植生史研究 Jpn. J. Histor. Bot. Vol. 19 Nos. 1–2 p. April 2011
©2011 Japanese Association of Historical Botany
Introduction
The identification of the living representatives of
the Taxodiaceae (now subsumed within the Cupres-
saceae) is based on a whole-plant concept, utilizing
an approach that recognizes significant morphologi-
cal features such as the shape, architecture, and age of
the tree; leaf arrangement on the leading and lateral
branches; leaf morphology; and the size and shape of
the seed cones (Silba, 1986; Rushforth, 1987; Wu &
Raven, 1999; Farjon, 2005). Regardless of the fea-
tures used, perhaps the single most useful feature that
facilitates species identification, description, and clas-
sification among the conifers is seed cone morphology
(Matsumoto et al., 1997; Aulenback & LePage, 1998;
LePage, 2001, 2003). Whereas leaf morphology alone
can sometimes be valuable, the identification and clas-
sification of isolated leaves that are preserved as com-
pression fossils is generally difficult in taxa where leaf
polymorphism is prevalent. Among the genera of the
Taxodiaceae, leaf polymorphism is present in Glyp-
tostrobus Endlicher, Metasequoia Hu et Cheng, Se-
quoiadendron Buchholz, Sequoia Endlicher, Taiwania
Hayata, and Taxodium Richard (Brown, 1936; Bůžek,
1971; Christophel, 1976; Farjon, 2005).
Although this is not necessarily problematic when
working with living representatives, variation in leaf
morphology together with preservation quality further
contribute to the difficulty of correctly identifying fos-
sil leaves that are not associated with reproductive or-
gans. Such difficulties are further amplified when sever-
al taxa that are known to produce polymorphic leaves
are preserved as part of the same floristic assemblage.
As a result, the paleobotanical literature is rife with
morphospecies names that are based on isolated leaves
that were incorrectly identified (Heer, 1868–1878;
Chaney, 1951; LePage et al., 2005; LePage, 2007). This
however does not mean that misidentified specimens
have no taxonomic and phylogenetic value. On the
contrary, the reassessment of fossil leaves and their ex-
tant counterparts may often reveal morphological and
anatomical features that were previously overlooked
as being diagnostic (Matsumoto et al., 1994, 1995;
Stockey & Ko, 1986; Stockey & Atkinson, 1993).
The problem of correctly identifying fossil taxodia-
ceous leaves is not unique to any generation of paleo-
botanists and the uncertainty of identifying isolated
leaves extends to the earliest plant classification sys-
tems. Brongniart established the genus Taxites Brong-
niart for fossil leaves that were “analogous in part to
the yews and Podocarpus, while in other cases they
could not be separated from the yews or Taxodium”
(Brongniart, 1828, p. 101), while Presl created the ge-
nus Taxodites Presl for “distichous twigs and leaves
strongly analogous to those of Taxodium” (Presl in
Sternberg, 1838, p. 204). Chaney (1951, p. 174) also
recognized this problem and stated that “As far back as
the middle of the past century, fossil foliage of Sequoia
has been confused with that of Taxodium. During all
109–116
Ben A. LePage
1
: A classification system to separate leaves
of the Cephalotaxaceae, Taxaceae, and Taxodiaceae
Abstract The identification and classification of isolated fossil leaves of the Taxodiaceae is generally difficult
when the remains are not associated with reproductive organs. These problems are further exacerbated when
more than one genus of the Taxodiaceae and/or genera of the Taxaceae or Cephalotaxaceae are preserved in the
same deposits or the quality or mode of preservation precludes confident identification. To help facilitate the
identification and separation of isolated taxodiaceous fossil leaves, the leaves of extant representatives of Cepha-
lotaxus Siebold et Zuccarini ex Endlicher, Glyptostrobus Endlicher, Metasequoia Hu et Cheng, Sequoia Endli-
cher, Taxodium Richard, Taxus L., and Torreya Arnott were examined. The results of this investigation provided
sufficient data about external morphological features to establish a system of classification for leaves that could
be used to assist in the identification and classification of isolated fossil leaves.
1
Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, Pennsylvania, 19103 USA and PECO Energy
Company, 2301 Market Street, S7-2, Philadelphia, Pennsylvania, 19103 USA
(e-mail: ben.lepage@exeloncorp.com)