RESEARCH PAPER
Apomixis does not affect visitation to flowers of
Melastomataceae, but pollen sterility does
F. R. Maia, I. G. Varassin & R. Goldenberg
Departamento de Bot^ anica, Setor de Ci^ encias Biol ogicas, Universidade Federal do Paran a, Curitiba, Paran a, Brazil
Keywords
Breeding systems; buzz-pollination; pollen
viability; pollination.
Correspondence
F. R. da Maia, Departamento de Bot^ anica,
Setor de Ci^ encias Biol ogicas, Universidade
Federal do Paran a, Caixa Postal 19031,
81.531-980 Curitiba, Paran a, Brazil.
E-mail: fabiano.botanica@gmail.com
Editor
N. Vereecken
Received: 6 November 2014; Accepted: 1 July
2015
doi:10.1111/plb.12364
ABSTRACT
Apomixis is an asexual seed reproduction mechanism thorough which embryos are
originated from material tissues inside the ovules, without precedent fertilisation. It
allows plants to colonise new habitats, even in places where flower visitors are scarce
or where plants are isolate. Apomixis seems to be related to pollen sterility and, in
species with flowers that offer pollen as a reward for pollinators, the amount or qual-
ity of the pollen offered by these species may influence the amount of the visits and
specific composition of the visitors. In order to test this hypothesis, we studied breed-
ing systems of 16 species of Melastomataceae and their flower visitors, evaluating
composition and abundance of the visits to apomictic and sexual species. Apomictic
plants with no viable pollen or with pollen with low viability did not receive visits
from pollinators, and consequently probably produce strictly apomictic fruits. On the
other hand, apomictic and sexual plants with high pollen viability do receive visits; in
this case, apomictic plants may produce fruits and seeds through both sexual and
apomictic methods. The species composition of insects visiting Melastomataceae with
high pollen viability was similar, regardless of whether the plants were apomictic or
not. It seems that pollen viability levels are important to determine visits to the flow-
ers irrespective of breeding system.
INTRODUCTION
Apomixis is an asexual seed reproduction mechanism through
which embryos are originated from material tissues inside the
ovules, without precedent fertilisation (Hanna 1995; Jefferson &
Bicknell 1995; Koltunow et al. 1995; Savidan 2000). Due to lar-
ger plasticity associated with polyploid lineages, apomixis allows
plant species to colonise new habitats and promote geographic
range expansions (H€ orandl & Paun 2007; Santos et al. 2012;
Hojsgaard et al. 2014). Apomixis may also lead to reproductive
assurance (Holsinger 2000) in places where flower visitors are
scarce or where plants are isolated, with no other conspecific
individuals to provide pollen exchange (Baker 1967; Jefferson &
Bicknell 1995; H€ orandl et al. 2008; Santos et al. 2012). Apomixis
seems to be prevalent in some larger families, suggesting a
positive role in diversification (Hojsgaard et al. 2014).
Melastomataceae is a large, mostly tropical family with both
apomictic and sexual species (Renner 1989; Goldenberg &
Shepherd 1998; Santos et al. 2012). Apomixis is frequent in the
family (Goldenberg & Shepherd 1998; Santos et al. 2012) and is
generally associated with polyploidy and hybridisation (Men-
des-Rodrigues & Oliveira 2012; Caetano et al. 2013a,b), which
in turn are related to different levels of low pollen viability or
even complete pollen sterility (Mogie 1992; Richards 1997;
Goldenberg & Shepherd 1998; Goldenberg & Varassin 2001).
On the other hand, sexual species have high pollen viability
(Goldenberg & Shepherd 1998; Goldenberg & Varassin 2001;
Hoffmann & Varassin 2011). The latter are mostly dependent
on pollinators, which in Melastomataceae are usually bees that
are specialised pollen gatherers from flowers with poricidal
anthers, performing so-called ‘buzz-pollination’ (Renner
1989). Nevertheless, Melastomateceae species can also be polli-
nated by other animals, like flies, wasps or even vertebrates,
usually in nectar-producing species (Varassin et al. 2008) and
large-pored anthers that release pollen more easily, even to
unspecialised visitors (Kribel & Zumbado 2014).
One event that has been constantly and informally observed
in Melastomataceae is that flowers of the apomictic species are
rarely visited, and sometimes have no visitors at all (Renner
1989; Goldenberg & Shepherd 1998; Fendrich T. G. unpub-
lished data). Sexual species, however, seem to be intensively
visited (Santos et al. 2010; Couto-Pereira et al. 2011; Franco
et al. 2011; Brito & Sazima 2012; Fendrich T. G. unpublished
data). Although it seems obvious that apomictic species that
do not depend on pollinators to set fruits and seeds may be
consequently less visited by them, there are species from other
angiosperm families in which apomixis seems to be facultative,
resulting in mixed breeding systems (Asker & Jerling 1992;
Koltunow 1993; Richards 1997; Bicknell & Koltunow 2004;
Good-Avila et al. 2008). Moreover, other angiosperm families
also have pseudogamic species, i.e. apomictic plants that
depend on pollination events to produce seeds, because of
endosperm formation (Koltunow 1993; Richards 1997; Sam-
paio et al. 2013). From the visitor’s standpoint, one can
assume that there is a relationship between apomixis, high
proportion of sterile pollen and scarce visits to the flowers,
considering that pollen is the reward explored by the visitors.
It would be expected that flowers with a low amount of reward
should attract no visitors, and visits would be consequently
scarce.
Therefore, it is expected that flower visitation may vary
depending on the breeding system in Melastomataceae species.
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