Secretory structures on the leaf rachis of Caesalpinieae and Mimosoideae (Leguminosae): implications for the evolution of nectary glands.

Cup- or sometimes slit-shaped nectary glands on the rachis are a widespread trait in the legume subfamily Mimosoideae, especially in derived tribes. Their spotty occurrence in genera that appear to be basal has led to uncertainty about when in the mimosoid radiation this character evolved. Until now, specialized rachis glands were unknown in caesalpinioids thought to be related to ancestral mimosoids. We report here the occurrence of rachis glands in seven of the ten species of the Paleotropical genus Erythrophleum, a member of the Dimorphandra group of caesalpinioids thought to include the sister group(s) of mimosoids. The histological structure and location of Erythrophleum glands suggest homology with those of mimosoids; these glands are simpler structurally than rachis glands of any known mimosoid. The Erythrophleum glands differ from those of most mimosoids in the following respects: (1) they are smaller than glands of mimosoids; (2) the secretory surface is sunken in a pit capped by a small round pore rather than exposed on a broad concave or flat surface; (3) a smaller number of cells are involved in production and secretion of nectar; (4) vascular supply to the nectary is less extensive; and (5) mechanical support tissue (sclerenchyma) is less extensive and less organized. Rachis glands appear to be absent in the nine other genera included in the Dimorphandra group. We also report the occurrence of other secretory structures (patches of glandular trichomes) on the rachis of some Caesalpinieae and Mimoseae that lack specialized nectary glands and suggest that these patches of trichomes are primitive homologues of more organized glands. We discuss the significance of these glands and of the patches of trichomes for understanding relationships among primitive mimosoids and related caesalpinioids, and for understanding the origin of ant-guard defenses typical of many mimosoids.

Molecular phylogenetic study of a myrmecophyte symbiosis: did Leonardoxa/ ant associations diversify via cospeciation?

The Leonardoxa africana (Leguminosae: Caesalpinioideae) complex is a group of four closely related taxa (L1 to L4) exhibiting various grades of specificity and specialization in mutualistic associations with ants. Each of the two most specialized species, Leonardoxa taxon 3 (L3) and L. africana sensu stricto (L4), interacts with a specific species of formicine ant, respectively Aphomomyrmex after and Petalomyrmex phylax, which nests in specialized swollen twigs. These two monotypic genera are the sole African members of the tribe Myrmelachistini, and their occurrence in closely related plants suggested thehypothesis that the two associations L4/Petalomyrmex and L3/Aphomomyrmex are derived by cospeciation from an ancestral association. Phylogenies based on DNA sequences were reconstructed for the ants and compared with phylogenies available for the plants in order to test for this hypothesis of cospeciation. The resulting topologies suggest either that the association with myrmelachistine ants arose several times or that a plant species (L2) and an ant population split off from an ancestral association. Furthermore, dates of speciation events appear to differ between ants and corresponding plants. An estimate of at least 4 million years was obtained for the separation of Aphomomyrmex and Petalomyrmex, whereas biological, biogeographic, and molecular-genetic data suggest a much more recent divergence for the plants. Thus, we reject the hypothesis of cospeciation and conclude that Aphomomyrmex and Petalomyrmex independently colonized different taxa of Leonardoxa. This striking instance of parallel evolution supports the notion that specific ant-plant associations originated by ecological fitting of preadapted partners. We discuss alternative evolutionary scenarios that are consistent with molecular data.

Factors affecting levels of some phenolic compounds, digestibility, and nitrogen content of the mature leaves ofBarteria fistulosa (Passifloraceae).

Levels of total phenolics, condensed tannins, acid detergent fiber, pepsin/cellulase digestibility, and nitrogen in mature leaves of 26 individuals of the ant-plant,Barteria fistulosa, have been determined. Analysis of the results in terms of the presence or absence of ants and the position of the branch from which the leaves were collected showed no relationship with concentrations of phenolics or fiber and only a weak relationship with digestibility and nitrogen. By contrast, light intensity strongly influenced levels of phenolics, notably condensed tannins, so that mature leaves of individuals growing in direct sunlight were less digestible and appeared to be of lower quality as food for herbivores than did mature leaves of individuals in shaded positions. Possible reasons for the variation in condensed tannin levels are discussed.