Understanding plant reproductive diversity

Flowering plants display spectacular floral diversity and a bewildering array of reproductive adaptations that promote mating, particularly outbreeding. A striking feature of this diversity is that related species often differ in pollination and mating systems, and intraspecific variation in sexual traits is not unusual, especially among herbaceous plants. This variation provides opportunities for evolutionary biologists to link micro-evolutionary processes to the macro-evolutionary patterns that are evident within lineages. Here, I provide some personal reflections on recent progress in our understanding of the ecology and evolution of plant reproductive diversity. I begin with a brief historical sketch of the major developments in this field and then focus on three of the most significant evolutionary transitions in the reproductive biology of flowering plants: the pathway from outcrossing to predominant self-fertilization, the origin of separate sexes (females and males) from hermaphroditism and the shift from animal pollination to wind pollination. For each evolutionary transition, I consider what we have discovered and some of the problems that still remain unsolved. I conclude by discussing how new approaches might influence future research in plant reproductive biology.     

Pollination by fungus gnats in four species of the genus Mitella (Saxifragaceae)

The first example of pollination by fungus gnats in the eudicots is reported. The genus Mitella (Saxifragales) is characteristically produces minute, inconspicuous, mostly dull-coloured flowers with linear, sometimes pinnately branched, petals. To understand the function of these characteristic flowers, we studied the pollination biology of four Mitella species with different floral traits and different sexual expression: dioecious M. acerina , gynodioecious M. furusei var. subramosa , and hermaphroditic M. stylosa var. makinoi and M. integripetala. Flower-bagging experiments showed that wind pollination did not occur in the dioecious and gynodioecious species. Two years of observations of flower visitors at six study sites in Japan revealed that the principal pollinators of all four Mitella were specific species of fungus gnats (Mycetophilidae), which landed on the flowers with their long spiny legs settling on the petals. Characteristically, numerous pollen grains were attached to the fungus gnats in specific locations on the body. Although, on average, 1.3–2.6 fungus gnats visited each inflorescence per day, the fruit set of both bisexual and female flowers exceeded 63%. These results suggest that fungus gnats are highly efficient pollinators of Mitella spp., and that Mitella flowers are morphologically adapted to pollination by fungus gnats.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 449–460.     

Experimental documentation of natural hybridization inCactaceae: Origin of Lloyd's Hedgehog Cactus,Echinocereus ×lloydii

The origin ofEchinocereus ×lloydii Britt. & Rose, pro sp. (Lloyd's Hedgehog Cactus) was investigated using comparative morphology, cytology, biochemistry, and particularly, artificial hybridization. Numerous artificial crosses between the putative parentsE. coccineus Engelm. (a species of claret-up cactus) andE. dasyacanthus Engelm. (Texas Rainbow Cactus) were successful, resulting in the production of hundreds of seeds with hybrid embryos. The F₁ hybrid progeny (i.e., syntheticE. ×lloydii) grew to sexual maturity in about four and one-half years, whereupon successful backcrosses and F₂ generation hybrids were also obtained. The known F₁ hybrids closely approximated naturalE. ×lloydii. The fertility of these syntheticE. ×lloydii was high, like their natural counterparts. The populations ofE. ×lloydii in Pecos County, Texas are inferred to have originated as the result of natural interspecific hybridization. It is assumed thatE. ×lloydii or similar plants may arise wherever the parental taxa grow sympatrically.     

Mussaenda yunnanensis sp. nov. (Rubiaceae), a new functionally dioecious species from Yunnan,China

Mussaenda yunnanensis, a new dioecious species of Rubiaceae from Yunnan Province, China, is described and illustrated. The new species can be recognized by its slender stem, congested‐cymose inflorescences and long corolla tubes. Differences between M. yunnanensis and two morphologically similar species (M. pubescens and M. antiloga) are presented. We also provide a key to all dioecious species of Mussaenda in China. The delimitation of the new species is further supported by molecular phylogenetic analyses based on eight plastid loci.     

Reproductive biology of Syzygiella rubricaulis (Nees) Steph. (Adelanthaceae,Marchantiophyta), a liverwort disjunctly distributed in high‐altitude Neotropical mountains

Syzygiella rubricaulis is a dioecious leafy liverwort disjunctly distributed and restricted to high‐altitude mountains in the Neotropics and the Azores. This study is part of a larger project examining the phylogeography of S. rubricaulis in the Neotropics, and our main goals were to understand its reproductive biology, where sex expression occurs, if vegetative propagules are frequently found, how the sexes are distributed in populations, how frequently sporophytes are formed and what environmental conditions influence sexual expression. S. rubricaulis patches are mostly female, but all patches also contain non sex‐expressing shoots. Out of 42 patches examined, 29 (69%) were sex‐expressing: 25 were unisexual (21 female and four male) and four of mixed sex (two male‐biased and two unbiased). At shoot level, out of 4200 shoots 18% were female and 7% male; among sex‐expressing shoots, 73% were female, representing a sex ratio of 0.8 (female‐biased). We encountered a total of 33 sporophytes in six patches (in Brazil, Venezuela and Ecuador). Leaf regenerants were found in one patch in Mexico. Low rates of sporophytes were likely related to low frequencies of male shoots and large distances between the sexes. As 25% of S. rubricaulis shoots expressed sex (occasionally producing sporophytes), we suggest that short‐distance (and rarely long‐distance) spore dispersal events occur in mountainous areas on a short‐term basis. On a long‐term basis, however, these events likely contribute to dynamic exchanges among populations in the Neotropics.