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A Reappraisal of the Order Corallinales (Rhodophyceae)

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A Reappraisal of the Order Corallinales (Rhodophyceae) British Phycological Journal ISSN: 0007-1617 (Print) (Online) Journal homepage: http://www.tandfonline.com/loi/tejp19 A reappraisal of the order Corallinales (Rhodophyceae) Paul C. Silva & H. William Johansen To cite this article: Paul C. Silva & H. William Johansen (1986) A reappraisal of the order Corallinales (Rhodophyceae), British Phycological Journal, 21:3, 245-254, DOI: 10.1080/00071618600650281 To link to this article: https://doi.org/10.1080/00071618600650281 Published online: 23 Feb 2007. Submit your article to this journal Article views: 318 View related articles Citing articles: 48 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tejp19 Br. phycol. J. 21:245 254 1 September 1986 A Reappraisal of the Order Corallinales (Rhodophyceae) By PAUL C. SILVA Department of Botany, University of California, Berkeley, California 94720, USA and H. WILLIAM JOHANSEN Biology Department, Clark University, Worcester, Massachusetts 01610, USA Articulated and non-articulated coralline alga e were brought together in the family Corallinaceae in essentially its present-day circumscription by Decaisne in 1842. Since that time, this family has been perceived as one of the most distinctive assemblages of Rhodophyceae. Alignment of families of red algae into orders based on criteria that today are considered to reflect natural relationships extends back only as far as 1892, when Schmitz presented a scheme founded largely on details of the female reproductive system and gonimoblast development. In that scheme, the Corallinaceae occupied an anomalous position in the Cryptonemiales. While attempts have been made to modify the definition of the Cryptonemiales to accommodate the Corallinaceae more comfortably, an ongoing accumulation of information supports the segregation of that family into its own order. At least three previous authors have adopted the taxonomic concept of the Corallinales, but the name has not yet been validated. In the present paper a Latin diagnosis is provided. Diagnostic characters of the Corallinales include the following: (1) walls of most vegetative cells are impregnated with calcite; (2) meristems are often intercalary and covered by a layer of cells; (3) plugs of primary pit-connections have two-layered, dome-shaped caps; (4) reproductive structures are produced in roofed conceptacles in all genera but one; (5) tetrasporocytes usually undergo simultaneous zonate division; (6) post-fertilization events involve a cluster of procarpial filament systems. HISTORICAL BACKGROUND lated forms had to be distinguished from corals. Second, the circumscription of For more than a century the coralline coralline algae had to be shorn of unrelated algae have been perceived by taxonomists as calcified algae (the red alga Galaxaura and one of the most distinctive assemblages of various siphonous Chlorophyceae). Third, Rhodophyceae. Prior to the period 1842- agreement had to be reached on the close 1853, when this perception was solidified, relationship between articulated and non- the taxonomic history of this assemblage articulated forms. As might be expected, the followed three interrelated lines. First, latter, being rocklike or coralloid, were agreement had to be reached on the nature accepted as plants less readily than the of corallines--whether they were plants or former. animals, or organisms intermediate between The two groups, articulated and non- animals and plants (zoophytes), or articulated, were brought together in organisms intermediate between minerals essentially their present-day circumscriptions and plants (lithophytes). Those with arti- by Decaisne (1842a, pp. 359, 365; 1842b) culated fronds had to be distinguished from and Chauvin (1842, pp. 109, 127), who sessile animals with flexible bodies, especially assigned them to the Corallinaceae, a family hydroids and bryozoans, while non-articu- established by Lamouroux (1812, p. 185, 0007-1617/86/030245 + 10 $03.00/0 © 1986 British Phycological Society 1 Published online 23 Feb 2007 246 P.C. Silva and H. W. Johansen "Corallineae") to include Galaxaura and (1837) and Lithothamnium Philippi (1837)]. calcified siphonous Chlorophyceae as well as Subsequent authors generally have Amphiroa, Corallina, Jania, and Melobesia. recognized only one family, Corallinaceae. Decaisne's belief that reproductive structures Exceptions include J. E. Gray (1864, p. 22) were of primary taxonomic significance led and his nephew S. O. Gray (1867, p. 133), to success in sorting out the true corallines, who recognized the family Hapalidiaceae, which he placed in his Algae based on Hapalidium Kiitzing (1843), a Choristosporeae alongside other genera now genus said to differ from the Corallinaceae assigned to the Florideophycidae. Lack of s. str. by its lack of calcification. Hapalidium knowledge regarding reproduction in calci- roseolum Kfitzing, the type of the genus, is in fied siphonous Chlorophyceae, however, led fact slightly calcified and is conspecific with to the dispersal of these genera between two Melobesia membranacea (Esper) Lamouroux, major categories, Algae Zoosporeae and according to Chamberlain (1983, p. 300). Algae Aplosporeae. A detailed account of Much later, Hylander (1928, p. 173), in a the taxonomic vicissitudes of coralline algae flora of Connecticut, assigned was given by Decaisne in the second part of Lithothamnium and Melobesia (and his thesis (1842b). presumably all other non-articulated Johnston, a British contemporary of corallines) to the Squamariaceae Decaisne, offered a contrasting treatment in (= Peyssonneliaceae). In recent years rather which the family Corallinaceae was reserved elaborate classifications have been proposed for articulated forms (erroneously including within the Corallinaceae (e.g., Cabioch, Halimeda), while non-articulated forms were 1971, 1972; Lebednik, 1977; Johansen, segregated as a distinct family, 1981). Nulliporaceae Johnston (1842, pp. 205, 231, 254, "Nulliporidae"). (Nullipora Lamarck TAXONOMIC POSITION OF (1801) is a superfluous name for Apora CORALLINACEAE Gunnerus (1768), the oldest available name for a genus of non-articulated corallines. Alignment of families of red algae into Johnston's recognition of four species of orders based on criteria that today are Nullipora in the British flora was purely considered to reflect natural relationships academic, however, as made clear by the extends back only as far as 1892, when following passage: "I believe that the Schmitz presented a scheme founded largely Nullipores are endowed with life, and with a on details of the female reproductive system vegetative growth, but that they are not and gonimoblast development. The four species. They are, to my conviction, merely orders of subclass Florideae proposed by the calcareous basis of Corallina officinalis Schmitz (Nemaliales, Gigartinales, modified by external circumstances, and Rhodymeniales, and Cryptonemiales, in that hindered from passing through the stages of sequence) are presently accepted, but with its normal developement"). Johnston's numerous modifications of varying account of the taxonomic history of the consequence, most notably the segregation corallines (Johnston, 1842, pp. 207 215, of the Ceramiales from the Rhodymeniales 233 237) is couched in colourful language by Oltmanns (1904, p. 683) and the segrega- lightly brushed with cynicism. tion of the Gelidiales from the Nemaliales by Non-articulated corallines were also Kylin (1923, p. 132). (The ordinal name placed in a distinct family by Kfitzing, who, Ceramiales should be credited, however, to however, employed the family name N~igeli, 1847, pp. 126, 253, "Ceramiaceae".) Spongitaceae Kfitzing (1843, pp. xxiii, 382, Among subsequently segregated orders, 385, "Spongiteae"). [Spongites was Nemastomatales Kylin (1925) and proposed by Kfitzing in 1841 to apply to a Sphaerococcales Sj6stedt (1926) have been genus that combined Lithophyllum Philippi abandoned (at least temporarily), while Reappraisal of the order CoraUinales 247 Bonnemaisoniales J. Feldmann et G. cells in a sorus become connected to form a Feldmann (1942), Acrochaetiales J. fusion cell from which gonimoblasts arise. Feldmann (1953), and Chaetangiales Whether ooblast filaments are involved was Desikachary (1963) have received negligible not known by Schmitz and remains a support. The recently proposed Palmariales controversial issue (Johansen, 1981, p. 105). Guiry et D. Irvine (in Guiry, 1978) is likely In deciding which of Schmitz's orders to fare better because of the very distinctive would most comfortably receive the life history exhibited by Palmaria and Corallinaceae, it is essential to clarify the Halosaccion (van der Meer & Todd, 1980; meaning of the term "procarp". This term van der Meer, 1981). Only the passage of was applied by Thuret & Bornet (1878) to time will reveal the fate of the fledgling the female reproductive system of Batrachospermales and Hildenbrandiales, Ceramiaceae and Rhodomelaceae at the both proposed by Pueschel & Cole (1982). stage of readiness for fertilization. It In addition to the recognition of new orders, included the supporting cell, the carpogonial there has been considerable movement of branch, and the auxiliary cell. Schmitz genera and families from one order to (1883) applied the term broadly to another, especially between the Gigartinales "Gruppen von Carpogon/isten und and the Cryptonemiales. Auxiliarzellen, die als selbstfindiges Ganzes Schmitz (1892) gave the following
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