Anatomically preserved Woodwardia virginica (Blechnaceae) and a new filicalean fern from the middle Miocene Yakima Canyon flora of central Washington, USA
Kb. Pigg et Gw. Rothwell, Anatomically preserved Woodwardia virginica (Blechnaceae) and a new filicalean fern from the middle Miocene Yakima Canyon flora of central Washington, USA, AM J BOTANY, 88(5), 2001, pp. 777-787
Anatomically preserved Woodwardia virginica (Blechnaceae) and a newly recog
nized onocleoid fern are described from the middle Miocene Yakima Canyon fl
ora of central Washington State, USA. Identification of the W. virginica fo
ssils is based on a combination of vegetative pinnules, rhizome and stipe a
natomy, and fertile pinnules with indusiate sori and sporangia Like those o
f extant W. virginica. Fronds are isomorphic. Vegetative pinnae are elongat
ed and pinnatifid, with a secondary vein paralleling the midvein. Secondary
veins of the pinnule lobe anastomose to form primary areoles and are eithe
r simple or dicholomize toward the margin. Rhizomes have a simple dictyoste
le with 3-5 cauline vascular bundles and often a sclerotic hypodermis. Leaf
traces contain two large adaxial vascular bundles that occur laterally and
adaxially, flanking an are of 4-6 smaller bundles. Fertile pinnules have l
inear sori that are somewhat embedded in the laminae and are enclosed by a
thin indusium. Leptosporangia display a vertical annulus and an elongated s
talk. A second fern, Wessiea yakimaensis gen. et sp. nov., is represented b
y anatomically preserved branching rhizomes and attached frond bases that c
onform to the Onoclea-type pattern of rhizome and frond-base vasculature. R
hizomes have a simple dictyostele of 4-5 cauline meristeles. Leaf divergenc
e is helical, with paired hippocampiform rachial traces. These two ferns oc
cur in the same matrix with specimens of Osmunda wehrii. They demonstrate t
hat filicalean fern assemblages similar to those of extant temperate floras
were well established in western North America by the middle Miocene and f
urther emphasize the exceptional species longevity of some homosporous pter
idophytes.