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Different position in the inflorescence. Demonstrated in Gaura mutabilis
(Onagraceae), where upper flowers contained three times as much nectar
as lower flowers (Cruden et al., 1983).
Differences in inflorescence size. In Asclepias quadrifolia (Asclepiada-
ceae) larger inflorescences produce less nectar per flower than smaller ones
(Pleasants & Chaplin, 1983).
Differences in microenvironment around the flower. A. castanea flow-
ers directly exposed to sun have a lower volume and higher concentration
than flowers in the shade (Nicolson & Nepi, 2005).
Differences in flower age. This may cause differences in nectar secretion
rate (nectar volume) and sugar concentration. These differences are docu-
mented in several species and it is common for older flowers to have a
lower volume of nectar. They may have higher or lower solute concentra-
tions than younger flowers (Wyatt & Shannon, 1986; Petanidou et al.,
1996; Navarro, 2001; Nicolson & Nepi, 2005).
Differences in the sexual phase of dichogamous flowers. In Delphinium
sp. (Ranunculaceae; Cruden et al., 1983), Lobelia cardinalis (Campanula-
ceae; Devlin & Stephenson, 1985), Echium vulgare (Boraginaceae; Klink-
hamer & de Jong, 1990), Alstroemeria aurea (Alstroemeriaceae; Aizen &
Basilio, 1998), and Euphorbia boetica (Euphorbiaceae; Narbona et al.,
2005) the total sugar content was higher in the male phase. On the other
hand, in Metrosideros collina (Myrtaceae; Cruden et al., 1983), Polyscias
sambucifolia (Araliaceae; Gillespie & Henwood, 1994) and C. carvi
(Apiaceae; Langenberger & Davis, 2002), nectar with a higher total sugar
content was produced during the female phase. Carlson and Harms (2006)
formulated two sets of adaptive hypotheses about the evolution of patterns
of gender-biased nectar production: sexual selection hypotheses and in-
breeding avoidance hypotheses.
Different flower morphs in individual plants. Linaria genistifolia
(Scrophulariaceae; Davis & Vogel, 2005) may have unspurred zygomor-
phic flowers, single-spurred zygomorphic flowers, and 3-5-spurred
actinomorphic flowers on the same individual. On average, multi-spurred
flowers produce three times more nectar sugar than single-spurred flowers,
and 400 times more than spurless flowers.
Differences in visitation by pollinators. This is important when polli-
nation induces changes in nectar secretion; e.g., in P. sambucifolia ,
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