Forest Ecology and Management 261 (2011) 391–399
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Forest Ecology and Management
journal homepage: www.elsevier.com/locate/foreco
Early-stage impacts of sika deer on structure and function of the soil microbial
food webs in a temperate forest: A large-scale experiment
Shigeru Niwa
a,∗
, Lucero Mariani
a
, Nobuhiro Kaneko
a
, Hiroaki Okada
b
, Kazunori Sakamoto
c
a
Soil Ecology Research Group, Graduate School of Environment and Information Sciences, Yokohama National University, 79-7 Tokiwadai, Hodogaya, Yokohama 240-8501, Japan
b
National Institute for Agro-Environmental Sciences, 3-1-3 Kan’nondai, Tsukuba 305-8604, Japan
c
Graduate School of Horticulture, Chiba University, 648 Matsudo, Chiba 271-8510, Japan
article info
Article history:
Received 1 July 2010
Received in revised form 20 October 2010
Accepted 20 October 2010
Available online 18 November 2010
Keywords:
Mammalian herbivore
Deer browsing
Dwarf bamboo
Soil nematode
Soil microorganism
Nitrogen mineralization
abstract
Modification of forest vegetation caused by an overabundance of mammalian herbivores has been
reported in temperate and subarctic regions all over the world. However, the indirect effects of these
herbivores on the structure and functions of soil decomposer systems are not fully understood, espe-
cially in temperate forests. We investigated the early effects of sika deer invasion on soil decomposer
systems in a Japanese temperate forest using two large-scale experimental enclosures with low and high
densities of deer (LD: 25 ha, 4 deer km
-2
; HD: 6.25 ha, 16 deer km
-2
) including control plots without deer
(WD). Three years after deer introduction the understory cover of dwarf bamboo (Sasa nipponica) declined
due to deer browsing in both enclosures. At the same time, measurements were made of the soil micro-
bial community, soil nematode community, soil nitrogen (N) mineralization rate, and carbon (C) and N
content in dwarf bamboo leaves. In LD, soil microbial biomass was lower from WD, probably due to the
decrease of fresh aboveground litter from dwarf bamboo. Surprisingly, there were no cascading effects
on total abundance of soil nematodes and soil total N mineralization potential which were unaffected by
deer in the LD treatment, while soil NH
4
+
-N content was lower and soil nematode community structure
was different (abundance of 4 families was higher and that of 3 families was lower, but the functional
structure was not different) from WD. Specifically, the responses to deer introduction varied between
microbes and nematodes, and the change of balance in the microbial food webs may have altered N min-
eralization processes. In contrast, in the HD treatment, all the variables measured were not significantly
different from those of WD treatment. Intensive browsing by deer may have cancelled out the effects of
the decrease in aboveground litter input on the soil decomposer systems through other pathways, such
as a transitory increase in belowground litter input caused by induced changes in allocation patterns of
bamboo. No changes in total N mineralization potential, leaf N, and composition of understory vegetation
in both enclosures indicated that deer introduction did not facilitate nor retard N cycling regardless of
deer density. This study showed that sika deer browsing can affect soil decomposer systems at an early
stage of invasion even at low density, which contrasts with previous studies on the subject. Linking our
findings of early-stage effects of deer on soil decomposer systems to longer-term dynamics of understory
vegetation and tree regeneration will be needed to evaluate the adequacy of deer management practices
with respect to the sustainability of soil nutrient supplies.
© 2010 Elsevier B.V. All rights reserved.
1. Introduction
Modification of forest vegetation caused by overabundance of
mammalian herbivores has been reported in temperate and sub-
arctic regions all over the world (Weisberg and Bugmann, 2003).
In temperate forests across Japan, Sika deer (Cervus nippon Tem-
∗
Corresponding author. Present address: Tomakomai Experimental Forest, Field
Science Center for Northern Biosphere, Hokkaido University, Takaoka, Tomakomai
053-0035, Japan. Tel.: +81 144 33 2171; fax: +81 144 33 2173.
E-mail address: sniwa@fsc.hokudai.ac.jp (S. Niwa).
minck) populations have been expanding for the past two decades,
and alterations of forest vegetation resulting from the heavy brows-
ing pressure of the deer have become a serious problem (Takatsuki,
2009). Understanding the impacts of mammalian herbivores on the
structure and functions of forest ecosystems is required as a basis
for the development of appropriate management systems of these
mammal populations.
Belowground systems play vital roles in terrestrial ecosystems,
such as organic matter decomposition and nutrient mineraliza-
tion, which enable sustainable primary production in aboveground
systems. Activities of large mammalian herbivores can alter the
structure and functions of soil decomposer systems resulting from
0378-1127/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2010.10.024