PETRIFIED CYCAD STEMS FROM ARGENTINA 121
Issued 00 August 2004
© 2004 The New York Botanical Garden
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121
The Botanical Review 70(2): 121–133
Two New Petrified Cycad Stems, Brunoa gen. nov. and Worsdellia
gen. nov., from the Cretaceous of Patagonia (Bajo de Santa Rosa,
Río Negro Province), Argentina
ANALÍA E. ARTABE,
1
ALBA B. ZAMUNER,
1
AND DENNIS WM. STEVENSON
2
1
División Paleobotánica
Facultad de Ciencias Naturales y Museo de La Plata
1900 La Plata, Argentina
2
New York Botanical Garden,
Bronx, NY 10458, U.S.A.
I. Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
II. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
III. Locality, Materials, and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
IV. Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
V. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
VI. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
VII. Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
I. Abstract
Polyxylic columnar stems covered by persistent leaf bases and found in sediments assign-
able to the Upper Cretaceous of Bajo de Santa Rosa, Río Negro Province, Argentina, are de-
scribed as two new generic entities in the Cycadales. Anatomical characters are the basis for
their being assigned to the Encephalartoideae of the Zamiaceae. Brunoa santarrosensis gen. et
sp. nov. is characterized by the presence of polyxyly, cone domes, mucilage cavities, and
uniseriate to triseriate araucaroid, scalariform, or bordered intervascular pitting. Worsdellia
bonettiae gen. et sp. nov. has polyxyly, anastomosing medullary vascular bundles, centripetal
xylem, mucilage canals, and concentric extraxylary bundles. Some characters (polyxyly, med-
ullary vascular bundles, and cone domes) were used to determine the systematic position, while
other characters (mucilage reservoirs and centripetal xylem) were used to establish the rela-
tionship between polyxylic and monoxylic forms.
II. Introduction
Cycadales are the most primitive order within gymnosperms with extant representatives.
The evolutionary history as known from fossils begins in the Upper Paleozoic (Mamay, 1969,
1976; Taylor, 1969; Zhu & Du, 1981) and reaches maximum distribution during the Mesozoic,
with decline beginning toward the end of the same period. Presently, Cycadales are a very well