Ecological Entomology (2015), DOI: 10.1111/een.12191
Does host-plant diversity explain species richness in
insects? A test using Coccidae (Hemiptera)
YEN-PO L I N,
1,2
DIANNE H. COOK,
3
PENNY J. GULLAN
4
and L Y N G. COOK
2
1
College of Life Science, Shanxi University, Taiyuan, China,
2
School of Biological Sciences,
The University of Queensland, Brisbane, Australia,
3
Department of Statistics, Iowa State University, Ames, Iowa, U.S.A. and
4
Division of Evolution, Ecology and Genetics, Research School of Biology, The Australian National University, Canberra, Australia
Abstract. 1. The megadiverse herbivores and their host plants are a major component
of biodiversity, and their interactions have been hypothesised to drive the diversiication
of both.
2. If plant diversity inluences the diversity of insects, there is an expectation that insect
species richness will be strongly correlated with host-plant species richness. This should
be observable at two levels (i) more diverse host-plant groups should harbour more
species of insects, and (ii) the species richness of a group of insects should correlate with
the richness of the host groups it uses. However, such a correlation is also consistent with
a hypothesis of random host use, in which insects encounter and use hosts in proportion
to the diversity of host plants. Neither of these expectations has been widely tested.
3. These expectations were tested using data from a species-rich group of insects – the
Coccidae (Hemiptera).
4. Signiicant positive correlations were found between the species richness of coccid
clades (genera) and the species richness of the host-plant family or families upon which
the clades occur. On a global scale, more closely related plant families have more similar
communities of coccid genera but the correlation is weak.
5. Random host use could not be rejected for many coccids but randomisation
tests and similarity of coccid communities on closely related plant families show that
there is non-random host use in some taxa. Overall, our results support the idea that
plant diversity is a driver of species richness of herbivorous insects, probably via
escape-and-radiate or oscillation-type processes.
Key words. Adaptive radiation, coccids, herbivory, host breadth, host-plant diversity,
insect-plant interactions, phytophagous insects, scale insects.
Introduction
Seed plants and the herbivorous insects that feed upon them
are spectacularly species-rich and abundant, constituting the
largest directly interacting component of terrestrial biodiversity
(Lewinsohn et al., 2005). Many phytophagous insect lineages
are more diverse than their non-plant-feeding relatives (e.g.
Farrell, 1998; Strauss & Zangerl, 2002), and this has con-
tributed to the idea that host-plant specialisation has driven
the diversiication of insects (e.g. Ehrlich & Raven, 1964).
In support of this idea, most insect groups have been found
Correspondence: Yen-Po Lin, College of Life Science, Shanxi
University, 92 Wucheng Road, Taiyuan 030006, China. E-mail:
yenpo.lin@uqconnect.edu.au
to be relatively host-speciic, with a majority of species each
restricted to a single host-plant family (Strong et al., 1984; van
Nieukerken, 1986; Schoonhoven et al., 2005; Lin et al., 2010).
If host-plant diversity is a causative or enabling factor in insect
diversiication, it would be expected that there would be more
herbivores on more diverse clades of plants, and this is generally
the case (e.g. Wright & Samways, 1998; Gonçalves-Alvim &
Fernandes, 2001; Cuevas-Reyes et al., 2004; Proche¸ s et al.,
2009; Lin et al., 2010). Thus, phytophagous insects and their
hosts have been the focus of hypotheses about how interactions
among organisms might drive diversiication of one or both part-
ners, and provide many empirical examples used to test these
hypotheses (Futuyma & Agrawal, 2009; Proche¸ s et al., 2009).
There are several broad hypotheses about how plant diver-
sity might drive insect species richness and all predict that the
© 2015 The Royal Entomological Society 1