Transferring Gracilaria irregularis (Gracilariaceae,
Rhodophyta) from Thailand to Gracilariopsis based on
morphological and molecular analyses
Narongrit Muangmai,
1
Yukimasa Yamagishi,
2
Giuseppe C. Zuccarello,
1
Anong Chirapart
3
and
Khanjanapaj Lewmanomont
3
*
1
School of Biological Sciences, Victoria University of Wellington, Wellington, New Zealand,
2
Department of Marine
Biotechnology, Faculty of Life and Biotechnology, Fukuyama University, Hiroshima, Japan, and
3
Department of Fishery
Biology, Faculty of Fisheries, Kasetsart University, Bangkok, Thailand
SUMMARY
Gracilaria irregularis, endemic to Thailand, was origi-
nally described from Ao Len, Trat peninsula, Thailand,
and was characterized by a short, succulent thallus
with irregular branching, verrucosa-type spermatangia
(a taxonomic criterion for Gracilaria) and the absence
of nutritive filaments (a taxonomic criterion for
Gracilariopsis). Due to the combination of characters
diagnostic of both the genera Gracilaria and Graci-
lariopsis, the taxonomic status of this species is
unclear. This present study reassesses the identity of
G. irregularis based on morphological features and
the large subunit of ribulose-bisphosphate carboxylase/
oxygenase gene sequences. Newly collected samples
from the type locality and KU collection were investi-
gated. Our morphological re-examinations were rela-
tively similar to the original type description, except
that the male plants showed superficial chorda-type
spermatangia, an important characteristic recognized in
Gracilariopsis. Molecular analysis placed G. irregularis
in the Gracilariopsis clade, forming a well-supported
clade with Gracilariopsis bailiniae. Detailed morphology
and molecular phylogenetic analyses clearly showed that
G. irregularis is congruous with the generic concept of
Gracilariopsis, and consequently the new combination of
Gracilariopsis irregularis is proposed. We believe that in
the original description a heterogeneous holotype was
used. Our data also shows the problems that can occur
when collections are made of multiple plants, of similar
external morphology, to describe new species.
Key words: Gracilariaceae, Gracilariopsis irregularis,
Gracilariopsis, morphology, rbcL, red algae,
spermatangia, Thailand.
INTRODUCTION
For more than a decade, the systematic of the marine
red algal family Gracilariaceae has been extensively
investigated and increasingly has relied on molecular
data, together with detailed comparative morphology
(e.g. Bellorin et al. 2002; Gurgel & Fredericq 2004;
Kim et al. 2006; Muangmai et al. 2012). Morphologi-
cally, identification of this family is based on internal
anatomy of mature cystocarps and spermatangial
arrangements (Fredericq & Hommersand 1990; Gurgel
& Fredericq 2004). The Gracilariaceae presently
contains seven distinct genera (Gurgel & Fredericq
2004), of which four genera, Congracilaria Yamamoto,
Gracilaria (G.) Greville, Gracilariopsis (Gp.) E.Y. Dawson
and Hydropuntia Montagne, are known in Thailand
(Terada et al. 2004; Chirapart 2008). Among the four
genera recognized in Thai waters, Gracilaria is the most
diverse and widespread genus: nine species have been
recorded so far, two of which, namely G. irregularis
Abbott and G. minuta Lewmanomont are reported as
endemic to Thailand (Abbott 1988; Lewmanomont
1994; Terada et al. 2004).
Gracilaria irregularis I.A. Abbott was originally
described based on collections from Ao Len, Trat
Peninsula, Thailand, and it was characterized by
inflated thalli with irregular branching, deep pot-shaped
spermatangial conceptacles (verrucosa-type), and the
absence of nutritive filaments (Abbott 1988). This
species is known only from the Gulf of Thailand (Terada
et al. 2004; Chirapart 2008) and commonly grows on
sandy-mud bottom in the lower intertidal (0.5 m
depth), attaching to shell and rock (Abbott 1988;
Lewmanomont 1994). Due to little research on this
species, detailed morphology, phylogenetic relation-
ships and its ecology are poorly characterized.
The taxonomic status of G. irregularis has long
been in confusion, owing to the inconsistent views of
*To whom correspondence should be addressed.
Email: ffiskpl@ku.ac.th
Communicating editor: S-M. Lin.
Received 28 February 2013; accepted 18 June 2013.
doi: 10.1111/pre.12032
Phycological Research 2014; 62: 29–35
© 2013 Japanese Society of Phycology