title: Pollen structure and development in Nymphaeales: insights into character evolution in an ancient angiosperm lineage.
abstract: OBJECTIVE
  A knowledge of pollen characters in early-diverging angiosperm lineages is essential for understanding pollen evolution and the role of pollen in angiosperm diversification. In this paper, we report and synthesize data on mature pollen and pollen ontogeny from all genera of Nymphaeales within a comparative, phylogenetic context and consider pollen evolution in this early-diverging angiosperm lineage. We describe mature pollen characters for Euryale, Barclaya, and Nymphaea ondinea, taxa for which little to no structural data exist.
  
  
  METHODS
  We studied mature pollen for all nymphaealean genera using light, scanning electron, and transmission electron microscopy. We reviewed published reports of nymphaealean pollen to provide a comprehensive discussion of pollen characters in water lilies.
  
  
  RESULTS
  Nymphaeales exhibit diversity in key pollen characters, including dispersal unit size, ornamentation, aperture morphology, and tapetum type. All Nymphaeales pollen are tectate-columellate, exhibiting one of two distinct patterns of infratectal ultrastructure-a thick infratectal space with robust columellae or a thin infratectal space with thin columellae. All genera have pollen with a lamellate endexine that becomes compressed in the proximal, but not distal wall. This endexine ultrastructure supports the operculate hypothesis for aperture origin. Nymphaeaceae pollen exhibit a membranous granular layer, which is a synapomorphy of the family.
  
  
  CONCLUSIONS
  Variation in pollen characters indicates that significant potential for lability in pollen development was present in Nymphaeales at the time of its divergence from the rest of angiosperms. Structural and ontogenetic data are essential for interpreting pollen characters, such as infratectum and endexine ultrastructure in Nymphaeales.
authors.affiliation: Department of Biology, Creighton University, 2500 California Plaza, Omaha, Nebraska 68178-0324 USA MackenzieTaylor@creighton.edu.
authors.forename: Mackenzie L
authors.initials: ML
authors.lastname: Taylor
authors.affiliation: Biology Department, Hillsdale College, 33 East College Street, Hillsdale, Michigan 49242 USA.
authors.forename: Ranessa L
authors.initials: RL
authors.lastname: Cooper
authors.affiliation: The University of Minnesota Landscape Arboretum, 3675 Arboretum Drive, Chaska, Minnesota 55318 USA.
authors.forename: Edward L
authors.initials: EL
authors.lastname: Schneider
authors.affiliation: School of Science, The College of New Jersey, P.O. Box 7718, 2000 Pennington Road, Ewing, New Jersey 08628-0718 USA.
authors.forename: Jeffrey M
authors.initials: JM
authors.lastname: Osborn
mesh_terms: D005075:Biological Evolution; D008296:Malaysia; D008855:Microscopy, Electron, Scanning; D046529:Microscopy, Electron, Transmission; D030721:Nymphaea / Q000033:anatomy & histology / Q000145:classification / Q000254:growth & development / Q000648:ultrastructure; D030702:Nymphaeaceae / Q000033:anatomy & histology* / Q000145:classification / Q000254:growth & development* / Q000648:ultrastructure; D010802:Phylogeny; D011058:Pollen / Q000033:anatomy & histology* / Q000254:growth & development* / Q000648:ultrastructure; D054817:Pollination; D014481:United States; D014904:Western Australia
keywords: Cabombaceae; Hydatellaceae; Nymphaeaceae; exine; morphology; pollen; pollination; tapetum; ultrastructure; water lilies