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Studies of Marine Algae in the Lesser-Known Families of the Gigartinales (Rhodophyta)

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Studies of Marine Algae in the Lesser-Known Families of the Gigartinales (Rhodophyta) Aust. J. Bot., 1977, 25, 97-140 Studies of Marine Algae in the Lesser-known Families of the Gigartinales (Rhodophyta). I. The Acrotylaceae Gerald T. Kraft School of Botany, University of Melbourne, Parkville, Vic. 3052. Contents Page Abstract ............................................................................................................................................................ 97 Introduction .................................................................................................................................................... 98 Methods ............................................................................................................................................................99 Terminology ..................................................................................................................................................... 99 Descriptions of taxa ..................................................................................................................................... 100 Key to the genera and species of the Acrotylaceae ...................................................................... 100 Family Acrotylaceae ............................................................................................................................... 100 1. Genus Acrotylus ............................................................................................................................... 100 2. Genus Ranavalona ............................................................................................................................ 107 3. Genus Hennedya ............................................................................................................................. 111 4. Genus Amphiplexia ......................................................................................................................... 116 Discussion of Amphiplexia ............................................................................................................. 123 5. Genus Reinboldia ............................................................................................................................ 123 Discussion of the Acrotylaceae ......................................................................................................... 126 Acknowledgments ........................................................................................................................................127 References ....................................................................................................................................................... 128 Abbreviations used in figures ................................................................................................................... 130 Index to taxa and synonyms ................................................................................................................... 131 Abstract The Acrotylaceae is composed of Acrotylus australis J. Ag., Hennedya crispa Harv., Amphiplexia hymenocladioides J. Ag. and A. racemosa (J. Ag.) comb, nov. (all endemic to southern Australia), Reinboldia polyearpa Schmitz from South Africa and Ranavalona duckerae gen. et sp. nov. from southern Madagascar. The species are multiaxial, zonately tetrasporangiate and monoecious. In Acrotylus and Ranavalona the tetrasporangia develop in raised nemathecia, whereas they are scattered in the other species. In Amphiplexia hymenocladioides, tetrasporangia are intercalary in contrast to the other species with known tetrasporophytes. Acrotylus and Ranavalona are polycarpo- gonial, with the carpogonial branches of Ranavalona being additionally nemathecial. The remaining genera are monocarpogonial, with scattered carpogonial branches. The carpogonia in all species are directed laterally or thallus-inwardly and provided with reflexed trichogynes. The species are all procarpic (except possibly Reinboldia), with supporting cells of carpogonial branches functioning as auxiliary cells. Fusion cells do not form, and several gonimoblast initials arise directly from diploidized auxiliary cells and enter an adjacent region of 'nutritive' cells. Gonimoblast development is thallus- outward in Amphiplexiu and thallus-inward in Acrotylus, RQ~QVQ~O~Q,Hennedya and possibly Reinboldia, the last genus being known only from a surviving fragment of the type specimen. In all but Amphiplexia, breakdown of the auxiliary cell, some adjacent cortical cells and some older goni- moblasts results in a cavity which is lined by gonimoblast derivatives connected to surrounding nutritive and other vegetative cells. With expansion of the cavity, branched gonimoblast filaments grow centripetally and produce terminal or short chains of carposporangia that are released through distinct ostioles. In Amphiplexiu, nutritive cells do not completely surround the auxiliary cell, and most gonimoblast filaments initially cover the interior surface of the pericarp rather than a cavity lined by specially produced nutritive cells. G. T. Kraft The genera are separated on wide differences in habit, vegetative cross section, carpogonial branches, gonimoblast orientation, carpospore morphology and mature cystocarp cross sections. The two African genera Ranavalona and Reinboldia appear on vegetative and certain reproductive grounds to be relatively primitive in the group, with possible links to the less reproductively complex family Solieriaceae. The most closely related genera within the Acrotylaceae, Acrotylus and Ranavalona, are also about the most geographically separated. The present-day distinctiveness, uncommonness and distribution of the genera of the Acrotylaceae perhaps suggest that it is a relict and isolated group. Introduction Red algae in the diverse order Gigartinales have become better known in recent years through detailed morphological studies of several of the 20 to 23 families (Feldmann 1954; Kraft 1973; Mikami 1965; Min-Thein and Womersley 1976; Schotter 1968; Searles 1968). The distinguishing criteria of the families are still largely those of Kylin (1956, p. 238), who placed major emphasis on whether (a) the gonirnoblast grows from the auxiliary cell or from the connecting filament at some remove from the auxiliary cell; (b) the gonimoblast grows from the auxiliary cell towards the centre of the thallus or towards its surface; (c) the tetrasporangia are cruciately or zonately divided; (d) the thalli are uni- or multiaxial; (e) several gonimoblast initials, or only one, are initiated by the diploidized auxiliary cell; (f) the female gametophytes possess or lack procarps; (g) the vegetative structure of mature axes is pseudoparenchymatous throughout or at least partly filamentous; and (h),in a few instances, whether the carpo- sporangia form in chains or as single cells at the ends of sporogenous filaments in the cystocarp. Since Kylin's time, however, doubts have been raised about the absolute importance of some of his major family-splitting characters (Kraft 1973, 1975a; Min-Thein and Womersley 1976; Searles 1968) which demonstrate the need for further, detailed work on the many remaining unstudied genera and families. Although there have been recent attempts to rank some of the gigartinalean families phylogenetically (Kraft 1973, 1975a, 1975b; Min-Thein and Womersley 1976), such speculations suffer from a lack of precise, comparative knowledge of vegetative and reproductive morphology within a large proportion of the order. Along the coasts of southern Australia there are representatives of at least 16 of the 20 to 23 gigartinalean families. The number of endemic species belonging to the order is high in this region, as is the case with the other red algal orders (Womersley 1959). Several entire families are virtually confined to southern Australia, among which are the poorly known Acrotylaceae, Dicranemaceae and Mychodeaceae. These were thought by Kylin (1956) to form a distinctive branch of the Gigartinales, along with the more cosmopolitan Gigartinaceae and Phyllophoraceae. All five families were said to exhibit highly condensed procarps in which the supporting cell of the carpogonial branch presumably connects directly with the fertilized carpogonium and becomes the auxiliary cell from which multiple gonimoblast initials issue that are directed towards the centre of the thallus. These reproductive features alone serve to isolate the five families from the rest of the order. The Acrotylaceae, Dicranemaceae and Mychodeaceae of Kylin (1956) are distinguished from the Gigartinaceae and Phyllophoraceae by their zonate, rather than cruciate tetrasporangia. The Mychodeaceae is also distinctive among the five families in being uniaxial, while the multiaxial Acrotylaceae and Dicranemaceae are separated from each other on vegetative features and the shape of their mature carposporophytes. Families of the Gigartinales. I Studies of the Gigartinaceae by Mikami (1965) and the Phyllophoraceae by Schotter (1968) indicate that these groups are quite specialized within the order and are without obvious links to families with lower levels of carposporophyte organization. Since comparable details of vegetative and reproductive morphology have not been available for genera of the Acrotylaceae, Dicranemaceae and Mychodeaceae, I have attempted monographic studies of these families, based on an abundance of fresh material of most known species. This paper reports investigations of the Acrotylaceae, with work on the Dicranemaceae and the Mychodeaceae
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