Phytotaxa 201 (4): 241–255
www.mapress.com/phytotaxa/
Copyright © 2015 Magnolia Press
Article
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Rafael Riosmena-Rodriguez: 14 Jan. 2015; published: 5 Mar. 2015
http://dx.doi.org/10.11646/phytotaxa.201.4.1
241
Taxonomic reassessment of Gracilaria cearensis (Rhodophyta, Gracilariales),
a poorly defined yet common flattened species based on morphological and
molecular analysis including topotype collections
LUANDA PEREIRA SOARES
1,5*
, CARLOS FREDERICO D. GURGEL
2,3,4
& MUTUE TOYOTA FUJII
5
1
Post-Graduate Program “Biodiversidade Vegetal e Meio Ambiente”, Instituto de Botânica, Av. Miguel Estéfano, 3687, São Paulo, SP,
04301-902, Brazil
2
Departamento de Botânica, Centro de Ciências Biológicas, Universidade Federal de Santa Catarina, Florianópolis, SC, 88040-970,
Brazil
3
School of Earth and Environmental Sciences, The University of Adelaide, Aidelaide, SA 5005, Australia
4
State Herbarium of South Australia, DEWNR, South Australia Government, PO Box 2732, Kent Town, SA 5071, Australia
5
Instituto de Botânica, Núcleo de Pesquisa em Ficologia, São Paulo, SP, 04301-902, Brazil
* Corresponding author: luanda87@gmail.com
Abstract
The genus Gracilaria is the major source of agarose in the world today and is one of the most species-rich genera in the
Rhodophyta. Flat Gracilaria species are among the most taxonomically challenging taxa due to their widespread phenotypic
plasticity among all species. Gracilaria cearensis is a flat species described in 1965 from Brazil. Its original description is
not conducive to accurate taxonomic identifications and the delineation of this taxon remains elusive. New samples of G.
cearensis were collected across a coastal length of 500 km including its type locality. Universal Plastid Amplicon (UPA) and
the rbcL gene were used to confirm the phylogenetic and taxonomic status of this poorly known species. Results showed
that this is a distinct species despite morphological similarities to G. hayi, G. cuneata, G. curtissiae, G. brasiliensis and G.
galetensis. G. cearensis is sister to G. hayi yet the genetic divergence between these two species was 2% for rbcL and 1.08%
for UPA, enough to consider them distinct taxa. However, newly generated molecular data placed G. smithsoniensis, an-
other morphologically similar species, as conspecific with G. cearensis with only 0.07% rbcL sequence divergence between
them.
Key words: biodiversity, rbcL, taxonomy, UPA
Introduction
Gracilaria cearensis (A.B. Joly & F.C. Pinheiro 1965: 81) A.B. Joly & F.C. Pinheiro (1966: 131) is a tropical intertidal
species with flat, strap-shaped branched thalli and commonly occurring in shallow reefs along the northeastern
Brazilian coast (Pinheiro-Vieira & Ferreira 1968). In Brazil, G. cearensis has industrial potential as a source of agar and
has attracted biotechnological interest due to its hemagglutination and antibiotic activity (Pinheiro-Vieira & Caland-
Noronha 1971, Ainouz & Sampaio 1991).
The species was originally described as Tylotus cearensis Joly & Pinheiro (1965: 81) based on material collected
in Fortaleza, Ceará state, Brazil. The genus Tylotus J. Agardh (1876: 428) was excluded from the Gracilariaceae by
Dawson (1949) based on the presence of zonate tetrasporangia in the type species, T. obtusatus (Sonder 1845: 56) J.
Agardh (1876: 428). Later, Pinheiro & Joly (1966) transferred T. cearensis to Gracilaria Greville (1830: 121) in the
light of its cruciately divided tetrasporangia. G. cearensis shows other main characters such as tubular nutritive cells
present in both pericarp and the cystocarp floor, and spermatangial conceptacles described as “usually not confluent”
as observed in most Gracilaria species. Since its description no further taxonomic or morphological studies have been
performed in order to ascertain the status of this taxon, despite its purportedly widespread distribution in northeastern
Brazil, from Ceará (Joly & Pinheiro 1965) to Bahia state (Nunes 2005).
Morphologically, G. cearensis is similar to G. brasiliensis Gurgel & Yoneshigue-Valentin (2008: 255), G. cuneata
Areschoug (1854: 351), G. galetensis Gurgel, Fredericq & J. Norris (2004: 181), G. hayi Gurgel, Fredericq & J. Norris