Reproduction and Breeding Systems

Among flowering plants, the Cactaceae have flowers that are strongly associated with animals, such as bees, hawk-moths, hummingbirds, and bats (Porsch 1939; Grant and Grant 1979; Gibson and Nobel 1986; Valiente-Banuet et al. 1996; Chapter 5). Plant-pollinator relationships are specialized for some cactus species, such as for opuntias with large solitary, diurnal, bowl-shaped flowers adaptable to bee pollination. Bee fauna in the Chihuahuan Desert of Mexico, where opuntia species are highly diversified, is particularly rich with at least 90 species of bees visiting opuntia flowers in North America (Grant and Hurd 1979). Other species pollinated by bees belong to tribe Pachy-cereeae, such as Myrtillocactus geometrizans.

Tubular, red, hummingbird-pollinated flowers are common in Andean and tropical American cacti, especially in genera such as Rathbunia, Peniocereus, Nopalea, and Pachycereus (Gibson and Nobel 1986). The hawkmoth-flower syndrome—characterized by strongly scented night-blooming flowers with white or whitish perianths and long, slender nectar-containing floral tubes—occurs in species of Acanthocereus, Cereus, Epiphyllum, Selenicereus, and Trichocereus (Grant and Grant 1979). The bat-flower syndrome—consisting of nocturnal anthesis, whitish bowl-shaped flowers with a strong unpleasant odor at night, large quantities of pollen and nectar, many anthers, and large-mouthed and strong single flowers (Faegri and van der Pijl 1979; Grant and Grant 1979)—occurs for about 70% of columnar cacti in tribe Pachycereeae (Valiente-Banuet et al. 1995a, 1996). Most such species studied in

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