Taxa–area relationship and neutral dynamics influence
the diversity of fungal communities on senesced
tree leaves
Larry M. Feinstein* and Christopher B. Blackwood
Department of Biological Sciences, Kent State
University, Kent, Ohio, USA
Summary
This study utilized individual senesced sugar maple
and beech leaves as natural sampling units within
which to quantify saprotrophic fungal diversity. Quan-
tifying communities in individual leaves allowed us to
determine if fungi display a classic taxa–area relation-
ship (species richness increasing with area). We
found a significant taxa–area relationship for sugar
maple leaves, but not beech leaves, consistent with
Wright’s species-energy theory. This suggests that
energy availability as affected plant biochemistry is a
key factor regulating the scaling relationships of
fungal diversity. We also compared taxa rank abun-
dance distributions to models associated with niche
or neutral theories of community assembly, and
tested the influence of leaf type as an environmental
niche factor controlling fungal community composi-
tion. Among rank abundance distribution models, the
zero-sum model derived from neutral theory showed
the best fit to our data. Leaf type explained only 5%
of the variability in community composition. Habitat
(vernal pool, upland or riparian forest floor) and site
of collection explained > 40%, but could be attributed
to either niche or neutral processes. Hence, although
niche dynamics may regulate fungal communities at
the habitat scale, our evidence points towards neutral
assembly of saprotrophic fungi on individual leaves,
with energy availability constraining the taxa–area
relationship.
Introduction
Fungal community composition has been shown to be an
important factor regulating decomposition (Balser and Fir-
estone, 2005; Waldrop and Firestone, 2006). Therefore,
our ability to predict decomposition rates and responses
to environmental changes may be enhanced with
increased understanding of the processes that regulate
fungal distributions. Variation in composition of ecological
communities is commonly divided into two components:
alpha diversity (number and evenness of taxa within a
sampling unit) and beta diversity (taxa turnover among
areas) (Gaston and Blackburn, 2000).
For macroorganisms, it has been frequently observed
that there is a correlation between the size of habitat
patches or survey areas and the most fundamental
measure of alpha diversity, number of taxa detected
(Rosenzweig, 1995; Connor and McCoy, 2001; Lomolino,
2001; Drakare et al., 2006). The ‘taxa–area relationship’
(TAR) refers to the shape of the increase in number of
taxa with increasing area, and has been most often mod-
elled as a power law (S = cA
z
) where S is number of
species, A is area, c is the intercept in log-log space, and
z is a constant related to the rate of species turnover
across space (Arrhenius, 1921; Gleason, 1922). Taxa–
area relationships have been used to extrapolate species
richness (Colwell and Coddington, 1994; He and Leg-
endre, 1996; Plotkin et al., 2000), estimate regional
diversity inventories (Chong and Stohlgren, 2007), and
compare species abundance distributions (May, 1975;
Harte et al., 1999; Pueyo, 2006). TARs are also used as
an important tool in conservation efforts to protect species
from habitat fragmentation and destruction (Faith et al.,
2008). Quantifying fungal TARs may help us understand
processes regulating fungal community assembly and
determine how sampling design affects our ability to
detect fungal diversity.
Communities in discrete habitat patches (e.g. islands)
have proven useful in studying TAR patterns because
such communities have well-defined boundaries (Mac-
Arthur and Wilson, 1967; Schoener, 1976; Bell et al.,
2005). Although a few studies have utilized leaves as
discrete habitats for microbial communities (Kinkel et al.,
1987; Jacques et al., 1995; Newell and Fell, 1997), most
previous molecular studies examining forest floor fungal
communities have utilized homogenized mixed ‘grab’
samples of leaves (O’Brien et al., 2005; Neubert et al.,
2006; Blackwood et al., 2007a; Keeler et al., 2009;
Redford et al., 2010). However, early after leaf fall,
individual leaf boundaries must limit the size of fungal
Received 29 July, 2011; revised 5 March, 2012; accepted 7 March,
2012. *For correspondence. E-mail feinsteinlm@gmail.com; Tel. (+1)
330 672 3895; Fax (+1) 330 672 3713.
Environmental Microbiology (2012) 14(6), 1488–1499 doi:10.1111/j.1462-2920.2012.02737.x
© 2012 Society for Applied Microbiology and Blackwell Publishing Ltd