Foliose Halymenia species (Halymeniaceae, Rhodophyta) from Southeast Asia, including a new species, Halymenia malaysiana sp. nov.Botanica Marina


Pui-Ling Tan, Phaik-Eem Lim, Showe-Mei Lin, Siew-Moi Phang, Stefano G.A. Draisma, Lawrence M. Liao
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Botanica Marina 2015; 58(3): 203–217 *Corresponding authors: Phaik-Eem Lim, Instititute of Ocean and Earth Sciences, University of Malaya, 50603 Kuala Lumpur,

Malaysia; and Institute of Biological Sciences, University of Malaya, 50603 Kuala Lumpur, Malaysia, e-mail:; and

Showe-Mei Lin, Institute of Marine Biology, National Taiwan

Ocean University, Keelung 20224, Taiwan, R.O.C., e-mail:

Pui-Ling Tan and Siew-Moi Phang: Instititute of Ocean and Earth

Sciences, University of Malaya, 50603 Kuala Lumpur, Malaysia; and

Institute of Biological Sciences, University of Malaya, 50603 Kuala Lumpur, Malaysia

Stefano G.A. Draisma: Instititute of Ocean and Earth Sciences,

University of Malaya, 50603 Kuala Lumpur, Malaysia; and Faculty of Science, Department of Biology, Prince of Songkla University,

Hatyai, Songkhla, 90112 Thailand

Lawrence M. Liao: Graduate School of Biosphere Science,

Hiroshima University, 1-4-4 Kagamiyama, Higashi-Hiroshima 739-8528, Japan

Pui-Ling Tan, Phaik-Eem Lim*, Showe-Mei Lin*, Siew-Moi Phang, Stefano G.A. Draisma and Lawrence M. Liao

Foliose Halymenia species (Halymeniaceae,

Rhodophyta) from Southeast Asia, including a new species, Halymenia malaysiana sp. nov.

Abstract: Despite the large number of species discovered in Halymenia, many remain poorly known due to the scarce information available. In order to facilitate species discrimination of foliose Halymenia species in

Southeast Asia, molecular analysis and morphological studies were made on Halymenia collections from

Malaysia, the Philippines, and Indonesia. The rbcL phylogenetic analyses showed that there are at least six taxa of foliose Halymenia occurring in Southeast Asia.

Among the six taxa, a new species, Halymenia malaysiana

P.-L. Tan, P.-E. Lim, S.-M. Lin et S.-M. Phang, is proposed based on both rbcL sequence analyses and morphological observations. Halymenia malaysiana is characterized by thalli possessing oblong or suborbiculate blades with a supple cartilaginous structure and gelatinous (slimy) texture, arising from a small discoid holdfast without a stipe, abruptly expanding into a broad blade and having a smooth surface with sinusoidally undulated margins.

The phylogenetic analyses also revealed that Halymenia is a polyphyletic genus, which requires further taxonomic studies.

Keywords: Halymenia malaysiana sp. nov.; red algae;

Southeast Asia; taxonomy.

DOI 10.1515/bot-2015-0004

Received 7 January, 2015; accepted 24 April, 2015


The red algal genus Halymenia C. Agardh, comprising 69 currently accepted species, is one of the largest genera in terms of species within the family Halymeniaceae (De

Smedt et al. 2001, Guiry and Guiry 2014). It is mostly distributed in tropical and subtropical regions (Gargiulo et al. 1986, Kawaguchi and Lewmanomont 1999, Hernández-Kantun et  al. 2009). The genus is mainly characterized by gelatinous thalli, presence of anticlinal filaments and refractive ganglionic cells in the medulla, stellate cells in the inner cortex, and auxiliary cell ampullae with branched secondary filaments (Balakrishnan 1961, Abbott 1967, Chiang 1970, De Smedt et al. 2001).

Halymenia was established by C. Agardh (1817) and the generitype is Halymenia floresii (Clemente) C. Agardh collected from Cádiz, Spain. Chiang (1970) used the architecture of auxiliary cell ampullae as a primary feature to group species at the generic level in the Halymeniaceae.

According to Chiang’s generic concept, simple or once or more branched secondary ampullar filaments may emerge from long and slender primary ampullary filaments in

Halymenia-type auxiliary cell ampullae. The auxiliary cell ampulla of Halymenia is flattish, expanded when mature, and is intermediate between the Grateloupiatype and the Cryptonemia-type of ampulla based on its shape and the degree of branching (Chiang 1970). Several attempts have been made to study foliose Halymenia species in Southeast Asia. Kawaguchi and Lewmanomont (1999) made a detailed morphological study of Halymenia dilatata Zanardini by comparing the vegetative and reproductive features of the material from Vietnam and

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Japan with Indian material, and by studying the pattern of spore development to establish a better classification system for the western Pacific species. Four Halymenia species in the Philippines including branched and foliose species were examined by De Smedt et al. (2001): Halymenia durvillei Bory de Saint-Vincent, H. dilatata Zanardini,

Halymenia maculata J. Agardh, and Halymenia porphyraeformis Parkinson. In the following year, Lewmanomont and Kawaguchi (2002) compared the morphological and anatomical structures of both H. dilatata and H. maculata from Thailand, while Kawaguchi et al. (2002) studied the morphology of H. maculata from Vietnam.

Foliose Halymenia are difficult to identify due to their simple thalli, often resulting in misapplied names (Abbott 1967, De Smedt et al. 2001). Our recent collections from Southeast Asia, including Malaysia, Indonesia, and the Philippines, comprised a mixture of foliose Halymenia resembling H. dilatata, H. maculata, and several possible new species. Most of these collections exhibit immense morphological plasticity, making their specific identification difficult. In this study, both morphological examination and rbcL analyses were undertaken in order to understand the species diversity of the foliose


Materials and methods

Specimens were collected from various localities in Malaysia, Indonesia, and the Philippines intertidally or subtidally by snorkelling or SCUBA diving. Specimens were pressed on herbarium sheets or preserved in 5% formalin seawater (samples from Tun Mustapha Park and Raja

Ampat), and a subsample of each specimen was preserved in silica gel for molecular studies. Voucher specimens are deposited in the University of Malaya Seaweeds and