Taxonomic importance of seed morphology in
Gentiana (Gentianaceae)
NINO DAVITASHVILI and GERHARD KARRER*
Institute of Botany, Department of Integrative Biology and Biodiversity Research, University of
Natural Resources and Applied Life Sciences, Gregor-Mendel Str. 33, Vienna 1180, Austria
Received 18 October 2009; accept for publication 10 November 2009
The seeds of Gentiana show high diversity in size, shape and surface pattern. Until now, seeds of only a limited
number of taxa have been studied in detail and their utility in the subgeneric classification of Gentiana has not
been evaluated. In this study we investigated seed micromorphology of the poorly known sections Pneumonanthe,
Frigidae and Isomeria. In order to evaluate the relevance of seed sculpturing for taxonomy, we selected qualitative
characters of seed shape and testa ornamentation of all documented sections and performed cluster analyses based
on presence and absence of coded characters. Several new scanning electron microscopy pictures of seeds provided
a number of valuable characters for the subgeneric differentiation of Gentiana. The results of the cluster analyses
of seed characters generally support the sectional classification given by former authors. However, the European
Gentiana asclepiadea does not group together with other taxa of section Pneumonanthe, but clusters near to the
strictly European section Gentiana. Gentiana froelichii, endemic to the south-east Alps and treated traditionally
within section Frigidae, clusters together with taxa of the European section Ciminalis. Our results reveal that seed
micromorphology supports the reassignment of some taxa incertae sedis within Gentiana. © 2010 The Linnean
Society of London, Botanical Journal of the Linnean Society, 2010, 162, 101–115.
ADDITIONAL KEYWORDS: cluster analysis – scanning electron microscopy (SEM) – seed micromorphology
– subgeneric classification – testa ornamentation.
INTRODUCTION
Data on features of seed micromorphology have
been reported to be useful for taxonomy in numer-
ous plant groups (e.g. Juan, Pastor & Fernández,
2000; Metzing & Thiede, 2001; Môro et al., 2001;
Segarra & Mateu, 2001; Plaza et al., 2004; Adams,
Baskin & Baskin, 2005; Song, Yuan & Küpfer,
2005). Seeds can provide relatively stable character
sets which are valuable for comparative studies at
every level of the taxonomic hierarchy. Most taxo-
nomic studies of seed micromorphology involve two
steps: (1) the description of seed testa characters
and (2) the definition of seed types. In general, such
descriptive ‘standardized’ work largely lacks the
comparative aspect, especially in the case of large
data sets. Utilization of the statistical approach for
establishing different types of seeds is an exception
rather than a rule in taxonomic literature.
The genus Gentiana L. (Gentianaceae) comprises
approximately 360 species distributed mainly in tem-
perate, arctic and alpine habitats of the northern
hemisphere. Gentiana was reduced several times
since its first description as a result of the segregation
of genera such as Gentianella Moench and Metagen-
tiana T.N.Ho and S.W.Liu. Current infrageneric clas-
sifications of Gentiana sensu Ho & Liu (2001) start at
the subgeneric (Halda, 1996) or sectional (Pringle,
1978, Ho & Liu, 2001) levels, using simple characters
of seed morphology and other features such as growth
form, flower morphology, chromosome number, etc.
The importance of seed morphology for taxonomic
studies of Gentiana was pointed out by Kusnezow
(1894), Müller (1982), Miège & Wüest (1984), Yuan
(1993), Omer & Qaiser (1995), Halda (1996), Ho & Liu *Corresponding author. E-mail: gerhard.karrer@boku.ac.at
Botanical Journal of the Linnean Society, 2010, 162, 101–115. With 6 figures
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 101–115 101