Laurencia (PROSEA)

Plant Resources of South-East Asia Introduction

List of species

Laurencia papillosa - 1, habit; 2, detail of ultimate part. L. cartilaginea - 3, detail of ultimate part. L. okamurae - 4, part of a cross-section through a branchlet. L. obtusa - 5, surface view of cortex cells with one cherry body in each cell and secondary pit connections; 6, longitudinal section of the same area. L. tronoi - 7, ultimate branchlet with cystocarps; 8, ultimate branchlet with spermatangial branchlets; 9, ultimate branchlet with tetrasporangial branchlets.

Laurencia J.V. Lamour.

Protologue: Essai Thalassioph.: 130 (1813).

Family: Rhodomelaceae

Chromosome number: x= unknown; L. papillosa: 2n= 40-52, L. okamurae: 2n= 64

Major species and synonyms

• Laurencia botryoides (C. Agardh) Gaillon, Résumé Thalassioph.: 363 (1828), synonyms: Fucus botryoides Turner (1809-1811, nom. illeg.), Chondria botryoides C. Agardh (1817).

• Laurencia cartilaginea Yamada, Univ. Calif. Publ. Bot. 16 (7): 230, fig. 0, pl. 19: fig. a (1931). Accepted name is now: Chondrophycus cartilagineus (Yamada) Garbary & J.T.Harper (1998).

• Laurencia composita Yamada, Univ. Calif. Publ. Bot. 16 (7): 236, figs R, S, pl. 23 (1931).

• Laurencia flexilis Setch., Univ. Calif. Publ. Bot. 12: 101, pl. 19 (1926).

• Laurencia nidifica J. Agardh, Spec. gen. ord. alg. 3 (1): 749-750 (1852) [1852-1863], synonym: L. obtusa (Huds.) J.V. Lamour. var. nana Harv. (1834).

• Laurencia obtusa (Huds.) J.V. Lamour., Essai Thalassioph.: 130 (1813), synonym: Fucus obtusus Huds. (1778).

• Laurencia okamurae Yamada, Univ. Calif. Publ. Bot. 16 (7): 206, figs J, K, pl. 7 (1931, as L. okamurai).

• Laurencia palisada Yamada, Univ. Calif. Publ. Bot. 16 (7): 196, figs C, D, pl. 4, fig. a (1931).

• Laurencia papillosa (C. Agardh) Grev., Alg. brit.: LII (1830), synonyms: Fucus papillosus Forssk. (1775, nom. illeg.), Chondria papillosa C. Agardh (1822). Accepted name is now: Palisada perforata (Bory) K.W.Nam (2007).

• Laurencia parvipapillata C.K. Tseng, Pap. Michigan Acad. Sci. 28: 204-205, pl. IV (1943).

• Laurencia patentiramea (Mont.) Kütz., Sp. alg.: 854 (1849), synonyms: Chondria obtusa (Huds.) C. Agardh var. paniculata C. Agardh (1822), C. obtusa (Huds.) C. Agardh var. patentiramea Mont. (1836), Laurencia paniculata (C. Agardh) J. Agardh (1852). Accepted name is now: Palisada patentiramea (Montagne) Cassano, Sentíes, Gil-Rodríguez & M.T.Fujii (2009).

• Laurencia similis K.W. Nam & Y. Saito, Brit. Phycol. J. 26: 375-382 (1991).

• Laurencia snackeyi (Weber Bosse) Masuda, in Masuda, Y. Takahashi, K. Okamoto, Matsuo & Min. Suzuki, Eur. Journal of Phycology 32: 296 (1997), synonyms: L. paniculata (C. Agardh) var. snackeyi Weber Bosse (1923), L. obtusa (Huds.) J.V. Lamour. var. snackeyi (Weber Bosse) Yamada (1931).

• Laurencia tronoi Ganz.-Fort., Kalikasan 11: 404, figs 1, 2 (1983).

Vernacular names

General:

• Laurencia
• Philippines: culot, kulot (Ilocano).

L. flexilis

• Philippines: culto (Ilocano).

L. papillosa

• Philippines: dipdipig, tartariptip, layalaya.

L. snackeyi

• Indonesia: sangau.

Origin and geographic distribution

Laurencia is a common component of seaweed communities in tropical and temperate waters. In general, however, detailed knowledge is needed to identify up to species. Most species listed here have been recorded from the Philippines, where these algae are frequently studied. Records from other countries in South-East Asia are more incidental than the result of detailed research. It is possible that records for L. botryoides (an Australian species) and of L. obtusa are based on mis-identifications. Especially reportedly cosmopolitan species need critical re-evaluation.

Uses

Several Laurencia (especially L. cartilaginea, L. patentiramea and L. snackeyi) are popular in areas in the Philippines and Indonesia for use in salads, in vegetable soup and as a medicine. In fact all Laurencia can be used for human food. Medical uses include as antibiotic (antibacterial or antifungal) and (in Indonesia for L. snackeyi) for treatment of stomach ailments. Occasionally, large amounts of washed-up Laurencia thalli are used as fertilizer.

Production and international trade

There are no data available on the amount harvested for food from wild populations of Laurencia.

Properties

Most Laurencia contain agar, but the quality of the phycocolloids of these algae is not yet fully known. Chemical investigations of the polysaccharide of L. flexilis, involving sulphate and 3,6-anhydrogalactose determinations as well as FT-IR (Fourier transform infrared spectroscopy) spectroscopic examinations, revealed an agar-type polysaccharide. Samples of L. flexilis from Hainan Island (China) contain 36.4% of agar (dry weight), with a gel strength of 458 g/cm²for the alkali-treated fraction. Detailed studies of the phycocolloid of L. papillosa in India suggest a relation to lambda carrageenan. However, many other researchers have suggested that the polysaccharides from cell walls of Laurencia should be considered as belonging to the agar group, being agarose derivates with a rather high content of alkali-stable sulphate groups and with a reduced amount of 3,6-anhydro-L-galactose residues. This has in some cases been verified by the use of agarose from the bacterium Cytophaga. It is obvious that different Laurencia produce different types of hot-water-soluble cell-wall polysaccharides.

The predominant saturated fatty acid in the lipids of L. papillosa is palmitic acid (16:0), while the predominant unsaturated fatty acid is linolenic acid (10.2%; 18:1).

Extracts of fresh L. snackeyi (as L. obtusa) show some antioxidant activity that might be used in the preservation of food. Laurencia spp., especially L. patentiramea, contain the phytohormones auxin, gibberellin and cytokinin, and the minerals (especially L. papillosa) B, Ca, Cu, Fe, I, K, Mg, Mn, N, Na, Zn, as well as chlorides and sulphates.

Many Laurencia contain special secondary metabolites which can be antibiotic and prevent grazing by fish and sea urchins. Despite the problems of classification and identification, the biochemical components of Laurencia make it biochemically one of the most studied of all algal genera. Sesquiterpenoids, diterpenoids and acetylenic non-terpenoids form the three major classes of Laurencia-derived compounds, often containing the halogens bromide or chlorine.

Description

• Thalli erect or decumbent, attached by disk or shovel-shaped holdfasts (haptera) or crustose base.
• Fronds fleshy, usually crisp, terete or compressed.
• Branches radially or pinnately divided; apices of ultimate branchlets depressed, with a tuft of usually inconspicuous, colourless hairs (trichoblasts).
• Central axial cells and pericentrals soon obscured by formation of other cells, thus cross-sections appearing pseudoparenchymatous; outer cortical cells in cross-section radially elongate (= palisade pattern) or nearly isodiametric and rounded to angular, with or without bright cell inclusions (cherry bodies), with or without lateral secondary pit connections; in some species surface of younger branches papillose, due to projecting outer tangential walls of cortex cells; inner cortical cells or medullary cells with or without lenticular thickenings of cell walls.
• Life cycle triphasic, diplo-haplontic and isomorphic.
• Tetrasporangia tetrahedral.
• Gametophytes dioecious.
• Spermatangia borne in stichidia on ultimate branchlets.
• Cystocarps conspicuous, usually nearly equal in diameter to bearing branchlet.

L. botryoides.

• Thalli dark red to red-brown, up to 17 cm tall, firm, drying cartilaginous, pyramidal in outline.
• Branches usually numerous; laterals alternate, distichous, 1-1.5 mm thick, arising from flexuous axis, 1.5-2(-3) mm thick; axes and branches terete or slightly compressed, bearing small reproductive ramuli; ramuli first distichous, later developing on all sides forming botryoidal clusters.
• Outer cortical cells about 1.5 times as long as broad in cross-section, with lateral secondary pit connections, without bright cell inclusions (cherry bodies) and not projecting.
• Medullary cells without lenticular thickenings.
• Tetrasporangia ovoid, 80-100 μm in diameter, on compound branchlets comprising botryoidal clusters of very short, wart-like single ramuli, arranged in right-angled manner; spermatangial stichidia very short, in botryoidal clusters; cystocarps crowded on ramuli, sessile, globular, 0.8-1 mm in diameter, with prominent ostiole.

L. cartilaginea.

• Thalli pink to brown, erect or caespitose, up to 9 cm tall.
• Main axis compressed, subcompressed or angular, cartilaginous, up to 1 mm in diameter, older portions always covered by calcareous material.
• Branching distichous with up to 3 orders of branching, ultimate branches terete, up to 1 mm in diameter with wart-like branchlets.
• Cortical cells not arranged in palisade pattern, not projecting, without cherry bodies, not connected to each other by secondary pit connections.
• Medullary cells with relatively thick cell walls and no lenticular thickenings.
• Tetrasporangiate branchlets abruptly inflated from usually long and slender bases, 0.7 mm in diameter; tetrasporangia in radial (= right-angled) arrangement; male stichidia simple, inflated, up to 1.2 mm at its widest diameter; spermatia in apex in shallow pits. No female specimens observed.

L. composita.

• Thalli dark purple, up to 10 cm tall, soft in texture.
• Branches irregularly spirally arranged, long, issuing on every side of percurrent main axis, terete, 0.8-2 mm thick; branchlets simple, up to 1 cm long, alternate or subopposite, paniculate.
• Outer cortical cells small, connected by longitudinal secondary pit connections, with one cherry body per cell, not palisade-like, not projecting. Medullary cells with rare lenticular thickenings.
• Tetrasporangia in cylindrical-clavate, paniculated, or cymosely fasciculated ultimate branchlets, 120-150 μm in diameter; cystocarps ovoid, 700-900 μm × 550-740 μm, with single, wide apical ostiole; spermatangia in apical pits of ultimate branchlets.

L. flexilis.

• Thalli purple to bleached, erect, up to 7(-13) cm tall. Main axis terete, wiry, rigid, up to 1.8 cm in diameter.
• Branching variable but generally radial, with up to 3 orders of branches; ultimate branches terete, simple, up to 8 mm in diameter.
• Cortical cells in palisade arrangement, not projecting, without cherry bodies, probably connected to each other by secondary pit connections.
• Medullary cells with relatively thick cell walls and no lenticular thickenings.
• Tetrasporangiate branches simple, up to 8 mm in diameter; tetrasporangia in a radial arrangement; male stichidia simple, rarely compound with truncate tips, up to 0.8-1 mm in its most inflated diameter; spermatia in apex in shallow pits; cystocarps globose, almost terminal up to 10 mm in diameter.

L. nidifica.

• Thalli erect, straw-pinkish, up to 10 cm tall, with several erect axes.
• Axes terete, alternate-opposite or paniculately branched, texture cartilaginous but not very rigid, 1-1.5 mm thick, arising from flexuous axis, 0.5-1 mm thick.
• Outer cortical cells subquadrate in cross-section, with lateral secondary pit connections and bright cell inclusions (cherry bodies), not radially elongated and not projecting.
• Medullary cells with lenticular thickenings.
• Tetrasporangial arrangement parallel.

L. obtusa.

• Thalli erect, up to 10 cm tall, deep purple, reddish, or greenish with pinkish tips, soft and fleshy in texture.
• Stipe short, terete, with many percurrent terete axes clothed with irregularly alternate to subverticillate branches; ultimate branchlets simple, paniculate or not, distinctly clavate, constricted at base, trichoblasts present.
• Outer cortical cells cuboidal to rounded in cross-section, with secondary pit connections and bright cell inclusions (cherry bodies) and not projecting.
• Medullary cells large, polygonal, without lenticular thickenings.
• Tetrasporangia arranged in parallel pattern.

L. okamurae.

• Thalli large, robust, purplish-green to dark purple, up to 20 cm tall, fleshy to cartilaginous, not very rigid, with percurrent axes.
• Axes and branches terete, paniculately branched; branching alternate, opposite or verticillate.
• Outer cortical cells not radially elongated, with secondary longitudinal pit connections and with cherry bodies, not projecting.
• Medullary cells with lenticular thickenings.
• Tetrasporangial arrangement of the parallel type, in terminal cylindrical branchlets; spermatangia on characteristically broadened ultimate branchlets, 6.7-9.7 μm × 4.2-5.6 μm; cystocarps on lateral surface of terminal branchlets, up to 820 μm in diameter.

• L. palisada . Thalli 5-15 cm tall, dark red, rigidly cartilaginous, heavily to moderately clustered. Primary axes 1-3 mm thick, terete, naked below, from middle upwards bearing short, clavate-truncate branchlets. Outer cortical cells elongate in cross-section (= palisade arrangement), radial, not projecting, without secondary pit connections or cherry bodies. Medullary cells without definite arrangement, roundish, up to 23 μm. Tetrasporangia in right-angled pattern on tetrasporangial branchlets, 120-140 μm × 100-120 μm; spermatangia in cup-shaped pits on ultimate and penultimate branchlets, ellipsoidal, 9-12 μm × 6-7 μm; cystocarps laterally on ultimate branchlets, solitary or in small groups, flask-shaped, 520-800 μm × 580-800 μm, with ostiole on 100-220 μm long neck.

L. papillosa.

• Thalli dark brown or purple, erect or partly prostrate.
• Main axis terete, cartilaginous, up to 1 mm in diameter; branching radial with up to 3 orders of branching; ultimate branches simple to compound, densely disposed, wart-like, generally 40-50 per 1 cm from apex, up to 1.1 mm in diameter, up to 2.8 mm long.
• Cortical cells in palisade arrangement, not projecting, without cherry bodies, without secondary pit connections.
• Medullary cells with relatively thick cell walls, with only occasional lenticular thickenings.
• Tetrasporangiate branches simple to compound, up to 1 mm in diameter; tetrasporangia in radial arrangement (= right-angled type); spermatangia formed in cup-shaped pits at apices of ultimate, penultimate branchlets; cystocarps globose with mamillate ostiole, up to 1 mm in diameter.

L. parvipapillata.

• Thalli spreading, dark red to dull purple, semi-prostrate, up to 4.5 cm tall, cartilaginous.
• Main axes up to 800 μm thick, 2 mm wide, with subopposite short branchlets along both sides of main branches; branches usually distichous, compressed, especially near basal portion, connected to other branches by tenacula.
• Outer cortical cells in cross-section 20 μm in diameter, radially elongated (= palisade pattern), protruding, with sporadic secondary pit connections and bright cell inclusions (cherry bodies).
• Medullary cells up to 100 μm in diameter, without lenticular thickenings.
• Tetrasporangia arranged in right-angled pattern; tetraspores (in culture) 80-100 μm in diameter.

L. patentiramea.

• Thalli green-red to brown-red, firm, rigid, decumbent or matted, sometimes forming cushions, attached to one another by hapteres.
• Main axis terete, fleshy, up to 2-3 mm in diameter; branching frequent, radial, with up to 4 orders of branching; ultimate branches terete, up to 1.8 mm in diameter, simple to compound or wart-like, closely arranged on branches.
• Outer cortical cells in palisade arrangement, not projecting, without cherry bodies, not connected to each other by secondary pit connections.
• Medullary cells with relatively thin cell walls and no lenticular thickenings.
• Tetrasporangiate branchlets simple to compound, with tetrasporangia in radial arrangement (= right-angled pattern); tetrasporangia 80-100 μm in diameter; spermatangia on short, branched spermatangial branchlets; cystocarps sessile, conical, 0.75-1 mm in diameter, single or with two on a branchlet.

L. similis.

• Thalli up to 15 cm tall, pale red, cartilaginous, with 1-2 terete, erect axes.
• Branching somewhat patent, irregularly alternate, subopposite, often repeatedly verticillate or subverticillate, from percurrent axis; ultimate branchlets densely covering all but lower axes, shortly cylindrical or clavate to wart-like, 0.3-0.5 mm in diameter, with truncate, swollen apices.
• Outer cortical cells quadrangular in cross-section, not projecting, connected by longitudinal secondary pit connections, containing two cherry bodies in each cell.
• Medullary cells without lenticular thickenings.
• Tetrasporangial branchlets cylindrical to cylindro-clavate, simple or compound, 0.4-0.6 mm in diameter, truncate, slightly swollen at the apices; tetrasporangia 100-120 μm in diameter, arranged in right-angled manner; spermatangia in cup-shaped pits on ultimate and penultimate branchlets; cystocarps on short branches, broadly ovoid, 640-800 μm × 680-840 μm.

L. snackeyi.

• Thalli greenish-purple to red, erect, up to 22(-50) cm tall.
• Main percurrent axis terete, firm, fleshy, up to 3.2 mm in diameter.
• Branching irregularly radial, with up to 4 orders of branching; ultimate branchlets simple to compound, up to 1 mm in diameter, very fragile when dried or preserved.
• Outer cortical cells usually not in palisade arrangement, although occasionally radially elongated in lower portions of branches, not projecting, with one cherry body per cell, connected by longitudinal secondary pit connections.
• Medullary cells with relatively thin cell walls, without lenticular thickenings.
• Tetrasporangial branchlets simple or compound, up to 0.6 mm in diameter; tetrasporangia in parallel arrangement, 120-160 μm in diameter; male stichidia 600-800 μm in diameter; spermatangia in apex in deep pits; cystocarps broadly ovoid, 640-880 μm × 640-960 μm, laterally on branches of every order.

L. tronoi.

• Thalli dark brown to purple, cartilaginous, with erect, decumbent branches.
• Branching irregular, somewhat secund or alternate; branches terete, gradually tapering towards the tips; ultimate branchlets 1.0-1.6 mm long, partly simple, cylindrical, partly bilobed, trilobed or paniculate.
• Outer cortical cells cuboidal, somewhat rounded to angular in cross-section, without secondary pit connections, not projecting.
• Medullary cells without lenticular thickenings.
• Tetrasporangia oval, borne on tip of tetrasporangial branchlets, arranged at right angles to axis of ultimate branchlets.

Growth and development

In culture, the compressed condition of thalli of Laurencia persists. Tetraspores are usually formed during the whole year and tetrasporophytes dominate over gametophytes.

Other botanical information

Species of Laurencia are notably difficult in their taxonomy and many different classifications and taxon definitions are in use. It is suggested that material from South-East Asia identified as L. botryoides most probably belongs to L. cartilaginea. It has been proposed that L. palisada , together with material identified as L. intermedia Yamada in South-East Asia belong to L. glandulifera (Kütz.) Kütz. For L. tronoi synonymy with L. glandulifera has also been suggested. However, in recent references, the presence of longitudinally oriented secondary pit connections in L. glandulifera , and parallel arrangement of its tetrasporangia exclude possible synonymy with L. patentiramea and L. tronoi.

As a result of recent research, former subgenera within Laurencia have been raised to generic status. Of these new genera, Chondrophycus (Tokida & Y. Saito) Garbary & J.T. Harper includes a number of taxa that have been described here as Laurencia spp.: Chondrophycus cartilaginea (Yamada) Garbary & J.T. Harper, C. palisada (Yamada) K.W. Nam, C. papillosa (C. Agardh) Garbary & J.T. Harper, C. parvipapillata (C.K. Tseng) Garbary & J.T. Harper, C. patentiramea (Mont.) K.W. Nam and C. tronoi (Ganz.-Fort.) K.W. Nam.

Many specimens in South-East Asia have been identified as L. papillosa although the anatomical structure is very different. In culture, it is very distinct: discoid holdfasts and very short suppressed branchlets which are densely disposed. In the field, the plants are less distinct morphologically, but are easy to differentiate from the papillose forms of L. patentiramea by making cross-sections and looking for cherry bodies in the cortical cells. Dried specimens are harder to differentiate, but if well rehydrated, medullary cell walls should be thicker than in L. patentiramea.

Ecology

Laurencia is found attached to many substrates, including rocks, limestone, corals, sponges, seagrasses, and algae, including other Laurencia spp. They can be found both in the subtidal and in the intertidal zones. Laurencia spp. form one of the more common components of seaweed communities on reefs. In seagrass beds, especially L. patentiramea can form mats.

There seem to be two ecological forms of L. papillosa. The more common one grows on sand in the seagrass bed or on limestone in quiet waters in the upper and lower intertidal and subtidal zones in central and southern Philippines. It has prostrate branches with very short, densely disposed papillae-like ultimate branchlets. In older branches, these ultimate branches become compound due to the initiation of branchlets but with very little elongation. This form is hard to distinguish from L. patentiramea unless fresh specimens are examined for the presence of cherry bodies. The other form is common in the northern Philippines, especially in the Batanes area, growing in clumps on rocks and sponges in the high-energy wave zone. The plants were heavily encrusted with the cyanobacterium Lyngbya and with crustose corallines.

Propagation and planting

Laurencia is not known in commercial phycoculture.

Diseases and pests

In the Caribbean, Laurencia is often grazed by the opistobranch sea hares (Aplysia spp.), which concentrate the metabolites of the algae in their digestive glands and possibly use them as a defence against predators. These sea hares cannot survive in culture without Laurencia.

Handling after harvest

Laurencia algae can be used fresh or dried.

Prospects

Although Laurencia has been much studied by biochemists, no promising chemical compounds for future commercial exploitation have been identified.

Literature

• Cordero, P.A., 1977. Studies on Philippine marine red algae. Special Publication from the Seto Marine Biological Laboratory, series 4: 1-258.
• Ganzon-Fortes, E. & Trono, G.C., 1982. Reproductive morphology and periodicity of Laurencia sp. at Calatagan, Batangas, Philippines. Kalikasan 11: 27-38.
• Garbary, D.J. & Harper, J.T., 1998. A phylogenetic analysis of the Laurencia complex (Rhodomelaceae) of the red algae. Cryptogamie, Algologie 19: 185-200.
• Masuda, M., Abe, T., Suzuki, T. & Suzuki, M., 1996. Morphological and chemotaxonomic studies on Laurencia composita and L. okamurae (Ceramiales, Rhodophyta). Phycologia 35: 550-562.
• Masuda, M., Kawaguchi, S. & Phang, S.M., 1997. Taxonomic notes on Laurencia similis and L. papillosa (Ceramiales, Rhodophyta) from the Western Pacific. Botanica Marina 40: 229-239.
• Masuda, M., Kogame, K., Abe, T. & Kamura, S., 1998. A morphological study of Laurencia palisada (Rhodomelaceae, Rhodophyta). Botanica Marina 41: 133-140.
• Masuda, M., Takkahashi, Y., Okamoto, K., Matsuo, Y. & Suzuki, M., 1997. Morphology and halogenated secondary metabolites of Laurencia snackeyi (Weber-van Bosse) stat. nov. (Ceramiales, Rhodophyta). European Journal of Phycology 32: 293-301.
• McDermid, K.J., 1988. Introduction to Laurencia (Rhodophyta, Rhodomelaceae). In: Abbott, I.A. (Editor): Taxonomy of economic seaweeds 2. pp. 221-229.
• Nam, K.W., 1999. Morphology of Chondrophycus undulata and C. parvipapillata and its implications for the taxonomy of the Laurencia (Ceramiales, Rhodophyta) complex. European Journal of Phycology 34: 455-468.
• Xia, B.M. & Zhang, J.F., 1982. A new raw material for the manufacture of agar-agar from Hainan Island, China. Oceanology and Limnology 13(6): 538-543.
• Zhang, J. & Xia, B., 1988. Laurencia from China: key, list and distribution of the species. In: Abbott, I.A. (Editor): Taxonomy of economic seaweeds 2. pp. 249-252.

Sources of illustration

Ganzon-Fortes, E.T., 1982. Laurencia tronoi (Rhodophyta: Ceramiales), a new species from Calatagan, Batangas, Philippines. Kalikasan 11: Fig. 1, p. 407 (L. tronoi); Saito, Y., 1965. Morphology of the genus Laurencia of Japan. 2. Bulletin of the Faculty of Fisheries, Hokkaido University 15: Plate 5, fig. 5 (L. okamurae); Saito, Y., 1982. Morphology of British Laurencia (Rhodophyta). Phycologia 21: Fig. 7 & 8, p.301 (L. obtusa); Trono, G.C., 1986. Philippine seaweeds. In: Guide to Philippine flora and fauna. Vol. 1. Natural Resources Management Center, Ministry of Natural Resources and University of the Philippines. Goodwill Bookstore, Manila, The Philippines. Fig. 94, p. 282 (L. papillosa); Trono, G.C. & Ganzon-Fortes, E.T., 1980. An illustrated seaweed flora of Calatagan, Batangas, Philippines. University of the Philippines Marine Science Center & Filipinas Foundation, Manila, The Philippines. Fig. on p. 101 (L. cartilaginea). Redrawn and adapted by P. Verheij-Hayes.

Authors

• W.F. Prud'homme van Reine