The Red Algal Genus Audouinella Bory Nemaliales: Acrochaetiaceae) from North Carolina SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 22 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES-NUMBER 22 The Red Algal Genus Audouinella Bory (Nemaliales: Acrochaetiaceae) from North Carolina Craig W. Schneider SMITHSONIAN INSTITUTION PRESS City of Washington 1983 ABSTRACT Schneider, Craig W. The Red Algal Genus Audouinella Bory (Nemaliales: Acrochaetiaceae) from North Carolina. Smithsonian Contributions to the Marine Sciences, number 22, 25 pages, 3 figures, 1983.—A monographic study of Audouinella in the Atlantic waters of North Carolina reports 15 taxa from coastal and continental shelf habitats. Three of these, A. affinis, A. hoytii, and A. ophioglossa, herein described as a new species, are endemic. Audouinella bispora and A. daviesii are reported from the Carolina flora for the first time. The taxonomy from several historical reports is elucidated, and taxonomic confu- sions in this complex are clarified. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the Atlantic coast of eastern North America. Library of Congress Cataloging in Publication Data Schneider, Craig W. The red algal genus Audouinella Bory (Nemaliales: Acrochaetiaceae) from North Carolina. (Smithsonian contributions to the marine sciences ; no. 22) Bibliography: p. Supt. of Docs, no.: SI 1.41:22 1. Audouinella—Classification. 2. Algae—North Carolina—Classification. I. Title. II. Se- ries. QK569.A27S36 1983 589.4'1 83-600261 Contents Page Introduction 1 Materials and Methods 2 Acknowledgments 2 Genus Audouinella 2 Key to the Species of Audouinella from North Carolina 2 Audouinella affinis (Howe and Hoyt), new combination 4 Audouinella bispora (B0rgesen) Garbary 5 Audouinella botryocarpa (Harvey) Woelkerling 5 Audouinella corymbifera (Thuret) Dixon 6 Audouinella dasyae (Collins) Woelkerling 7 Audouinella daviesii (Dillwyn) Woelkerling 8 Audouinella densa (Drew) Garbary 8 Audouinella hallandica (Kylin) Woelkerling 9 Audouinella hoytii (Collins), new combination 10 Audouinella hypneae (B0rgesen), new combination 11 Audouinella infestans (Howe and Hoyt) Dixon 12 Audouinella microscopica (Nageli) Woelkerling 12 Audouinella ophioglossa, new species 13 Audouinella saviana (Meneghini) Woelkerling 15 Audouinella secundata (Lyngbye) Dixon 16 Literature Cited 17 Figures 1-3 20 in The Red Algal Genus Audouinella Bory (Nemaliales: Acrochaetiaceae) from North Carolina Craig W. Schneider Introduction The Acrochaetiaceae has received a great deal of systematic attention in the last half century (Drew, 1928; Hamel, 1928b; Papenfuss, 1945, 1947; Abbott, 1962, 1968; Feldmann, 1962; Woelkerling, 1971, 1972, 1973a, 1973b; Dixon and Irvine, 1977; Garbary, 1978, 1979). Never- theless, many of the species reported or described from North Carolina have received little or no attention since their early notation here (Howe and Hoyt, 1916; Hoyt, 1920; Williams, 1948, 1951). The only taxa updated in a recent sys- tematic listing for North Carolina (Searles and Schneider, 1978) were those species taxonomi- cally treated in other floras. Therefore, the North Carolina checklist contained both recent and an- tiquated taxonomy in this family, as well as sev- eral unsubstantiated reports. This monograph was undertaken to verify and update the taxo- nomic record. In the process of this work addi- tional taxa have been discovered in the North Carolina flora. Several genera have been described in Acro- chaetiaceae and species have been variously as- signed to them on the basis of chromoplast mor- phology and presence or absence of sexual repro- duction (for example, see Garbary, 1979). Because Craig W. Schneider, Department of Biology, Trinity College, Hart- ford, CT 06106. generic concepts vary from study to study and because frequently utilized generic characters have proved unreliable (Woelkerling, 1971, 1973b), I find that it is more pragmatic and less confusing to follow the single-genus concept for the Acrochaetiaceae previously utilized by B0r- gesen (1915) and first argued for by Drew (1928). This approach has recently been followed by Dixon and Irvine (1977) and Garbary (1979). The genus with historical precedence is Audoui- nella Bory (1823:340), and its usage requires some new nomenclatural combinations herein pro- posed. With the advent of further culture work in this family, it is possible that in the future we will be able to resegregate the species into more than one genus based on more reliable characters, such as life history characteristics (West, 1968, 1969; Stegenga, 1978; Stegenga and van Wissen, 1979). Generic confusion aside, species concepts have of late also come under more critical examination. Several a priori standard specific characters such as mode of attachment and reproductive state have been criticized as taxonomic criteria (Borsje, 1973; Stegenga and Borsje, 1976, 1977; Stegenga and Vroman, 1976; Stegenga and Mulder, 1979; Stegenga and van Erp, 1979). Broad circumscrip- tions of species utilized by some (e.g., Woelker- ling, 1971, 1972, 1973a, 1973b; Stegenga and Vroman, 1976) are not universally accepted (e.g., Dixon and Irvine, 1977; Kornmann and Sahling, 1 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 1978; Humm, 1979). The cultural experiments of Stegenga and his associates have paired "distinct" species as heteromorphic generations of one taxon in this family. For the present, however, I con- tinue to follow Woelkerling's classical interpreta- tion of species concepts within Audouinella until a more complete understanding of species based upon life history and other features can be syn- thesized. Included in the scheme listed below, for example, is Audouinella dasyae, which is probably a gametophytic generation of either A. saviana, A. hypneae, or A. bispora, or possibly, all of them (Stegenga and Borsje, 1976). One of the asexual taxa mentioned below could possibly be the spo- rophytic generation of Scinaia complanata (Collins) Cotton (van den Hoek and Cortel-Breeman, 1970), though that remains to be determined. MATERIALS AND METHODS.—Where possible, observations, measurements, and drawings were made using living field-collected material or liq- uid-preserved material. Dried specimens were wetted and removed from herbarium sheets using 10% aqueous sodium carbonate. Permanent slides were produced by either mounting the specimens in 20% Karo syrup, 1% aniline blue, 1 N HC1, 3%-4% formalin in a ratio of 95: 1:2:2, or by staining with 1% aniline blue, fixing with 1% HC1, and mounting in 20% Karo with phenol added as a preservative. Drawings were made using a Zeiss camera lucida. Aside from those listed below for DUKE and US, duplicates of some of the author's collection numbers have been deposited in AHFH, GALW, HBFH, MASS, MICH, NCU, NFLD, NHA, and WNC. Herbarium abbreviations follow Holmgren et al. (1981). Specimen measurements are accurate to 0.1 micrometer. ACKNOWLEDGMENTS.—I thank Duke University for space and assistance at the Marine Laboratory and Dr. W. Kirby-Smith of the Reference Mu- seum for sponsoring an offshore collecting trip and allowing access to the collections. I acknowl- edge the Duke University Cooperative Program in Oceanography as some of my earlier collections were made on R/V Eastward cruises. I also thank fellow scientists (and the herbaria with which they are associated): Drs. R.B. Searles (DUKE), J.N. Norris (US), and D.F. Kapraun (WNC) for loans; Dr. L.G. Williams for sending specimens; and the Duke University Botany Department for access to that herbarium. Dr. M.H. Hommersand assisted with the literature and Dr. J.S. Ramus allowed use of his personal library. Ginny Schnei- der deserves special thanks for typing during my leave. Drs. Searles, Norris, and W.J. Woelkerling kindly reviewed the manuscript. This research was sponsored by Trinity College as a sabbatical leave. This monograph is dedicated to the memory of Dr. G.F. Papenfuss, both for his early work with the Audouinella complex and for his recent encour- agement for me to unravel the taxa in North Carolina. Genus Audouinella Key to the Species of Audouinella from North Carolina1 (Species alphabetically arranged in text) 1. Plants attached to the substrate by an obvious, persistent, undivided basal spore which may give rise to one or several accessory cells, an endophytic 1 Several specimens from a population should be observed when utilizing the above key. Some characters, such as presence or absence of unicellular hairs, are variable and by themselves are not considered to be taxonomically significant. In practice, however, a combination of several characters, some apparently more stable than others, is useful in delineating taxa. This key reflects distinctions found in North Carolina specimens; it should also be helpful in distinguishing these "purported" taxa elsewhere. NUMBER 22 branched filament, or a prostrate filamentous system 2 Plants attached to the substrate by a multicellular prostrate system, the basal spore once septate and persistent, or not obvious, obscured by divisions in the earliest stages 8 2. Plants minute, less than 250 jum tall A. microscopica Plants larger, greater than 500 jum tall 3 3. Basal spores consistently globose or subglobose, obviously larger than erect filaments 4 Basal spore ovoid to panduriform, globose or subglobose, the same size or only a few micrometers larger than erect filaments 7 4. Basal spore producing a single penetrating endophytic branched filament in addition to one or more erect filaments 5 Basal spore remaining undivided or occasionally producing a few accessory cells 6 5. Basal spore 12-15 jum diam., erect filaments 8-13 jum diam.; dioecious, spermatangia forming corymbose clusters on lateral branches A. corymbifera Basal spore 7.5-12.5 /xm diam., erect filaments 3-5 jum diam.; monoecious, spermatangia paired, produced on opposite or whorled branches in a long series A. ophioglossa 6. Monosporangia 5.0-7.5 jum diam., 11-15 jum long A. hoytii Monosporangia 10-18 jum diam., 18-27 jum long A. affinis 7. Basal spore remaining undivided, ovoid, elongate to rectangular, occa- sionally with a flared disciform base A. hallandica Basal spore, pyriform to panduriform, producing a prostrate filamentous system A. dasyae 8. Plants mostly endophytic or endozoic, upright axes less than 150 jum tall A. infestans Plants mostly epiphytic, epizoic, or saxicolous, upright axes greater than 200 jum tall 9 9. Plastids with more than one pyrenoid per cell A. botryocarpa Plastids with only one pyrenoid per cell 10 10. Sporangia, for the most part, borne in clusters of 3 or more on branched stalks at the base of lateral branches A. daviesii Sporangia, for the most part, borne singly or in pairs, not clustered at the base of lateral branches 11 11. Monosporangia 9-13 jum long 12 Monosporangia 15-40 jum long 14 12. Monosporangia 5-6 jum diam.; bisporangia, if present, 7-12 jum diam., 12-18 jum long A. bispora Monosporangia 6-10 jum diam.; bisporangia not present 13 13. Plants producing only single monosporangia, erect filaments to 1 mm tall arising from a uniseriate prostrate filament to small compact pseudo- parenchymatous disc A. hypneae Plants producing single or 2-3 seriate monosporangia, erect filaments to 0.5 mm tall arising from a wide-spreading, much-branched prostrate system, often with a persistent septate basal spore A. densa SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 14. Plastids stellate, parietal or axial, unicellular hairs common but not present on all plants, plants arising from an orbicular pseudoparenchy- matous disc that is often later obscured by proliferating filaments .... A. secundata Plastids lobate, parietal or axial, unicellular hairs absent or rare, but terminal cells occasionally taper sharply to the tips, plants not arising from an orbicular pseudoparenchymatous disc in early stages 15 15. Prostrate system a widespreading mass of uniseriate filaments from an enlarged central pyriform to panduriform cell that is persistent but often obscured in older plants, uniseriate prostrate filaments giving rise to numerous erect filaments A. dasyae Prostrate system a compact, often penetrating mass of uniseriate filaments, usually coalesced centrally to form a pseudoparenchymatous disc, basal spore not persistent A. saviana Audouinella affinis (Howe and Hoyt), new combination FIGURE \a,b Acrochaetium affine Howe and Hoyt, 1916:118, pi. 15.—Hoyt, 1920:471, pi. 69.—Searles and Schneider, 1978:100. Audouinella alariae sensu Kapraun, 1980:40, figs. 15-17. [Not (Jonsson) Woelkerling, 1973b:541, figs. 1-9] DESCRIPTION.—Plants epiphytic or epiendo- phytic, to 4 mm tall, caespitose, arising from a persistent, large, globose to subglobose spore, 14- 26 jum diam. including an obvious wall 2.5-5.0 jum thick, one half again as big as the cells which emanate from it; if embedded in host tissue, the spore becoming vertically elongate to subpyri- form or panduriform, if not, producing a subcy- lindric obtuse or truncate foot that penetrates to 24 jum, the extended spore 20-33 jum high; spore usually remaining simple, occasionally develop- ing a few smaller accessory cells, rarely producing short prostrate irregularly branched filaments up to 5 cells long; basal spore, and occasionally accessory cells or basal filament cells, producing 1-4 erect filaments, cells cylindrical, 5-14 jum diam., 17-78 jum long, each with a parietal lobate plastid and one pyrenoid; for the most part plants immediately branched from the distal end of the first cell of the erect filament, branching subdi- chotomous or subtrichotomous, appearing rigid below, more flexuous above, branching mostly subdichotomous to distinctly lateral, ultimate branches elongate-virgate, 3.0-5.5 jum diam., gradually tapering toward the apices, often ter- minating with inconspicuous hairs. Monosporangia infrequent, lateral, secund, or less commonly terminal on upper portions of the plant, sessile or on one-celled pedicels, ovoid, obovoid, or oblong, 10-18 jum diam., 18-27 jum long, occasionally on gametophytic plants. Monoecious, spermatangia in proximity to sessile carpogonia, lateral or terminal, sessile or pedicel- late, solitary or in groups of 2 or 3; carposporo- phytes common, 3-12 spored, carpospores 8-25 jum diam., 13-28 jum long. TYPE-LOCALITY.—Onslow Bay, North Caro- lina, USA. HOLOTYPE.—US 56005. DISTRIBUTION.—Endemic to North Carolina, as far as currently known. NORTH CAROLINA SPECIMENS STUDIED.—Ons- low Bay, reef, 11 Aug 1914, Radcliffe (Holotype, US 56005; Isotypes, US 33567, 33584, 33646; Isotypes, DUKE 111, 1458). Beaufort, Radio Is. jetty, 5 Jul 1960, Aziz (DUKE 109). Cape Look- out jetty, 23 Jul 1961, Aziz (DUKE 102, 108). Beaufort, Radio Is. jetty, 4 Aug 1961, Aziz (DUKE 103-107, 110, 120, 121). Onslow Bay, reef, 2 Aug 1971, Searles (DUKE 13190). Onslow Bay, reef, 20 Jul 1973, Schneider 718 (DUKE 12606, 12716; US 50776; CWS). Onslow Bay, reef, 11 Aug 1981, Schneider 2711 (DUKE 16525; US 88906; CWS). Beaufort, Radio Is. jetty, 20 NUMBER 22 Jul 1982, Schneider 2867 (CWS). REMARKS.—This species is an obvious epiphyte of Dictyota, Sargassum, and other algae from deep (to 30 m) offshore and from shallow subtidal habitats in the Beaufort and Wilmington areas, at times turning the host alga red with a complete or marginal covering of its fine red filaments. To date, found from June to August. Hoyt (1920) listed several morphological char- acters useful in distinguishing Audouinella affinis from the earlier described A. hoytii. Although they are generally helpful, I find the size of monospor- angia as the most reliable in the separation of these two closely allied taxa. A. affinis appears related to A. alariae (Jonsson) Woelkerling (1973b), but can be distinguished by the produc- tion of accessory cells from the basal spore and the larger monosporangia. A comparison of type material is needed to clarify the limits of species with persistent globose basal spores. Audouinella bispora (B0rgesen) Garbary FIGURE Ic-f Audouinella bispora (B0rgesen) Garbary, 1979:490. Chantransia bispora B0rgesen, 1910:178, fig. 1. DESCRIPTION.—Plants epiendophytic to 1.5 mm tall, arising from an irregularly ramified, spread- ing prostrate system that remains superficial or penetrates host tissue, prostrate cells cylindrical to irregular, to 17.5 jum in greatest dimension, all capable of producing an erect filament; erect filaments radially to irregularly branched and rebranched, infrequent below, more above, the branches sharply to narrowly angled from the axes that bear them, tapering toward the apices; cells cylindrical, 2.5-8.0 jum diam., 10.0-32.5 jum long in the main axes, 1.0-3.0 jum diam., in the ultimate segments, occasionally ending in long multicellular, lightly pigmented hair-like exten- sions, unicellular hairs unknown; each cell con- taining a well-developed parietal plastid with a single large pyrenoid. Monosporangia terminal or lateral on short branches to adaxial on longer axes, sessile or on one-celled pedicels, single or paired, narrow, ovoid to oblong, 5-6 jum diam., 9.5-12.5 jum long; bisporangia situated as monosporangia, single or paired, broad, ovoid to ellipsoidal, 7.5-11.5 jum diam., 12.5-17.5 jum long. Gametangia unknown. TYPE-LOCALITY.—St. Thomas, Virgin Islands. HOLOTYPE.—C. DISTRIBUTION.—North Carolina, St. Thomas (Virgin Islands). NORTH CAROLINA SPECIMENS STUDIED.—Ons- low Bay, reef, 30 June 1972, Searles (DUKE 16526). Onslow Bay, reef, 5 May 1981, Handsel, BLM 65818086 (Duke Marine Lab Reference Museum). REMARKS.—Collected to date from water 25- 30 m deep in Onslow Bay, May-June. This is the first report of this taxon outside the type-locality. The North Carolina specimens perfectly fit the descriptions of B0rgesen (1910, 1915), and I do not hesitate to place them under Audouinella bis- pora. The questions raised by B0rgesen (1915) as to whether this plant is indeed a form of Audoui- nella hypneae (B0rgesen) Schneider cannot be as- certained without additional comparative mate- rial of the latter named species (see discussion of A. hypneae). Audouinella bispora occasionally pro- duces bispores and has narrower monospores than reported for A. hypneae, a feature also seen in North Carolina plants. Nevertheless, it is interest- ing to note that in Garbary's (1979) cluster anal- ysis, A. bispora was found to be quite similar to A. seriata (B0rgesen) Garbary, a species I consider a synonym of A. hypneae in this paper. Audouinella hypneae was not treated by Garbary (1979). The description of Audouinella bispora by B0r- gesen is broadened here based on the variation seen within North Carolina plants, but no char- acteristics were found that contradict the earlier descriptions. Audouinella botryocarpa (Harvey) Woelkerling FIGURES 1^-/ Audouinella botryocarpa (Harvey) Woelkerling, 1971:37.— Searles and Schneider, 1978:100. Callithamnion botryocarpum Harvey, 1854:563. DESCRIPTION.—Plants epiphytic, epiendophytic SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES or saxicolous to 1 cm tall, caespitose, arising from an irregularly branched prostrate system with entangled filaments free from one another or coalesced into a pseudoparenchymatous disc, oc- casionally supplemented by corticating rhizoids descending cells or lower portions of erect fila- ments; base giving rise to several erect filaments, much and irregularly to unilaterally branched, sometimes terminating abruptly in multicellular hair-like extensions or gradually tapering toward the tips; cells of main axes cylindrical (10—) 15— 20(-30) jum diam., 30-120 jum long and 6-15 jum diam. in the ultimate segments, each cell contain- ing a single parietal lobate plastid with one to several pyrenoids, unicellular hairs unknown. Monosporangia terminal on short branches and adaxial, near the bases of branches, secund or scattered, sessile or on one-celled pedicels, sin- gle, paired, or rarely in groups of 3 to 5, ovoid, 12-28 jum diam., 18-43 jum long; tetrasporangia cruciate, with range of dimensions, shape, and position similar to monosporangia, in general somewhat larger than monosporangia on the same plant. Sexual plants unknown in the Caro- linas. TYPE-LOCALITY.—King George's Sound, west- ern Australia. HOLOTYPE.—Harvey 324, travelling collection; not in TCD. DISTRIBUTION.—North Carolina, Australia, New Zealand. NORTH CAROLINA SPECIMENS STUDIED.—Beau- fort, Radio Is. jetty, 2 Mar 1974, Schneider 829 (DUKE 13250, 13251, 13264; CWS). Beaufort, Radio Is. jetty, 16 Feb 1975, Searles 75-1-1 (DUKE 14178; CWS). Beaufort, Radio Is. jetty, 14 Sep 1981, Schneider 2720 (DUKE 16524; CWS). Beaufort, Pivers Is. seawall, 10 Nov 1981, Schneider 2746 (DUKE 16590; US 88908; CWS). Beaufort; Pivers Is. seawall, 7-8 Dec 1981, Schneider 2753 (DUKE; US 88909; CWS). Beaufort, Radio Is. jetty, 20 Jul 1982, Schneider 2864 (DUKE; US 88910; CWS). REMARKS.—Known only as an obvious epi- phyte of Codium from the Pivers Island seawall and Radio Island jetty, Beaufort, but probably more widespread. Collected July-March. This taxon is at present known in the Atlantic Ocean only from North Carolina and also repre- sents the only member of the acrochaetoid com- plex with more than one pyrenoid per cell (Figure 1^) from the east coast of North America (Searles and Schneider, 1978). Specimens have been com- pared with type material by W.J. Woelkerling. The only plants thus far collected in North Carolina are tetrasporic and/or monosporan- giate. Monosporangia, found in all collections, are large, falling in the upper size ranges of those listed by Woelkerling (1970, 1971). Many of the monosporangia are terminal on short branches, some later becoming subtended by the short lat- eral filaments that produce them, becoming dis- placed to a lateral position. Later the monospor- angia are seen as clustered near the base of branches, presumably due to the subtended fila- ments forming beneath monosporangia. Tetra- sporangia have been found only in December collections. Plants from New Zealand and Aus- tralia are no longer than 6 mm, while those in the Carolina region reach 1 cm, by far the largest of the acrochaetoids from this area. Multicellular hair-like extensions have thus far been observed only from the September Radio Island popula- tions; none of these minor differences warrants a separation from the taxon previously known only from the Southern Hemisphere, Audouinella botryocarpa. Audouinella corymbifera (Thuret) Dixon Audouinella corymbifera (Thuret in Le Jolis) Dixon in Parke and Dixon, 1976:590. Chantransia corymbifera Thuret in Le Jolis, 1863:107 [in part]. Acrochaetium corymbiferum (Thuret in Le Jolis) Batters, 1902:59.—Hoyt, 1920:473. Acrochaetium bornetii Papenfuss, 1945:313.—Searles and Schneider, 1978:100. DESCRIPTION.—Plants epiendophytic, 2-3 mm tall, arising from a persistent, large, globose spore, 12-15 jum diam., which produces an irregular, contorted branched filament that deeply pene- trates host tissue with cells 8-13 jum diam., 25-40 jum long; basal spore and occasional cells of the internal filament giving rise to erect branched axes, secund to alternate with branches more NUMBER 22 abundant above and not often rebranched; cells of erect filaments cylindrical, 7-16 jum diam., 22- 55 jum long, each cell containing a well-developed parietal lobed plastid with a single pyrenoid, unicellular hairs unknown. Monosporangia terminal on short branches or lateral, near the bases of branches, sessile or on one-celled pedicels, single, ovoid, 8-10 jum diam., 15-18 jum long, occasionally associated with ga- metangia. Dioecious, spermatangia lateral on main axes, formed in a corymbose cluster on a 1- 2 celled pedicel, colorless, 4-5 jum diam.; carpo- gonia lateral on a main axis, on a 1-2 celled pedicel near the bases of branches, producing a corymbose carposporophyte with terminal car- posporangia, ovoid, 9-15 jum diam., 14-18 jum long. TYPE-LOCALITY.—Belle-Ile-en-Mer, Atlantic France. HOLOTYPE.—Cannot be located (Dixon and Irvine, 1977:88). DISTRIBUTION.—North Carolina, Bermuda, Mediterranean, England, France. NORTH CAROLINA SPECIMENS STUDIED.—Beau- fort harbor, 20 May 1907, Hoyt (DUKE 135; US 33648). REMARKS.—Abundant sexually reproductive plants found on one specimen of Dasya from Beaufort harbor, to date collected only once, in May. For a clarification of the correct epithet refer to Dixon and Irvine (1977). Audouinella dasyae (Collins) Woelkerling FIGURE \m-p Audouinella dasyae (Collins) Woelkerling, 1973b:545, figs. 10- 31.—Searles and Schneider, 1978:100.—Kapraun, 1980:40 [in part]. Acrochaetium dasyae Collins, 1906:191.—Aziz, 1967:408. Acrochaetium robustum sensu Williams, 1948:690; 1951:153.— Taylor, 1960:315 [in part]. [?Not B0rgesen, 1915:40, figs. 38-40.] DESCRIPTION.—Plants epiphytic, to 3 mm tall, caespitose, arising from a basal spore and wide- spreading uniseriate, prostrate filamentous axis it produces; original spore elongated pyriform to panduriform, not greater in size than the cells which it produces, rarely remaining simple, often becoming obscured by descending rhizoids and entangled prostrate filamentous axes; creeping prostrate system giving rise to several erect fila- mentous, moderate to much and irregularly branched, often tapering to the tips, cells cylin- drical, 6-12(-16) jum diam., 25-70(-90) jum long in lower portions, 6-10 jum diam., 15-60 jum long in the ultimate segments, each having a single parietal plastid with one pyrenoid, hairs un- known. Monosporangia mostly single, occasionally in pairs, secund to scattered, sessile or on one-celled pedicels, ovoid, 7—12(—16) jum diam., 16-27 jum long, borne on vegetative or gamete-producing plants. Monoecious or dioecious, spermatangia terminal or lateral on specialized short lateral branched axes, globose to ovoid, 2-4 jum diam., 3-5 jum long; carpogonia scattered, sessile or stalked; carposporophytes with numerous termi- nal and lateral ovoid carposporangia, 9-12 jum diam., 16-24 jum long. TYPE-LOCALITY.— Woods Hole, Massachu- setts, USA. HOLOTYPE.—FH. DISTRIBUTION.—Florida to Prince Edward Is- land (discontinuous; not reported from South Carolina and Georgia). NORTH CAROLINA SPECIMENS STUDIED.—Cape Lookout jetty, 16 Jun 1947, Williams (CWS). Cape Lookout jetty, 14 Aug 1962, Aziz (DUKE 125). Beaufort, Pivers Is., 6 Apr 1963, Aziz (DUKE 158). Onslow Bay, reef, 16 Jul 1968, Searles (CWS). Beaufort, Radio Is. jetty, 8 May 1971, Schneider 172a (CWS). Onslow Bay, reef, 11 Aug 1981, Schneider 2687 (DUKE 16523; CWS). REMARKS.—Epiphytic on Dasya, Sargassum, and other hosts from shallow and deep subtidal envi- ronments, April-August in the area. After an examination of Williams's collections of Acrochaetium robustum, I find those specimens easily fall within the circumscription of Audouinella dasyae (Woelkerling, 1973b); any slight variation in their holdfasts is probably due to the differ- ences in surface textures of the host plants. An examination of Collins type material of A. robus- 8 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES turn is necessary to test whether the two taxa are indeed conspecific. Stegenga and Borsje (1976) have found that A. dasyae alternates with a het- eromorphic tetrasporophyte provisionally identi- fied as A. saviana; however, they felt it premature to deal with the nomenclatural problems arising from their study. North Carolina plants are often found with an early formed basal system of the ovoid to pyriform spore and one smaller accessory cell from which an upright filament rises obliquely (Figure \m,n). Audouinella daviesii (Dillwyn) Woelkerling FIGURE 2a-c Audouinella daviesii (Dillwyn) Woelkerling, 1971:28, figs. 7, 22. Conferva daviesii Dillwyn, 1809:73, suppl., pi. F. Audouinella hallandica sensu Kapraun, 1980:41, fig. 26 [in part]. [Not Audouinella hallandica (Kylin) Woelkerling, 1973a:82.] DESCRIPTION.—Plants epiphytic, epiendophy- tic, epizoic, or epiendozoic, to 6 mm tall, caespi- tose, arising from spreading to penetrating branched filamentous systems, centrally coalesced into irregular pseudoparenchymatous discs or en- tangled fungiform masses, original spores not per- sistent; erect filaments sparsely to freely and ir- regularly to unilaterally branched, branches often ending in long multicellular hair-like projections, 3-4 jum diam., 60-120 jum long; cells of main axes cylindrical (6-)9-12(-20) jum diam., (8-) 15-50 (-70) jum long, and only slightly tapering to tips, which are 7.5-10.0 jum diam. where extensions not formed, each cell containing a single parietal lobate plastid with one pyrenoid; unicellular hairs unknown. Monosporangia terminally clustered in groups of 3 or more on branched stalks or paired on 1-2 celled stalks adaxially on the lowermost cells of lateral branches, rarely single and scattered, ovoid, 7-13 jum diam., 8-20 jum long; tetraspo- rangia cruciate, paired on unicellular pedicels, solitary or in groups of 3, in similar positions to monosporangia, 13-22 jum diam., 12-36 jum long. Sexual plants unknown in the Carolinas. TYPE-LOCALITY.—Bantry Bay, Ireland. HOLOTYPE.—NMW. DISTRIBUTION.— Nearly cosmopolitan. As yet, not known from South Carolina and Georgia. NORTH CAROLINA SPECIMENS STUDIED.— Wrightsville Beach, Masonboro Inlet jetty, 18 Jan 1973, Kapraun (WNC). Wrightsville Beach, Ma- sonboro Inlet jetty, Apr 1977, 6 Apr 1978, Kapraun (WNC; DUKE). Ft. Macon jetty, 30 Jun 1982, Schneider 2831 (DUKE; US 88911; CWS). REMARKS.—To date, known only from the shal- low subtidal of Masonboro Inlet jetty, Wrights- ville Beach, January and April, and the intertidal of Fort Macon jetty, June. This is the first report of this taxon from North Carolina, although commonly known to the north (Woelkerling, 1973b). Only sporangiate plants have been found to date and these agree in all respects to recently published descriptions (Woelkerling, 1971, 1973b; Dixon and Irvine, 1977). Multicellular hair-like projections were found only in the January collection and tetra- sporangia only in April. Audouinella densa (Drew) Garbary FIGURE 2d-i Audouinella densa (Drew) Garbary, 1979:490. Rhodochorton densum Drew, 1928:168, pi. 38: figs. 17-24. Acrochaetium densum (Drew) Papenfuss, 1945:308.—Stegenga and Vroman, 1976:276, figs. 1-9. Acrochaetium thuretii sensu Kapraun, 1980:41, fig. 27. [Not Acrochaetium thuretii (Bornet) Collins and Hervey, 1917: 98.] Acrochaetium hummii Aziz, 1965:59, pi. 8: figs. 1-6, pi. 9: figs. 1-4, pi. 10: fig. 4 [nomen nudum]. DESCRIPTION.—Plants epiphytic or epizoic, 0.2- 0.3(-0.5) mm tall, caespitose, arising from an irregularly branched prostrate system, often orig- inally developed from a persistent septate spore, occasionally coalesced to form a pseudoparenchy- matous disc, especially centrally, giving rise to several erect filaments, sparsely to much branched, irregular, alternate, opposite and se- cund, usually with a moderate branching above, the branches sometimes tapering toward the tips, often ending in colorless unicellular hairs; cells of the upright filaments cylindrical to clavate, 4-8 NUMBER 22 (-13) jum diam., 7-18 jum long, each containing a single lobate to stellate plastid with a single pyrenoid. Monosporangia single or 2-3 seriate, terminal, alternate secund, rarely opposite, sessile or on one-celled pedicels, single or in pairs, rarely clus- tered; monosporangia ovoid to ovoid-ellipsoid, 7- 10 jum diam., 10-13 jum long. TYPE-LOCALITY.—Fort Point, San Francisco, California, USA. HOLOTYPE.—UC 294560 (Gardner 4607). DISTRIBUTION.—Atlantic: North Carolina, South Carolina, Brazil, Netherlands. Pacific: Mexico, California, Japan, Kurile Islands. NORTH CAROLINA SPECIMENS STUDIES.—Ft. Ma- con jetty, 27 Jun 1960, Aziz (DUKE 112). Beau- fort, Radio Is. jetty, 15 Apr 1973, Kapraun (NCW). Wrightsville Beach, Masonboro Inlet jetty, 26 Feb 1978, 20 May 1978, 10 Sep 1978, Kapraun (NCW). Ft. Macon jetty, 1 Jul 1978, Schneider 78- 12-2 (DUKE; US 88913; CWS). Wrightsville Beach, Masonboro Inlet jetty, 4 Apr 81, Searles (DUKE, slide). Beaufort, Pivers Is., 7-8 Dec 1981, Schneider 2760 (CWS). Beaufort, Radio Is. jetty, 21 Jun 1982, Schneider 2790 (DUKE; US 88915; CWS). Ft. Macon jetty, 30 Jun 1982, Schneider 2826 (DUKE; US 88912; CWS). REMARKS.—Epiphytic on various algae and an- imals, in particular Chaetomorpha aerea (Dillwyn) Kutzing, throughout the year from the intertidal and shallow subtidal of Cape Lookout jetty, Fort Macon jetty and the Pivers Island seawall, Beau- fort and Masonboro Inlet jetty at Wrightsville Beach. Also a single collection from 4.5 m in Port Royal Sound, South Carolina. Germinating spores divide into two equal cells after attaching to a host (Figure 2d,e). One or both of these initials gives rise to an erect and/or prostrate filament, occasionally one initial re- maining undivided for some time. The prostrate system consists of either irregular filamentous branched axes or becomes compacted into a pseu- doparenchymatous disc (see Figure 2/). This taxon was first reported from North Car- olina in an unpublished thesis as "Acrochaetium hummii" (Aziz, 1965). Stegenga and Vroman (1976) recognized the Aziz plants as members of a widespread heteromorphic acrochaetoid with seriate monosporangia tentatively identified by them as Acrochaetium densum (Drew) Papenfuss (= Audouinella densa [Drew] Garbary). In their report, Stegenga and Vroman (1976) call the report of gametangia in A. hummii by Aziz (1965) "strange." I could not find any sexual structures on Aziz's specimens (slides); therefore, to date only monosporangiate plants are known from here. This is further evidence that the North Carolina specimens belong in the A. densa com- plex outlined by Stegenga and Vroman (1976). They suggest A. microscopica as a possible game- tophytic generation if seriate monosporangia are not considered taxonomically important at the species level. Additional study is needed to clarify this relationship. For the present, I consider ser- iate monosporangia taxonomically significant, and leave the plants under the oldest validly published name, A. densa. Audouinella hallandica (Kylin) Woelkerling Audouinella hallandica (Kylin) Woelkerling, 1973a:82, figs. 1- 4.—Schneider, 1976:138.—Searles and Schneider, 1978: 100.—Kapraun, 1980:41, figs. 18-20 [in part]. Chantransia hallandica Kylin, 1906:123, fig. 8. Acrochaetium dufourii (Collins) B0rgesen, 1915:19.—Hoyt, 1920:470, fig. 26.—Taylor, 1960:305. Acrohaetium sargassi Bdrgesen, 1915:17, figs. 7-10.—Taylor, 1960:306. DESCRIPTION.—Plants epiphytic, to 1 mm tall, arising from an ovoid, obovate subglobose, disci- form, elongated to rectangular basal spore; if disciform, the flared base of the spore to 20 jum, otherwise 5-8 jum diam. and the same size or slightly larger than cells of the filaments it pro- duces; spore remaining simple producing 1-2(-3) erect filaments, branching irregularly lateral to secund, not closely set, beginning in the lower- most portions of the plant, occasionally tapering toward the apices, hairs unknown; lower cells cylindrical, 3-7 jum diam., 10-30 jum long, each containing a single parietal lobate plastid with one pyrenoid. Monosporangia single or paired, lateral to se- 10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES cund, more rarely terminal, sessile or on one- celled pedicels, obovate to ovoid, 5-10 /xm diam., 8-11 (-16) jum long. Dioecious, spermatangia op- posite on short lateral branches, rare; carpogonia sessile, rare. TYPE-LOCALITY.—Hogardsgrund, Halland, Sweden. HOLOTYPE.—LD (See Woelkerling, 1973a:84). DISTRIBUTION.—North Carolina, South Caro- lina, Bermuda, Sargasso Sea, Florida to Venezu- ela, Brazil, Europe. NORTH CAROLINA SPECIMENS STUDIED (repre- sentative).—Ft. Macon jetty, 13 Jul 1908, Hoyt, Acrochaetium dufourii (Collins) Hoyt (Isotype, DUKE 136). Beaufort, Pivers Is., 25 Jun 1960, Aziz (DUKE 113, 142). Beaufort, Shackleford jetty, 29 Jul 1961, Aziz (DUKE 143). Ft. Macon jetty, 5 Aug 1961, Aziz (DUKE 154-157). Beau- fort, Shackleford jetty, 6 Aug 1961, Aziz (DUKE 144, 149-153). Beaufort, Shackleford jetty, 12 Aug 1962, Aziz (DUKE 145-147). Cape Lookout jetty, 8 Jun 1975, Schneider 947 (DUKE 15077, 15133; CWS). Onslow Bay, wreck, 9 Jun 1975, Schneider 998a (CWS). Wrightsville Beach, Ma- sonboro Inlet jetty, 11 Nov 1977, Kapraun (WNC). Onslow Bay, reef, 11 Aug 1981, Schneider 2683 (CWS). REMARKS.—Known as an epiphyte of Dictyota and Sargassum from shallow subtidal habitats near Beaufort and Wilmington, and on a variety of red and brown algae from deep water (to 35 m) environs, June-November. This taxon is most easily recognized when at- tached to its host by the distinctive broad disc of cell-wall materials. The disc does not stain and the original spore remains as a simple elongated basal cell the same size or slightly larger than the cells of erect filaments it produces. A recent study by Stegenga and Borsje (1977) demonstrates that a plant provisionally deter- mined as Audouinella hallandica is the gametophyte of a heteromorphic plant whose sporophyte is tentatively determined as Acrochaetium polyblastum (Rosenvinge) B0rgesen. No formal taxonomic changes for this species complex were offered, however. Audouinella hoytii (Collins), new combinaton FIGURE 2j Acrohaetium hoytii Collins, 1908:134.—Hoyt, 1920:470, figs. 27, 28.—Williams, 1948:690; 1951:153.—Searles and Schneider, 1978:100.—Wiseman, 1978:29. DESCRIPTION.—Plants epiphytic, 0.2-1.3 mm tall, arising from a persistent, large, globose spore, 9-15 (-28) jum diam. including an obvious wall 2- 3 jum thick; spore superficial to slightly embedded in host tissue, elongating to 30 jum, being ob- viously broader than the cells it produces, re- maining simple, rarely forming one or a few accessory cells, producing 1—2(—4) erect filaments; erect branching infrequent to frequent, often se- cund, ultimate branches elongated, tapering to the apices for the most part, 2-3 jum diam., hairs unknown; cells of the main axes cylindrical, 5-7 jum diam., 10-20 jum long, each with a single parietal plastid and a large central pyrenoid. Monosporangia secund or lateral on upper por- tions of the plant, sessile or on one-celled pedicels, oblong, 5.0-7.5 jum diam., 11-15 jum long. Mon- oecious, spermatangia lateral and terminal on short lateral branching systems; carpogonia ses- sile, lateral; carposporophytes developing as short branching systems lateral on main axes or near the base of branches, with 4-16 terminal carpos- pores, 4-9 jum diam., 7.0-13.5 jum long. TYPE-LOCALITY.—Fort Macon jetty, vicinity Beaufort, North Carolina, USA. HOLOTYPE.—US 33650. DISTRIBUTION.—North Carolina, South Caro- lina. NORTH CAROLINA SPECIMENS STUDIED.—Ft. Macon jetty, 20 Sep 1905, Hoyt (US 43499). Ft. Macon jetty, 12 Oct 1905, Hoyt (Holotype, US 33650). Ft. Macon jetty, 15 Aug 1906, Hoyt (DUKE 137; US 43500a, 33651). Beaufort har- bor, 5 Oct 1906, Hoyt (DUKE 1460; US 43500b). Ft. Macon jetty, 13 Jul 1908, Hoyt (DUKE 1461; US 33652, 2803 as P.B.-A. 1540). Beaufort, Shac- kleford jetty, 14 Jul 1938, Blomquist (DUKE 133). Cape Lookout jetty, 28 Aug 1946, Williams (CWS). Ft. Fisher outcropping, July 1955, Rho NUMBER 22 11 (DUKE 139). Beaufort, Shackleford jetty, 4 Jul 1960, Aziz (DUKE 119, 122, 123). Beaufort, Ra- dio Is. jetty, 5 Jul 1960, Aziz (DUKE 148). Beau- fort, Pivers Is., 18 Jul 1974, Searles (DUKE 16091). Beaufort, Radio Is. jetty, 13 Jun 1978, Schneider 78-4-5b (CWS). Beaufort, Duncan breakwater, 27 Jun 1978, Schneider 78-10-20 (CWS). Beaufort, Radio Is. jetty, 20 Sep 1981, Schneider 2738 (DUKE 16522; US 88914; CWS). Beaufort, Ra- dio Is. jetty, 21 Jun 1982, Schneider 2789 (DUKE; US 88915; CWS). Beaufort, Radio Is. jetty, 20 Jul 1982, Schneider 2861 (DUKE; US 88916; CWS). REMARKS.— Known as an epiphyte of Dictyota, Padina, and other seaweeds from shallow subtidal habitats in the Beaufort area, June-October. Although similar in morphology to Audouinella alariae (Jonsson) Woelkerling (1973b) from New England and Europe, main axes, carposporangia, and monosporangia of A. hoytii are one-half the size of the first mentioned taxon. A further com- parison is necessary to clarify the relationship of these two species. Audouinella hoytii has also been linked to A. pectinata (Kylin) Papenfuss by Woelk- erling (1971). Audouinella hypneae (B0rgesen), new combination FIGURE 2k-l Chantransia hypneae B0rgesen, 1909:2, fig. 2. Acrochaetium hypneae (B0rgesen) B0rgesen, 1915:51, figs. 53, 54. Acrochaetium seriatum B0rgesen, 1915:32, figs. 25-28.—Wil- liams, 1948:690—Searles and Schneider, 1978:100. DESCRIPTION.—Plants epiphytic or epiendo- phytic to 1 mm tall arising from small compact to spreading filamentous prostrate system com- posed of one or more irregularly bent, branched filaments that can be coalesced centrally into a pseudoparenchymatous disc; each cell of the disc or ramified filaments capable of producing an erect filament; erect branches little to radially and much branched below, alternate, often pec- tinate secund above, cells cylindrical, 6-10 jum diam., 16-22 jum long below, tapering to 4-7 jum diam., 16-30 jum long in the ultimate segments, each containing a single parietal plastid with an obvious pyrenoid, occasionally bearing unicellu- lar hyaline hairs to 60 jum long. Monosporangia single, rarely paired, lateral in long secund series or rarely terminal, occasionally scattered, sessile or on one-celled pedicels, oblong- ovoid, 6-10 jum diam., 9-13 jum long. Gametangia unknown. TYPE-LOCALITY.—St. Thomas, Virgin Islands. HOLOTYPE.—C. DISTRIBUTION.—North Carolina, South Caro- lina, Georgia, Bermuda, Florida, Virgin Islands, Brazil, East Indies. NORTH CAROLINA SPECIMENS STUDIES.—Ft. Ma- con jetty, 15 Jul 1907, Hoyt (US 42553). Cape Lookout jetty, 28 Sep 1946, Williams (CWS). Beaufort, Shackleford jetty, 4 Jul 1960, Aziz (DUKE 124). Cape Lookout jetty, 8 Aug 1961, Aziz (DUKE 186-187). Onslow Bay, reef, 11 Jul 1976, Searles (CWS). Onslow Bay, reef, 15 Jul 1981, Handsel, BLM 65818144, 65818145, 65818148 (Duke Marine Lab Reference Mu- seum). Onslow Bay, reef, 28 Jul 1981, Amsler, BLM 65818152 (Duke Marine Lab Reference Museum). Onslow Bay, reef, 11 Aug 1981, Schnei- der 2712 (DUKE 16523; CWS). Cape Lookout jetty, 5 Jul 1982, Schneider 2841 (DUKE; US 88917; CWS). REMARKS.—On a variety of plant and animal hosts from Beaufort area jetties and on Arthrocla- dia, Spatoglossum, and other hosts from deep off- shore, July-September. North Carolina plants show variation in their attachment, ranging from the small compact, epiphytic disc illustrated for Acrochaetium seriatum (B0rgesen, 1915:32) to the short branched or simple prostrate epiphytic (A. gracile, B0rgesen, 1915:26) or endophytic (A. hypneae, B0rgesen, 1915:52) filament. In his discussion of A. hypneae, B0rgesen (1915:52) states, "If the basal part is left out of consideration it cannot be denied that the above, [newly] described Acrochaetium seriatum shows great likeness to this species. But while the present plant has an endophytic base the other is epiphytic." I have observed epiphytic, epi-endo- phytic and endophytic bases of this small spored 12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES acrochaetoid on the same host, Arthrocladia, from 30 m offshore. Basal system variation has previ- ously been used as a taxonomic criterion, even though this variation may have been due to the texture of the host. Recent studies have shown great variation within the attachment morphol- ogy for Audouinella daviesii (Dillwyn) Woelkerling, and other species (Woelkerling 1971, 1973b). I therefore propose that plants previously reported from North Carolina as Acrochaetium seriatum B0r- gesen by Williams (1948) are the same as those I have observed. Because the single criterion that separates them appears unreliable, Acrochaetium seriatum is synon- ymized under the earliest available taxon, Au- douinella hypneae. A recent report of somewhat larger spore dimensions for A. seriatum than shown in this and classical reports requires additional study. If the plants reported by Hall and Eiseman (1981) conform to B0rgesen's A. seriatum, then spore size distinction between A. hypneae and A. saviana is clouded, and further taxonomic reduc- tions may be necessary. Until additional studies are completed, the key and description of A. hypneae in this report conform to Carolina speci- mens and classical reports (e.g., B0rgesen, 1915; Taylor, 1960). It is, at this stage, still possible to segregate Audouinella bispora from A. hypneae on the basis of size and type of sporangia. I expect however, as did B0rgesen, that these two taxa are one and that further collections and life history studies will bear this out. The necessity of a taxonomic study of the many B0rgesen Caribbean acrochae- toid species is obvious to me. Several of the small spored, partially or wholly endophytic or epiphy- tic plants will no doubt prove to be conspecific. At present, I consider the evidence sufficient to synonymize two of the species in this complex. Audouinella infestans (Howe and Hoyt) Dixon FIGURE 2m Audouinella infestans (Howe and Hoyt) Dixon in Parke and Dixon, 1976:590. Acrochaetium infestans Howe and Hoyt, 1916:116, pi. 14.— Hoyt, 1920:473, pi. 68. Kylinia infestans (Howe and Hoyt) Papenfuss, 1947:438.— Taylor, 1960:301. Colaconema infestans (Howe and Hoyt) Woelkerling, 1973a:89, figs. 5, 6.—Searles and Schneider, 1978:100. DESCRIPTION.—Plants endophytic or endozoic with emergent axes to 100 jum tall exclusive of hairs, internal axes consisting of tortuous, serpen- tine, or labyrinthine irregularly branched fila- ments, sparse and straight for considerable dis- tances, often crowded between host cells or com- pacted and subparenchymatous, cells irregular to elongate, (2-)4-8(-13) jum diam., 6-60 jum long; emergent axes to 10 or more cells high but gen- erally less, simple or sparingly and irregularly branched, occasionally bearing terminal hairs to 170 jum long; cells cylindrical to ellipsoid, 3-7 jum diam., 6-30 jum long, containing a single parietal lobate plastid with a single pyrenoid. Monosporangia single or paired, terminal or lateral, often on the adaxial surface, sessile or on one celled pedicels, ovoid to ellipsoid, 4-6(-9) jum diam., 6-15 jum long. Gametangia unknown. TYPE-LOCALITY.—Onslow Bay, North Caro- lina, USA. HOLOTYPE.—US 56009. DISTRIBUTION.—North Carolina, Bermuda, Sargasso Sea, British Isles, Mediterranean, Japan. NORTH CAROLINA SPECIMENS STUDIES.—Ons- low Bay, reef, 11 Aug 1914, Radclijfe, (Holotype, US 56009, Isotypes, US 33514, 33653, 336534, DUKE 140). Cape Lookout jetty, 8 Aug 1961, Aziz (DUKE, slide). Onslow Bay, wreck, 8 Jul 1976, Searles (CWS). Beaufort, Radio Is. jetty, 13 Jun 1978, Schneider 78-4-5a (CWS). Onslow Bay, reef, 15 Jul 1981, Searles BLM 65818144 (Duke Marine Lab Reference Museum). Onslow Bay, reef, 28 Jul 1981, Amsler BLM 65818152 (Duke Marine Lab Reference Museum). REMARKS.—Collections on various hydroids, Dictyota, and Sargassum from deep offshore habi- tats and shallow subtidal habitats near Beaufort and Wilmington, June-August. Audouinella microscopica (Nageli) Woelkerling Audouinella microscopica (Nageli) Woelkerling, 1971:33, figs. 10, 23A.—Searles and Schneider, 1978:100.—Kapraun, 1980:41, figs. 21, 22. Callithamnion microscopicum Nageli in Kutzing, 1849:640. NUMBER 22 13 Acrochaetium trifilum (Buffham) Batters, 1902:58. Acrochaetium parvulum (Kylin) Hoyt, 1920:470, fig. 25. Acrochaetium compactum Jao, 1936:241, pi. 10: figs. 6-14.— Williams, 1948:690. Kylinia crassipes (B0rgesen) Kylin, 1944:13.—Taylor, 1960:300. DESCRIPTION.—Plants minute epiphytes, 20- 150(-220) jum tall exclusive of hairs, arising from a persistent subglobose, isodiametric, to cylindri- cal spore with or without a thick cell wall, 5-15 jum diam., close to the same size as the cells it produces; spore giving rise to 1-6 erect and/or lateral filaments, commonly arcuate, simple to much branched, secund, irregular or opposite, branches short, tapering; cells barrel-shaped, iso- diametric to cylindrical, 3-10 jum diam., 3-11 jum long, having a single lobate to stellate plastid with one pyrenoid, often bearing hyaline hairs to 50 jum long. Monosporangia terminal and secund to oppo- site, sessile or on one-celled pedicels, ovoid, 4- 10(-15) jum diam., 6-15(-22) jum long. TYPE-LOCALITY.—Torquay, England. HOLOTYPE.—L 940285 . .. 306. DISTRIBUTION.—Almost cosmopolitan in distri- bution. NORTH CAROLINA SPECIMENS STUDIED.—Beau- fort Inlet, sea buoy, 27 Jul 1909, Hoyt (DUKE 141; US 33655). Beaufort, Shackleford jetty, 29 Nov 1940, Blomquist 11472 (DUKE 676, as Calli- thamnion polyspermum). Ft. Macon jetty, 27 Feb 1971, Schneider 50A (CWS). Wrightsville Beach, Masonboro Inlet jetty, 9 Jun 1977, Kapraun (WNC). Beaufort, Radio Is. jetty, 20 Jul 1982, Schneider 2865 (CWS). REMARKS.—Known as an epiphyte on a variety of intertidal and shallow subtidal seaweeds from the Beaufort and Wilmington areas, throughout the year. Tetrasporangia have not been observed on North Carolina specimens and only rarely elsewhere (Woelkerling, 1972). Although some authors have chosen to retain the similar Audouinella microscopica and A. parvula (Kylin) Dixon as separate entities (e.g., Dixon and Irvine, 1977), the grounds for their separation seem unclear and unsubstantial in light of the morphological diversity shown for the A. micros- copica complex based upon studies of several type collections (Woelkerling, 1972). Stegenga and Mulder (1979), however, find some of the mem- bers of this complex synonymized by Woelkerling (1972) belong in the separate genera based upon their utilization of a multiple generic scheme for the family. There is, however, no doubt that all previously and recently collected Carolina speci- mens easily fit into the circumscription of A. microscopica proposed by Woelkerling (1971, 1972, 1973b), which for the present I continue to follow. Examination of type material oi A. parvula (Kylin, 1906 as Chantransia parvula) is still necessary if a formal transfer is to be made. Audouinella microscopica is reported as Acrochae- tium trifilum (Buffham) Batters for adjacent Vir- ginia (Humm, 1979). That listing follows Aziz (1965), which finds A. parvulum synonymous with several Caribbean taxa, of which A. trifilum is the oldest available binomial. The synonyms in- cluded A. compactum Jao and A. crassipes (B0rge- sen) B0rgesen, both of which also appear as syn- onyms of A. microscopica (Woelkerling, 1972). Aziz did not mention A. microscopica in his paper, and at that time it was not listed for the Southeast or Caribbean (Taylor, 1960), but it is clear that the two are the same, with A. microscopica having historical precedence over A. trifilum. I therefore reject usage of the latter name. Audouinella ophioglossa, new species FIGURE 3 Plantae epiendophyticae, 1.0-2.2 mm elatae, spora persistenti magna globosa exorientes, 7.5- 12.5 jum diametro; spora basilis filum singulare unramosum aut ramosum furcatum irregulare contortum producens qui hospitem cum cellulas 2-4 jum diametris profunde penetrans, vel basem unicellulam remanens; spora basalis 1-2 axes er- ectas ramosas producens, secundas ad alternas et oppositas, rami plures communes super; cellulae filorum erectorum cylindricae, 4-5 jum diametris, 45-55 jum longae segmentis infernis, cellula omnis plastum singularem lobatum ad spiralem parie- talem cum pyrenoide singulari inconspicua cap- iens; trichomata unicellularia terminalia vel la- teralia, ad 2 mm, saepe cum ramis gametangiis consociata. 14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Monosporangia terminalia in ramis brevibus aut lateralia, secunda ad alterna aut opposita, sessilia aut in pedicellis unicellularis, singularia vel binata, ovoidea ad ellipsoidea, 6.0-12.5 jum diametris, 12.5-20.0 jum longa, plerumque cum gametangiis consociata. Plantae monoeciae, rami spermatangiis quadriverticillati, oppositi vel mi- nus plerumque ramus unicus ad uterque nodum in seriebus longis in partibus superis plantae; spermatangia binata in pedicellis unicelluris, glo- bosa, 2.5 jum diametris; carpogonia lateralia, op- posita vel juxta monosporangium, aliquando in verticillo ramorum spermatangiorum facientia, non nisi unum carposporophytum ad uterque nodum producentia, saepe in seriebus longis in- terrupts; carposporangia terminalia globosa, 10- 15 jum diametris. Specimen holotypicum a SCUBA lectum, 20 VII 1973, 34°23.0'N, 76°36.0'W (North Caro- lina), in profunditatae 22 metrorum, in Dudresnaya crassa Howe, leg. R.B. Searles, in herbariis Univ- ersitatis Dukensis, No. 16585, 96; Isotypi DUKE, US 88918. DESCRIPTION.—Plants epiendophytic, 1.0- 2.2mm tall, arising from a persistent, large, glo- bose spore, 7.5-12.5 jum diam., which produces a single unbranched or forked branched irregularly contorted filament that deeply penetrates host tissue with cells 2-4 jum diam., or remains as a unicellular base; basal spore giving rise to 1-2 erect branched axes, secund to alternate and opposite, more common above; cells of the erect filaments cylindrical, 4-5 jum diam., 45-55 jum long in lower segments; each cell containing a single lobate to spiral parietal plastid with a single inconspicuous pyrenoid, unicellular hairs termi- nal or lateral, to 2 mm, often associated with gametangial branches. Monosporangia terminal on short branches or lateral, secund to alternate or opposite, sessile or on one-called pedicels, single or in pairs, ovoid to ellipsoid, 6.0-12.5 jum diam., 12.5-20.0 jum long, commonly associated with gametangia. Monoe- cious, spermatangial branches in a whorl of 4, opposite paired or less commonly one per node, in long series in the upper parts of the plant; spermatangia paired on one-celled pedicels, glo- bose, 2.5 jum diam.; carpogonia lateral, opposite or adjacent to a monosporangium, occasionally forming in a whorl of spermatangial branches, producing only one carposporophyte per node, often in a long interrupted series; carposporangia terminal, globose, 10-15 jum diam. TYPE-LOCALITY.—Onslow Bay, North Caro- lina, USA. HOLOTYPE.—DUKE 16585. DISTRIBUTION.—Endemic to North Carolina as far as currently known. NORTH CAROLINA SPECIMENS STUDIED.—Ons- low Bay, reef, 18 Sep 1972, Schneider 553e (CWS). Onslow Bay, reef, 19 Jul 1973, Schneider 665a (CWS). Onslow Bay, reef, 20 Jul 1973, Searles (Holotype, DUKE 16585; Isotypes, DUKE 16586, 16587, US 88918). Onslow Bay, reef, 20 Jul 1973, Schneider 716a (Topotypes, CWS). Ons- low Bay, reef, 19 Jun 1980, Searles (DUKE 16589). REMARKS.—Audouinella ophioglossa is known 22- 35 m offshore on Dudresnaya crassa, June-Septem- ber. The species is named for the general appear- ance of the basal spore and often forked endo- phytic filament, which resembles the head of a snake with extended tongue (see Figure 3b-d). Of the several species worldwide with a persistent, obvious basal spore which produces a deep pen- etrating endophytic filament, this newly de- scribed plant is most similar to Audouinella subtil- issima (Kutzing) Garbary, an epiendophyte of Dudresnaya and other species along the French coast. The new taxon, A. ophioglossa, differs from A. subtilissima in that its branches do not taper toward the tips, in having blunt apices, a smaller filament diameter (4-5 vs. 6-7 jum), and conspic- uous unicellular hairs (see Figure 3fh,j). Hamel (1928a) reports A. subtilissima as having some filaments end in pseudohairs that do not appear to be unicellular, a feature not illustrated by him or by Kutzing (1861). Hamel (1928a) describes and illustrates other characters of A. subtilissima that vary from the North Carolina plant: oval to oblong carposporangia, carpogonia often distal to spermatangia on the same branch, and laterally disposed spermatangial branches. In A. ophiog- NUMBER 22 15 lossa, the carpogonia are either proximal to sper- matangial branches (Figure 3/), paired with one (Figure 3i), or are found as one branch in a whorl of 4, the remainder being male axes (Figure 3h). The verticillate arrangement of male branches in the new taxon (Figure 3hJ) is unique for this species cluster of Audouinella. In A. ophioglossa, monosporangia, male branches or carpogonia are commonly oppositely arranged (Figure 3fl,m), or a carpogonium is disposed opposite to a mono- sporangium (Figure 3/). Opposite spermatangia are found occasionally in Acrochaetium kuckuck- ianum Hamel (1928b), which has a similar mode of attachment to A. ophioglossa. Its size and dioe- cious habit still separate it clearly from the new species. From Audouinella corymbifera, the only other member from North Carolina having a persistent spore with a penetrating endophytic filament, A. ophioglossa, is distinguished by a smaller basal spore, narrower erect filaments, its monoecious habit and the disposition of spermatangial branches. Audouinella saviana (Meneghini) Woelkerling Audouinella saviana (Meneghini) Woelkerling 1973b:560, figs. 56-60.—Schneider, 1976:138.—Searles and Schneider, 1978:100. Callithamnion savianum Meneghini, 1840:511. Acrochaetium thuretii (Bornet) Collins and Hervey, 1917:98. Audouinella thuretii (Bornet) Woelkerling, 1971:36, figs. 12, 24. [Not Audouinella thuretii sensu Kapraun, 1980:41.] DESCRIPTION.—Plants epiphytic to epiendo- phytic, to 4 mm tall, arising from a compact, mostly superficial prostrate system of short, sim- ple or branched filaments, free from each other to coalesced into an irregular pseudoparenchy- matous disc, the original spore not recognizable; prostrate filaments the same size or distinctly larger than erect ones; erect filaments with irreg- ular or alternate to secund branching, cells of main axes and branches cylindrical 7-14 jum diam., 20-60 jum long, sometimes 4-6 jum diam. in the ultimate branches, unicellular hairs absent or rare, each cell containing a single parietal lobate plastid. Monosporangia single or paired, adaxially se- cund to irregularly arranged, sessile or on 1 (or 2) celled elongate pedicels, ovoid to ellipsoid, 10-15 jum diam., 18-27 jum long. Tetrasporangia cru- ciate, with shape and positions similar to mono- sporangia, often borne simultaneously, 17-24 jum diam., 26-34 jum long. Monoecious, spermatangia sparse, on short branchlets near the base of branches; carpogonia sparse and solitary on lower segments near the base of branches, carpospores 9-13 jum diam., 18-21 jum long. TYPE-LOCALITY.—Genoa, Italy. HOLOTYPE.—FL. DISTRIBUTION.—Atlantic North America, Ber- muda, Caribbean, Atlantic Europe, Mediterra- nean, Juan Fernandez Islands, Australia, Califor- nia. NORTH CAROLINA SPECIMENS STUDIED (repre- sentative).—Beaufort, Duncan Breakwater, 19 Oct 1940, Blomquist 11447A (DUKE 134, 1459), Ft. Macon jetty, 27 Jun 1960, Aziz (DUKE 114, 116, 159). Marshallberg, Cowper Is., 29 Aug 1962, Wirtensen (DUKE 115). Beaufort, Bird Shoals, 10 Jun 1971, Brauner (DUKE 12459). Wrightsville Beach, Figure 8 Is., Jun 1972, Kapraun (NCW). Onslow Bay, reef, 9 Jun 1974, Schneider 904 (DUKE 13514; CWS). Beaufort, Radio Is. jetty, 13 Jun 1978, Schneider 78-4-17 (CWS). Onslow Bay, wreck, 18 Jun 1978, Schneider 78-6-9 (DUKE 16271; CWS). Beaufort, Radio Is. jetty, 14 Sep 1981, Schneider 2727 (CWS). Cape Lookout jetty, 5 Jul 1982, Schneider 2852 (CWS). REMARKS.—Epiphytic usually on Codium but also known on Dictyopteris and other coarse algae from shallow subtidal and deep offshore habitats to 30 m, May-October. Dixon and Irvine (1977) and Garbary (1979) question the conspecificity of Audouinella thuretii (Bornet) Woelkerling with A. saviana as proposed by Woelkerling (1973b). In a numerical taxo- nomic approach to the Acrochaetiaceae, Garbary (1979) finds dissimilarity in some published de- scriptions of A. thuretii and A. saviana and states "that [Woelkerling's] conclusions are so different from those reached by other workers is perhaps a reflection of the limitations of dealing with her- 16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES barium specimens of acrochaetioid algae." I reject the inference that taxa can be distinguished on the basis of written description alone, especially if these are not type descriptions, as is often the case in Garbary's (1979) report. Therefore, I con- tinue to follow the results of Woelkerling (1973b) who, using herbarium specimens, found type ma- terial of the two taxa conspecific, and accept A. saviana as the epithet for North Carolina plants. This taxon may be an alternate generation of A. dasyae (see prior discussion and Stegenga and Borsje, 1976). Although I have not found unicellular hairs on specimens from the Carolinas, in conformity with those from New England (Woelkerling, 1973b), Hall and Eiseman (1981) reported them on plants during the winter months in Florida. Audouinella secundata (Lyngbye) Dixon Audouinella secundata (Lyngbye) Dixon in Parke and Dixon, 1976:590.—Kapraun, 1980:41, figs. 23-25. Callithamnion daviesii var. secundatum Lyngbye, 1819:129, pi. 41: figs. B4-B6. Colaconema secundata (Lyngbye) Woelkerling, 1973a:94, figs. 7, 8.—Searles and Schneider, 1978:100. Acrochaetium virgatulum (Harvey) Bornet, 1904:xxii.—Wil- liams, 1948:690; 1951:153. Kylinia virgatula (Harvey) Papenfuss, 1947:437.—Taylor, 1962:214. DESCRIPTION.—Plants epiphytic or epizoic, to 3 mm tall, scattered to caespitose, arising from a uni- to bistratose, generally compact pseudopar- enchymatous disc, the original spore not persist- ent; disc giving rise to several erect, usually freely and irregularly branched with laterals often con- sisting of only 1-5 cells, with or without lateral and teminal unicellular hairs to 300 jum long; cells of main axes cylindrical 8-15 (-20) jum diam., 15-100 jum long, each with an axial or parietal stellate plastid and single central pyrenoid. Monosporangia lateral or terminal on shorter lateral branches, sessile or pedicellate, single, paired or rarely in groups of 3, ovoid, (6-) 10-20 jum diam., (10-)15-26(-32) jum long. TYPE-LOCALITY.—Kvivig, Faeroes Islands. HOLOTYPE.—C. DISTRIBUTION.—Atlantic coast of North Amer- ica (not reported from Georgia), Sargasso Sea, Europe, Mediterranean, Canary Islands, Aus- tralia. NORTH CAROLINA SPECIMENS STUDIED.—Beau- fort, Pivers Is., 10 Aug 1907, Hoyt (DUKE 188 D; US 7l721a,b). Beaufort, Shackleford jetty, 12 Aug 1907, Hoyt (US 71721c). Ft. Macon jetty, 15 Apr 1909, Hoyt (DUKE 188A; US 33656). Beau- fort, Shackleford jetty, 19 Jun 1960, Aziz (DUKE 126, 131, 132). Beaufort, Bird Shoals, 10 Jun 1971, Brauner (DUKE 12457, 12459). Beaufort, Bird Shoals, 20 Jun 1971, Brauner (DUKE 12480). REMARKS.—Epiphytic on a variety of algae intertidally, April-August. Although some authors maintain a specific dis- tinction between Audouinella secundata and A. vir- gatula (Harvey) Dixon (e.g., Dixon and Irvine, 1977; Kornmann and Sahling, 1978), I have ob- served morphological plasticity in North Carolina plants that corroborates the findings leading to the recent synonomy of the two taxa (Woelker- ling, 1973a, 1973b). Easiest to distinguish in young stages, the pros- trate system develops from an orbicular, paren- chymatous group of cells. This disc may later proliferate as prostrate filamentous axes, obscur- ing the central disc and becoming more similar to later stages of Audouinella dasyae and A. saviana. Literature Cited Abbott, I.A. 1962. Some Lza^ora-inhabiting Species of Acrochaetium. Bemice P. Bishop Museum, Occasional Papers, 23:77- 120. 1968. An Examination of the Type Specimens of Some Species of Acrochaetium (Rhodophyceae). Taxon, 17:518-520. Aziz, K.M.S. 1965. Acrochaetium and Kylinia in the Southwestern North Atlantic Ocean. 245 pages, 16 plates. Doctoral dissertation, Duke University, North Carolina. 1967. The Life Cycle of Acrochaetium dasyae Collins. British Phycological Bulletin, 3:408. Batters, E.A.L. 1902. A Catalogue of the British Marine Algae. Journal of Botany (London), 40(supplement): 1-107. B0rgesen, F. 1909. Some New or Little Known West Indian Flori- deae. Botanisk Tidsskrift, 30:1-19, 11 figures, 2 plates. 1910. Some New or Little Known West Indian Flori- deae, II. Botanisk Tidsskrift, 30:177-207, 20 figures. 1915. The Marine Algae of the Danish West Indies, Part III: Rhodophyceae (1). Dansk Botanisk Arkiv, 3(1 a): 1-80, 86 figures. Bornet, E. 1904. Deux Chantransia corymbifera Thuret: Acrochaetium et Chantransia. Bulletin de la Societe Botanique de France, 51 (supplement) :xiv-xxiii. Borsje, W.J. 1973. The Life History of Acrochaetium virgatulum (Harv.) J. Ag. in Culture. British Phycological Journal, 8:205. Bory de Saint Vincent, J.B.M. 1823. Auduinella. In Dictionnaire classique d' Histoire Natu- relle, volume 3, pages 340, 341. Paris: Rey et Gravier. Collins, F.S. 1906. Acrochaetium and Chantransia in North America. Rhodora, 8:189-196. 1908. Two New Species of Acrochaetium. Rhodora, 10:133- 135. Collins, F.S., and A.B. Hervey 1917. The Algae of Bermuda: Contributions of the Ber- muda Biological Station for Research, number 68. Proceedings of the American Academy of Arts and Sciences, 53:1-195. Dillwyn, L.W. [1809.] British Confervae. 87 pages, London. W. Phillips. Dixon, P.S., and L.M. Irvine 1977. Introduction, Nemaliales, Gigartinales. Part I of Volume I (Rhodophyta) in Seaweeds of the British Isles, xi + 252 pages. London: British Museum (Natural History). Drew, K.M. 1928. A Revision of the Genera Chantransia, Rhodochorton, and Acrochaetium, etc. University of California Publi- cations, Botany, 14:139-225, plates 37-48. Feldmann, J. 1962. The Rhodophyta Order Acrochaetiales and its Classification. Proceedings of the Ninth Pacific Science Congress {1957), 4:219-221. Garbary, D. 1978. On the Phylogenetic Relationships of the Acro- chaetiaceae (Rhodophyta). British Phycological Jour- nal, 13:247-254. 1979. Numerical Taxonomy and Generic Circumscrip- tion in the Acrochaetiaceae (Rhodophyta). Botan- ica Marina, 22:477-492. Hall, M.O, and N.J. Eiseman 1981. The Seagrass Epiphytes of the Indian River, Flor- ida I: Species List with Descriptions and Seasonal Occurrences. Botanica Marina, 24:139-146. Hamel, G. 1928a. Floridees de France, V. Revue Algologique, 3:99-158. 1928b. Sur les Genres Acrochaetium Naeg. et Rhodochorton Naeg. Revue Algologique, 3:159-210. Harvey, W.H. 1854. Some Account of the Marine Botany of the Colony of Western Australia. Transactions of the Royal Irish Academy, 22:525-566 Hoek, C. van den, and A.M. Cortel-Breeman. See van den Hoek, C, and A.M. Cortel-Breeman Holmgren, P.K., W. Keuken, and E.K. Schosield 1981. Index Herbariorum, I: The Herbaria of the World. Sev- enth edition, 452 pages. Utrecht, Netherlands: Bohn, Scheltema, & Holkema. Howe, M.A., and W.D. Hoyt 1916. Notes on Some Marine Algae from the Vicinity of Beaufort, North Carolina. New York Botanical Gar- den, Memoirs, 6:105-123, plates 11-15. Hoyt, W.D. 1920. Marine Algae of Beaufort, N.C. and Adjacent Regions. Bulletin of the Bureau of Fisheries (U.S.), 36:365-556, 47 figures, plates LXXXIV-CXIX. Humm, H.J. 1979. The Marine Algae of Virginia. Special Papers in 17 18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Marine Science, 3: viii + 263 pages. Charlottesville: University Press of Virginia. Jao, C.C. 1936. New Rhodophyceae from Woods Hole. Bulletin of the Torrey Botanical Club, 63:237-257, plates 10-13. Kapraun, D.F. 1980. An Illustrated Guide to the Benthic Marine Algae of Coastal North Carolina. Volume I: Rhodophyta. 206 pages, 227 figures. Chapel Hill. Kornmann, P., and P.-H. Sahling 1978. Meeresalgen von Helgoland, iii -I- 289 pages. Ham- burg: Biologische Anstalt Helgoland. Kutzing, F.T. 1849. Species algarum. vi + 922 pages. Leipzig. F.A. Brock- haus. 1861. Tabulae phycologicae .... Volume 11, ii + 32 pages. Nordhausen. Kylin, H. 1906. Zur Kenntnis einiger Schwedischen Chantransia- Arten. In Botaniska Studier til F.R. Kjellman, pages 113-126, figures 1-9. Uppsala. 1944. Die Rhodophyceen der Schwedischen Westkuste. Lunds Universitets Arsskrift, ny foljd, avdeling 2, 40(2): 1-104. Le Jolis, A.c.F. 1863. Liste des algues marines de Cherbourg, . ., Paris: J.B. Bailliere et Fils. [Also published, 1864, Memoires de la Societe Imperiales des Sciences Naturelles de Cherbourg, 10:1-168.] Lyngbye, H.C. 1819. Tentamen hydrophytologiae Danicae, . . . xxxii + 248 pages, 70 plates. Copenhagen: Typis Schultzianis. Meneghini, G. 1840. Botanische Notizen. Flora (Regensburg), 23:510- 512. Papenfuss, G.F. 1945. Review of the Acrochaetium-Rhodochorlon Complex of the Red Algae. University of California Publications, Botany, 18:299-334. 1947. Further Contributions toward an Understanding of the Acrochaetium-Rhodochorlon Complex of the Red Algae. University of California Publications, Bo- tany, 18:433-447. Parke, M., and P.S. Dixon 1976. Check-List of British Marine Algae—Third Revi- sion. Journal of the Marine Biological Association, United Kingdom, 56:527-594. Schneider, C.W. 1976. Spatial and Temporal Distributions of Benthic Marine Algae on the Continental Shelf of the Carolinas. Bulletin of Marine Science, 26:133-151. Searles, R.B., and C.W. Schneider 1978. A Checklist and Bibliography of North Carolina Seaweeds. Botanica Marina, 21:99-108 Stegenga, H. 1978. The Life Histories of Rhodochorton purpureum and Rhodochorton floridulum (Rhodophyta, Nemaliales) in Culture. British Phycological Journal, 13:279-289. Stegenga, H., and W.J. Borsje 1976. The Morphology and Life History of Acrochaetium dasyae Collins (Rhodophyta, Nemaliales). Acta Bo- tanica Neerlandica, 25:15-29. 1977. The Morphology and Life History of Acrochaetium polyblastum (Rosenv.) B0rg. and Acrochaetium hallan- dicum (Kylin) Hamel (Rhodophyta, Nemaliales). Acta Botanica Neerlandica, 26:451-470. Stegenga, H., and A.S. Mulder 1979. Remarks on the Audouinella microscopica Complex with a Brief Survey of Chromastrum (Rhodophyta, Nemaliales). Acta Botanica Neerlandica, 28:289-312. Stegenga, H., and N.D. Van Erp 1979. Morphological Variation in the Genus Acrochaetium (Rhodophyta, Nemaliales). Acta Botanica Neerlan- dica, 28:425-448. Stegenga, H., and M.J. van Wissen 1979. Remarks on the Life Histories of Three Acrochae- toid Algae (Rhodophyta, Nemaliales). Acta Botan- ica Neerlandica, 28:97-115. Stegenga, H., and M. Vroman 1976. The Morphology and Life History of Acrochaetium densum (Drew) Papenfuss (Rhodophyta, Nemali- ales). Acta Botanica Neerlandica, 25:257-280, figures 1-14. Taylor, W.R. 1960. Marine Algae of Eastern Tropical and Subtropical Coasts of the Americas, ix + [iii] + 870 pages. Ann Arbor: University of Michigan Press. 1962. Marine Algae of the Northeastern Coast of North America. Second edition with corrections, viii + 509 pages. Ann Arbor: University of Michigan Press, van den Hoek, C, and A.M. Cortel-Breeman 1970. Life-History Studies on Rhodophyceae: III, Scinaia complanata (Collins) Cotton. Acta Botanica Neerlan- dica, 19:457-467. West, J.A. 1968. Morphology and Reproduction of the Red Alga Acrochaetium pectinatum in Culture. Journal of Phycol- ogy, 4:89-99. 1969. The Life Histories of Rhodochorton purpureum and R. tenue in Culture. Journal of Phycology, 5:12-21. Williams, L.G. 1948. Seasonal Alternation of the Marine Floras at Cape Lookout, North Carolina. American Journal of Bo- tany, 35:682-695. 1951. Algae of the Black Rocks, pages 149-159. In A.S. Pearse and L.G. Williams, The Biota of the Reefs off the Carolinas. Journal of the Elisha Mitchell Scientific Society, 67:133-161. NUMBER 22 19 Wiseman, D.R. 1978. Benthic Marine Algae. In R.G Zingmark, An An- notated Checklist of the Biota of the Coastal Zone of South Carolina, pages 23-36. Columbia: University of South Carolina Press. Woelkerling, W.J. 1970. Acrochaetium botryocarpum (Harv.) J. Ag. (Rhodo- phyta) in Southern Australia. British Phycological Journal, 5:159-171. 1971. The Morphology and Taxonomy of the Audouinella Complex (Rhodophyta) in Southern Australia. Australian Journal of Botany, Supplement, 1:1-91, fig- ures 1-27. 1972. Studies on the Audouinella microscopica (Naeg.) Woelk. Complex (Rhodophyta). Rhodora, 74:85- 96. 1973a. The Audouinella Complex (Rhodophyta) in the Western Sargasso Sea. Rhodora, 75:78-101, figures 1-8. 1973b. The Morphology and Systematics of the Audouinella Complex (Acrochaetiaceae, Rhodophyta) in Northeastern United States. Rhodora, 75:529-621, figures 1-83. 20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 1.—Audouinella affinis: a, surface view of a basal spore with a single erect filament; b, transverse section of Dictyota cells displaced by the penetrating spore. A. bispora: c, epiphytic and endophytic (broken lines) attachment filaments; d, bisporangia in various stages of development; ef, monosporangia. A. botryocarpa: g, detail of two cells showing lobate parietal plastids and associated multiple pyrenoids (clear); h,i, tetrasporangia; j, branch bearing monosporangia (some spent) and a single tetrasporangium; k,l, monosporangia. A. dasyae: m,n, axis rising on oblique angle from two-celled basal system; o, basal cells immersed in cell wall of Ceramium; p, maturing basal system with original panduriform basal cell still obvious. NUMBER 22 21 22 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 2.—Audouinella daviesii: a-c, various arrangements of monosporangia. A. densa: d, e, early prostrate system showing septate spore; /, habit on Chaetomorpha, single and seriate monospor- angia; g-i, single and 2-3 seriate monosporangia. A. hoytii: j, habit of a monosporangiate plant, showing obvious persistent basal spore. A. hypneae: k, basal system with erect filaments; /, monosporangia in series. A. infestans: m, endophytic system and emergent axis with monospor- angia and a single unicellular hair. NUMBER 22 23 24 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 3.—Audouinella ophioglossa, new species: a, basal spore, without endophytic penetrating filament, adjacent to Dudresnaya cortical cells; b-e, basal spores with erect and forked penetrating filaments;/, habit, showing paired spermatangia (s) and a spent monosporangium (SM) opposite to a carpogonium (c); g, monosporangia (M) and a whorl of spermatangia; h,i, primarily male axis bearing carpogonia (c) with extended trichogynes, hair cell present; j, male axis with numerous unicellular hairs (H) among whorls of spermatangia; k, carpogonium with 4 sper- matangia on trichogyne; /, opposite carpogonia; m, opposite or single monosporangia, some spent (SM) ; n-p, development of carposporophyte with carpospores (cs). 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If several illustrations are treated as components of a single composite figure, they should be designated by lowercase italic letters on the illustration; also, in the legend and in text references the italic letters (underlined in copy) should be used: "Figure 9b." Illustrations that are in- tended to follow the printed text may be termed Plates, and any components should be similarly lettered and referenced: "Plate 9b." Keys to any symbols within an illustration should appear on the art rather than in the legend. Some points of style: Do not use periods after such abbre- viations as "mm, ft, USNM, NNE." Spell out numbers "one" through "nine" in expository text, but use digits in all other cases if possible. Use of the metric system of measurement is preferable; where use of the English system is unavoidable, supply metric equivalents in parentheses. Use the decimal sys- tem for precise measurements and relationships, common frac- tions for approximations. Use day/month/year sequence for dates: "9 April 1976." For months in tabular listings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc. Omit space between initials of a personal name: J.B. Jones." Arrange and paginate sequentially every sheet of manu- script in the following order: (1) title page, (2) abstract, (3) contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes section, (8) glossary, (9) bibliography, (10) legends, (11) tables. Index copy may be submitted at page proof stage, but plans for an index should be indicated when manuscript is submitted.