Wood anatomy of the subfamily Crotonoideae (Euphorbiaceae s.s.)
from India: systematic implications with special reference to the
taxonomic delimitation of Givotia and Vernicia
Prem Prakash Jangid and Sangeeta Gupta
P. P. Jangid (ppjangid2010@gmail.com) and S. Gupta, Wood Anatomy Discipline, Botany Division, Forest Research Inst., IN-248006 Dehradun,
India.
Te wood anatomy of 15 representative species belonging to 12 genera of nine tribes of the subfamily Crotonoideae
(Euphorbiaceae) are comprehensively described with focus on systematic implications. In addition, ecological and evo-
lutionary aspects are evaluated. An identification key to the species based on wood anatomical features is presented. Te
wood microstructure of the tribes was found to be considerably heterogeneous reflecting an unnatural classification of the
subfamily. However, the results confirm the generic relationship within subtribe Aleuritinae and tribe Ricinodendreae.
Vernicia and Givotia may be recognized based on wood anatomical and morphological characters. Te tribes Micrandreae
and Adenoclineae have considerable similarity in wood anatomy. Te wood structure of the monogeneric tribes Trigonos-
temoneae and Geloneae idicate a close relationship with the tribe Crotoneae.
Te pantropical subfamily Crotonoideae is one of three
subfamilies of the spurge family Euphorbiaceae s.s . Te
subfamily include notable economic products such as
cassava ( Manihot esculenta), rubber ( Hevea brasiliensis) and
tung oils ( Vernicia spp.), as well as many medicinal plants
(Balakrishnan and Chakrabarty 2007).
Crotonoideae is traditionally circumscribed to contain
laticiferous taxa with crotonoid pollen. Morphology-based
classifications for the subfamily have been proposed by
Müller (1866), Bentham (1880), Pax (1890), Hurusawa
(1954), Hutchinson (1969), Webster (1975, 1994) and
Radcliffe-Smith (2001). Webster (1994) recognised 12
tribes and 68 genera in Crotonoideae. Te classification
of Radcliffe-Smith (2001) was almost identical to that of
Webster (1994) as far as the composition of Crotonoideae is
concerned (12 tribes, 74 genera). Wurdack et al. (2005) stud-
ied the molecular phylogeny of this subfamily and divided
it into four clades, viz Adenoclineae, Geloneae, ‘articulated
crotonoids’ and ‘inaperaturate crotonoids’. Systematic and
phylogenetic palynological studies have been important as
guides of the current classifications (Erdtman 1952, Punt
1962, Webster and Rupert 1973, Nowicke 1994, Lobreau-
Callen and Sua´Rez-Cervera 1997, Tokuoka and Tobe 1998,
Lobreau-Callen et al. 2001).
Prior to the current definition of Euphorbiaceae, Hayden
and Hayden (2000) and Mennega (2005) briefly compared
the wood structure of Crotonoideae with the uni-ovulate
subfamilies Acalyphoideae and Euphorbioideae, and also
with what was then recognized as the bi-ovulate subfami-
lies Phyllanthoideae and Oldfieldioideae. Tese latter sub-
families are now roughly equivalent to Phyllanthaceae and
Picrodentraceae. From an Indian perspective, an overview
of gross wood structure of the subfamily has been provided
by Gamble (1922), Pearson and Brown (1932), Metcalfe
and Chalk (1950) and Raturi et al. (2001). Rudall (1994)
studied the homology and evolution of laticifers in Croto-
noideae. Weidenhoeft et al. (2008) studied the phylogeny by
wood anatomy of tribe Crotoneae. Although several wood
anatomical studies have been done on Euphorbiaceae, the
wood anatomy in particular of subfamily Crotonoideae and
its systematic implication has not yet been studied in detail.
Te systematics of Crotonoideae in the present paper is
based on Webster (1994). Te subfamily is represented in
India by 60 species belonging to 16 genera and 10 tribes
(Balakrishnan and Chakrabarty 2007). Te present paper
deals with the wood anatomical study of 37 specimens
belonging to 15 representative species of 12 genera covering
9 tribes of the subfamily.
Te aims of this study was to evaluate the systematic
relationships of supra-generic taxa and evaluate the morpho-
logical and molecular classification of the subfamily Croto-
noideae based on a wood anatomical approach. A species
identification key should be developed based on wood ana-
tomical characters. Also ecological and evolutionary aspects
of the subfamily should be considered. As a result, we will
suggest possible modest taxonomic changes at generic and
© 2016 Te Authors. Nordic Journal of Botany © 2016 Nordic Society Oikos
Subject Editor and Editor-in-Chief: Torbjörn Tyler. Accepted 1 January 2016
Nordic Journal of Botany 000: 001–018, 2016
doi: 10.1111/njb.00990, ISSN 1756-1051
Early View (EV): 1-EV