pre- This research was supported by the Natural Sciences and
sent than with ground cover removed. Engineering Research Council of Canada. B . Beisner, J. Buck,
This interpretation of experin~entalresults is consistent with C . Clark, and P. Kelly kindly helped with the fieldwork. T w o
the observation by Mittelbach and Gross (1984) that ants and anonymous reviewers plus P. Krannitz, D. Larson, and A.
rodents removed more seeds from experimental dishes set in Maun provided constructive criticisms of an earlier version of
undisturbed vegetation than on bare ground in the old field the paper.
they studied. Ants are also important seed predators in the
abandoned pasture studied here. When ants were excluded ALEX,J. F.. and SWITZER, C. M . 1976. Ontario weeds. Ont. Minist.
from experimental dishes containing seeds. significantly fewer Agric. Food Publ. No. 505.
seeds disappeared than from dishes that were accessible to ants BORCHERT, M. I.. and J A I NS., K. 1978. The effect of rodent seed
(R. J . Reader and B . Beisner. unpublished data). In contrast, predation on four species of California annual grasses. Oecologia
excluding rodents and birds had little effect on the number of (Berlin), 33: 101-1 13.
seeds that disappeared from experimental dishes. GRIEG-SMITH, J. T., and SAGAR, R. G. 1981. Biological causes of
rarity in Car-litla vulgaris. It1 Biological aspects of rare plant con-
The plastic tube and screen could have improved seedling servation. Edited by H. Synge. John Wiley & Sons, New York.
emergence not only by excluding predators but also by creating pp. 389-400.
more favourable microclimatic conditions for seed germina- GROSS,K. L., and WERNER, P. A. 1982. Colonizing abilities of
tion. In the greenhouse, where seed predators were absent, "biennial" plant species in relation to ground cover: implications
seedling emergence did not improve significantly with the for their distributions in a successional sere. Ecology, 63: 921-
addition of the plastic tube and screen. Therefore, it is unlikely 93 1 .
that the plastic tube and screen improved seedling emergence HARPER, J. L. 1977. Population b~ologyof plants. Academic Press,
in the field by simply improving the microclimate for seed London.
germination. HOGENBIRK, J. C., and READER, R. J. 1989. Biotic versus abiotic
This is the first study where the effect of removing ground control of plant density: studies of Medicngo l~cp~clina L. on a
topographic gradient. J . Biogeogr. 16: 269-277.
cover on seedling emergence was examined with seeds pro- KEIZER. P. J., VANTOOREN, B. F., and DURING, H. J. 1985. Effects
tected from seed predators. Consequently, it is difficult to of bryophytes on seedling emergence and establishment of short-
judge whether seed predation was a confounding factor in other lived forbs in chalk grassland. J. Ecol. 73: 493-504.
studies. Since results of the present study suggest that the MCPHERSON, J. K . , and THOMPSON, G. L. 1972. Competitive and
removal of ground cover can reduce seed predation, it would allelopathic suppression of understory by Oklahoma oak forests.
be unwise to assume that ground cover must restrict seedling Bull. Torrey Bot. Club, 99: 293-300.
 NOTES 2087
MITTELBACH, G. G., and GROSS,K. L. 1984. Experimental studies
of seed predation in old-fields. Oecologia (Berlin), 65: 7- L3.
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m

PUTWAIN, P. D., and HARPER, J. L. 1970. Studies in the dynamics

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READER,R. J.. and BUCK,J . 1986. Topographic variation in the dovd
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abundance of Hieraciumfloribundutn: relative importance of dif- .'i
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RICE,K . J. 1985. Responses of Erodiilm to varying microsites: the c, '9+
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Erratum: Postglacial vegetation history of oak

savanna in southern Ontario1
J. M. Sz~rczAND G. M.MACDONALD
Departmetlt of Geography, McMaster University, Hatniltotl, Otzt.,
Catzada L8S 4 K l
(Ref. Can. J. Bot. 69: 1507-1519. 1991.)
O n p. 1513, the caption for Fig. 6 was inadvertently omit-
ted. This figure with its caption is reprinted in the right-hand
column.

> -
-7-. -?-
a .
. E
2.? 0
L ln
'Received at NRC October 7, 1991. 5