The Crustose Coralline Al^ae (Rhodophyta: Corallinaceae) of the Hawaiian Islands THE MARINE SCIENCES NUMBER 15 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 Anthropo/ogy 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 Zoo/ogy 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 15 The Crustose Coralline Algae (Rhodophyta: Corallinaceae) of the Hawaiian Islands Walter H. Adey, Roberta A. Townsend, and William T. Boykins SMITHSONIAN INSTITUTION PRESS City of Washington 1982 ABSTRACT Adey, Walter H., Roberta A. Townsend, and William T. Boykins. The Crustose Coralline Algae (Rhodophyta: Corallinaceae) of the Hawaiian Is- lands. Smithsonian Contributions to the Marine Sciences, number 15, 74 pages, 47 figures, 1982.—Crustose corallines were collected from a wide range of depths (intertidal to about 300 m) throughout the Hawaiian Archipelago. A total ol 25 species in 10 genera are recognized on the basis of habit, anatomy, morphology, and ecology, including one new genus and 10 new species. Generic and specific keys for the differentiation of the Hawaiian crustose corallines are also provided. The ecology of each species, in terms of depth distribution and habitat, is also given, and the potential use of these plants in determining paleoenviron- ments in the Hawaiian Neogene is discussed. The Caribbean and Hawaiian crustose coralline floras are briefly compared. The large number of "pair species" and the parallelism in subfamily, generic, and "pair species" ecology indicate that coralline evolution is very slow. The crustose corallines are potentially excellent paleoecological indicators for the Tertiary. OFFICIAL PUBLICATION DATE is handslamped 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 Adey, Walter H. The crustose coralline algae (Rhodophyta: Corallinaceae) of the Hawaiian Islands. (Smithsonian contributions to the marine sciences ; no. 15) Bibliography: p. Includes index. 1. Corallinaceae. 2. Marine algae—Hawaii. 3. Algae—Hawaii. I. Townsend, Roberta A. II. Boykins, William T. III. Title. IV. Series/ QK569.C8A33 589.4'1 81-21247 AACR2 Contents Page Introduction 1 Acknowledgments 2 Materials and Methods 2 Taxonomy 5 Presentation of Material 5 CORALLINACEAE 5 Key to the Subfamilies and Genera 5 MASTOPHOROIDEAE (Svedelius) Setchell, 1943 7 Porolithon (Foslie) Foslie, 1909 7 Key to the Species 7 Porolithon onkodes (Heydrich) Foslie 7 Porolithon gardineri (Foslie) Foslie 10 Paragomolithon, new genus 12 Paragomolithon conicum (Dawson), new combination 13 Neogoniolithon Setchell & Mason, 1943 15 Key to the Species 15 Neogoniolithon rugulosum, new species 17 Neogoniolithon rufum, new species 17 Neogoniolithon clavacymosum, new species 21 Neogoniolithon fosliei (Heydrich) Setchell & Mason 23 Hydrolithon (Foslie) Foslie, 1909 25 Key to the Species 25 Hydrolithon reinboldii (Weber-van Bosse & Foslie) Foslie 25 Hydrolithon breviclavium (Foslie) Foslie 26 Hydrolithon laeve, new species 31 Hydrolithon megacystum, new species 32 Lithoporella (Foslie) Foslie, 1909 34 Lithoporella melobesioides (Foslie) Foslie 34 LITHOPHYLLOIDEAE Setchell, 1943 35 Tenarea Bory, 1832 35 Tenarea tessellatum (Lemoine) Littler 35 Lithophyllum Philippi, 1837 37 Key to the Species 37 Lithophyllum kotschyanum Unger 37 Lithophyllum pallescens (Foslie) Foslie 40 Lithophyllum ganeopsis, new species 42 Lithophyllum insipidum, new species 44 Lithophyllum punctatum Foslie 47 in IV SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES MELOBESIOIDEAE (J.E. Areschoug) Yendo, 1902 47 Archaeolithothamnium Rothpletz, 1891 47 Key to the Species of Archaeolithothamnium 48 Archaeolithothamnium erythraeum (Rothpletz) Foslie 48 Archaeolithothamnium episoredion, new species 51 Lithothamnium Philippi, 1837 53 Key to the Species 53 Lithothamnium pulchrum Weber-van Bosse & Foslie 53 Lithothamnium australe Foslie 53 Mesophyllum Lemoine, 1928 58 Key to the Species of Mesophyllum 58 Mesophyllum madagascariensis (Foslie) Adey 58 Mesophyllum prolifer (Foslie) Adey 61 Mesophyllum purpurascens (Foslie) Adey 61 Mesophyllum syrphetodes, new species 63 Mesophyllum fluatum, new species 63 Discussion 66 Glossary 68 Literature Cited 69 Index 73 The Crustose Coralline Algae (Rhodophyta: Corallinaceae) of the Hawaiian Islands Walter H. Adey, Roberta A. Townsend, and William T. Boykins Introduction In the early part of the present century, the marine botanists Howe (1912) and Setchell (1926) stressed the critical importance of crustose corallines as well as other calcareous algae in reef formation. More recently, Littler (1971b, 1973a,b) measured the surface area coverage of organisms at several sites on Oahu. He deter- mined, for the Waikiki fringing reef, that the "crustose coralline algae cover 39% of the reef surface and exceed all other organisms as the major builders and consolidators of reef material" (Littler, 1973a: 103), while at 8-28 m, "the deep- water crustose Corallinaceae (38% mean cover) overshadow all other calcareous organisms in terms of standing stock and also seem to have more biological influence than do other limestone producers" (Littler, 1973b:381). Doty (1974) summarized the recent Hawaiian studies of the role of crustose corallines in reef construction. During the summer of 1965, two holes were cored through the upper Tertiary-Recent lime- stone cap of Midway Atoll and into the under- Walter H. Adey and William T. Boykins, Department of Paleobiol- ogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Roberta A. Townsend, Smithsonian Insti- tution Predoctoral Fellow, School of Biological Sciences, University of Sydney, N.S. W. 2006, Australia. lying basalt. Ladd, Tracey, and Gross (1967, 1970) and Gross, Milliman, Tracey, and Ladd (1969) concluded that the crustose coralline algae had been the most abundant carbonate building elements in the limestone cap, a situation com- parable to that reported for the Recent (Littler, 1971b). At about the same time, a series of borings of a Holocene "algal ridge" and its associated reef flat were begun at the other end of the Hawaiian Archipelago on Hanauma Reef in Oahu (Easton and Olson, 1973). This reef is at present strongly dominated by coralline algae, a condition that seems to have existed during much of the latter half of its 7000-year history. The present study was conceived to provide the necessary systematic and ecological information needed for the paleoenvironmental interpretation of the crustose coralline algae found in the Mid- way cores. Previous studies of Pacific corings and exposed limestones (Johnson, 1954, 1958, 1961, 1964) have been of limited value for stratigraphic or paleoecological interpretation because funda- mental systematic information has been lacking. In addition, diagnostic characteristics at the spe- cific and even generic levels have been difficult to use in the study of fossil corallines, since the major systematic papers to date are based on a small number of specimens with little information SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES about the ecological variation expected within a species. To rectify this situation, the emphasis of this study has been to delineate the major elements of the Recent crustose coralline algal flora and to describe each species in sufficient detail to allow ecologic treatment of crustose coralline algae by both marine biologists and geologists. As we shall point out, not only are the major coralline genera depth stratified, but to a large extent the depth patterns are evident even at the subfamily level. The following environments of deposition should be differentiable in cores of reefs and limestone caps: quiet lagoon, wave-swept back reef flat, shallow (< 30 m depth) fore reef or algal ridge, mid-depth (30-50 m depths) fore reef, and deep- water banks (50-150 m depths). This investigation is by no means to be re- garded as a comprehensive systematic-ecological study. Although the collection is large, there are a number of taxa represented by only a few specimens. This investigation treats 25 species, using an approach that is a simplified population analysis. The total Holocene crustose coralline flora of the area probably consists of 40-50 spe- cies. Concurrent with the study of the Hawaiian collections, the first author was engaged in an intensive field program in the Caribbean. Consid- erable emphasis will be placed on the comparison of the floras from these two regions, for example, the marked parallelism in the morphology and ecology of "pair species," and in the spatial ecol- ogy of the entire coralline flora at both generic and subfamily levels; however, for purposes of taxonomic identification, the Caribbean species were purposely not considered, because we feel that adequate population data are not available at this time to establish whether or not the many species pairs are identical or evolutionarily diver- gent. We have kept them in separate taxa. Indi- vidual cases are discussed in the species descrip- tions below. Tropical provinces of the Atlantic and Pacific have been separated since the Miocene, and ge- netic interchange for most species prevented. The abundance of "pair species" indicates that crus- tose coralline algae evolve slowly under tropical conditions. Although this suggests that they would be of little value for tropical stratigraphic correlation, at least for periods of 10-20 million years, it enhances their value for paleoecological investigations. The species composition and ecol- ogy presently existing in the Hawaiian chain are probably little different from that which existed throughout the time of deposition of the Midway limestone cap, though additional central Indo- Pacific elements are to be expected in the Mio- cene, the lower Pliocene, and occasionally in the Pleistocene. Hawaiian geologists have long considered the reefs in the northwestern part of the Hawaiian Archipelago to be limestone caps on older vol- canic islands and that the pattern of evolution of the chain was from the northwest to the southeast. The Midway boring established this pattern and placed it within a time scale of about 20 million years. In terms of plate tectonics, the Pacific plate has been moving WNW over a hot spot in the mantle at a rate of about 10 cm/year during the formation of the Hawaiian chain. A detailed understanding of the ecology of crustose coralline algae, a major component of Hawaiian reefs and banks, could potentially provide considerable in- sight into the changing patterns of environments occurring while these limestone caps have devel- oped. ACKNOWLEDGMENTS.—The Latin diagnoses were kindly provided by Dr. Hannah T. Croas- dale (Dartmouth College). Field studies for col- lections were made possible through the support of the National Science Foundation (research grant GA-27343) to the Hawaii Institute of Geo- physics of the University of Hawaii. We wish to thank Drs. Isabella Abbott, Michael Borowitzka, Susan Brawley, William Johansen, and James Norris for critically reading the manuscript. Materials and Methods The collections for this study were taken by David Child during April and October of 1971 NUMBER 15 at the locations shown in Figure 1. Transect collections were taken by SCUBA diving from the following zones: intertidal, 0-3, 3-9, 9-15, 15-21, 21-28, 28-37, and 37-46 meters. Dredge collections were taken primarily aboard the Uni- versity of Hawaii's R/V Tentu; those from Nero Bank and Kure taken by the R/V Agassiz in 1964 were received from H. Ladd. Collections by SCUBA were made within each depth zone by taking representative proportions of the different substrate types available to crus- tose corallines. Because light is also critical in determining coralline populations, an attempt was made to include cryptic areas in the collec- tions in proportion to their occurrence in the sampling area. The SCUBA collections were all made by a project field assistant, David Child, who was familiar with the collecting techniques regularly employed in coralline sampling (see, e.g., Adey, 1971). He was not familiar with the Hawaiian species and would not have uncon- sciously biased the collections; however, some plants, such as branching species and Porolithon onkodes (Heydrich) Foslie, are more conspicuous than others and are perhaps positively biased in our collections. Potential problems of this nature are discussed for each species. Specimens collected were returned in buckets to a variety of base laboratories and examined alive. Representative samples of each apparent species were live-fixed in Susa's fixative (Suneson, 1937), decalcified with trichloroacetic acid, and prepared for paraffin sectioning by standard tech- niques (Gray, 1958). Phosphotungstic hematoxy- lin was used as a stain in all cases (Adey and Johansen, 1972). Paraffin sectioning of fresh material is critical in the initial study of a poorly known crustose coralline flora; however, once the taxa are delin- eated in a regional flora, it is not always necessary to paraffin section live material for identification. In many species, surface characteristics, color, and conceptacle dimensions and shape are quite diagnostic for the subfamilies treated herein; however, properly dried and maintained herbar- ium specimens are essential. Also, microscopic examination of hand sections and fresh vertical fractures of dried crusts are invaluable for taxo- nomic decisions. External morphological data such as color, type, and size of branching were taken from dried specimens. The light microscope measurements included in the taxonomic study presented below are based on about 400 sectioned specimens, de- posited in the Non-Articulate Coralline Algal Herbarium, National Museum of Natural His- tory, Smithsonian Institution (USNC). Specimens were prepared for scanning electron microscopy by washing a small piece of crust in distilled water, drying, and mounting on stubs. Carbon coating by evaporation using an E.F.F.A. vacuum evaporator and sputter coating with gold palladium alloy (S.E.M. Laboratory, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560, USA) were done prior to specimen observation with a Cambridge S4-10 scanning electron microscope. Ecology and the surface aspects of the species treated are based on about 700 specimens, many of which had two to three identifiable species present. Plants of less than two to three square centimeters surface area and lacking reproductive structures or surface characteristics known to us from larger specimens were not treated. Almost 1000 units of species occurrence are employed in our ecological data. The unit of occurrence is defined as the pres- ence of an identifiable species on a specimen that is in almost all cases a block of "coralgal" sub- strate ranging from about 5-20 cm diam. The abundance figures cited in the following discus- sions refer simply to the number of samples within a given collection unit upon which a particular species occurs. Tabulation of the number of in- dividual plants found within a collection would be less meaningful because of the tendency of adjacent plants of the same species to grow to- gether and fuse without a trace of their former margins. Surface area occupied per species is a more desirable unit, but considering that the typical sample of coralline occupied substrate is a complex three-dimensional shape with several SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES NUMBER 15 TRH UC US USNC spatial niches represented on each specimen, areal measurement would be time consuming and probably unnecessary in demonstrating the basic patterns of species distribution. The technique used by Littler (1971b) based on projecting sur- face coverage found within randomly selected areas is very effective on a relatively smooth surface lacking cryptic niches but is of limited usefulness on typical "coralgal" bottoms. Ecological factors such as light, grazing, and epiphytes often alter the external appearance of coralline algae, and there can be considerable variation within a species. While we describe such morphological variation for certain taxa, other taxa are poorly known because of their limited number in the collections. A number of taxa for which we have inadequate data have been omit- ted from this paper. More intensive study in Hawaii and in more central parts of the Indo- Pacific will probably provide enough information on many of these less-abundant species to allow positive identification. Taxonomy One hundred seven species of crustose coralline algae previously described in many publications for the tropical-subtropical Indo-Pacific and trop- ical East Pacific were considered in this study. Fifty-four of these were described by Foslie from 1895 to 1929, isotypes of which are deposited in USNC. Nineteen species were described by Le- moine (1929), all but one from the tropical East Pacific; nine of these are represented by isotype fragments in our collection (USNC). Heydrich, from 1897 to 1901, described 12 of the species, though we have only two isotype fragments in our possession. Dawson, from 1944 to 1961, de- scribed 10 of the species considered, and although we have not been able to consult his type mate- rials, his descriptions are extremely good and generally lead us to feel confident in their use. The Lemoine (1929) and Heydrich (1897a,b,c; 1901a,b,c) descriptions are often minimal, and where type material was not available, it was felt that we lacked an adequate understanding of the taxon in question. In these cases we followed the descriptions of Foslie. All specimens, including the holotypes of the newly described species, are deposited in USNC. Presentation of Material In the taxonomic treatments, not all literature citations of previous species descriptions are given, but rather only those of particular signifi- cance or value in delineating the species in ques- tion. Only those taxa that are common have been treated fully. Neogoniolithon frutescens (Foslie) Setchell & Mason and Mesophyllum siamense (Fos- lie) Adey, although present, have not been in- cluded. The following abbreviations are used in this study. PC Museum d'Histoire Naturelle, Labora- toire de Cryptogamie, Paris herbarium of M. Foslie, Trondheim, Norway herbarium of the University of Califor- nia U.S. National Herbarium, Smithsonian Institution Non-Articulate Coralline Algal Herbar- ium, National Museum of Natural History, Smithsonian Institution CORALLINACEAE Key to the Subfamilies and Genera (Applies only to genera we encountered in Hawaii; see "Glossary" and Adey and Macintyre (1973) for discussion of terminology) 1. Tetrasporangia without caps, i.e., tetrasporangial conceptacles single pored 2 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Tetrasporangia with caps, i.e., tetrasporangial conceptacles multipored (Melobesioideae) ' 2. Perithallial secondary pit connections present, cell fusions more or less rare or lacking (Lithophylloideae) 3 Perithallial secondary pit connections absent, cell fusions present (Mastophoroideae) 4 3. Perithallium palisade, single-layered hypothallium Tenarea Perithallium nonpalisade, single-layered or multilayered coaxial hypothallium Lithophyllum 4. Heterocysts absent Lithoporella Heterocysts present 5 5. Hypothallium single layered Hydrolithon Hypothallium multilayered 6 6. Heterocysts vertical rows or single, hypothallium coaxial or simple-parallel to substrate Neogoniolithon Heterocysts loosely grouped into horizontal fields, hypothallium coaxial Paragomolithon, new genus Heterocysts grouped into tight horizontal pustulous fields, hypothallium simple-parallel or plumose Porolithon 7. "Lithothamnium-typo.'" cover cells not present, hypothallium coaxial Mesophyllum "Lithothamnium-type" cover cells present, hypothallium noncoaxial 8 8. Tetrasporangia in broad sori, partially calcified walls between adjacent sporangia Archaeolithothamnium Tetrasporangia in conceptacles, hypothallium simple-parallel Lithothamnium The first element in the key requires the pres- ence of tetrasporic conceptacles. Although these are very often lacking in boreal-arctic plants (at least during the summer), they are frequently present in tropical corallines—if not on the sur- face, often buried in the thallus and visible upon fracture. If the nature of a single-pored concep- tacle is uncertain, a quickly made slide (wet mount) will often indicate whether or not the contents are tetrasporangia. Even in the complete absence of diagnostic reproductive structures, very much can still be done on the generic level without paraffin sections. The following applies especially to the use of a high-powered dissecting microscope or reflecting compound microscope; hand sections, though time consuming, are equally useful. The consistent presence of heterocytes and their grouping, along with the form of the hypothal- lium, delineates the four genera of Mastophoro- ideae: Porolithon, Paragomolithon, Neogoniolithon, and Hydrolithon. Heterocysts do occur in Mesophyllum, although they tend to be small and scattered. A leafy plant with coaxial growth and few small heterocysts could be either a Mesophyllum or a Neogoniolithon. Reference to the species key and collection of more material for reproductive struc- tures is required in this case. In the Hawaiian saxicolous flora, the only Ten- area found is T. tessellatum (Lemoine) Littler, easily distinguished by its overlapping whorls (Figure 21). Similarly, no difficulty should be encoun- tered in distinguishing the large-celled and leafy Lithoporella melobesioides (Foslie) Foslie (Figure 20). Of the remaining species, the presence of sec- ondary pit connections in abundance will place a plant in Lithophyllum or in the melobesioids. Some care is needed here, as Archaeolithothamnium NUMBER 15 has abundant perithallial cell fusions, along with scattered secondary pit connections, or narrow or incipient fusions resembling secondary pit con- nections. Lithophyllum species also have a distinc- tive surface texture, which, although difficult to describe, is very helpful in initial separation of specimens: species with several epithallial cells are dull "chalky" in appearance; those with only a single epithallial cell are iridescent "chalky." Porolithon is the only genus with similar surface texture, but here the distinctive pustulous heter- ocyst fields are almost always present. Archaeolith- othamnium plants can appear similar to the irides- cent Lithophyllum plants, though the former are usually much darker red. A hand section is useful in this instance. Among the remaining species, Mesophyllum plants are almost invariably leafy in habit and even if subsequently branched will often revert to the leafy habit between branches. The plants are characterized by a coaxial hypothallium, though careful orientation of a section parallel to the growth direction may be required to see this. Archaeolithothamnium plants are characterized by a dark red-brown color and a quite glossy and smooth texture, whereas Lithothamnium and Meso- phyllum are mostly pink to bluish-pink and dull in texture, often with surface micro-ridges. The "Lithothamnium-type" cover cells are quite distinc- tive for both Lithothamnium and Archaeolithotham- nium but can be hard to determine with certainty without paraffin sections. MASTOPHOROIDEAE (Svedelius) Setchell, 1943 Porolithon (Foslie) Foslie, 1909 Key to the Species Plants crustose P. onkodes Plants branching P. gardineri Porolithon onkodes (Heydrich) Foslie FIGURES 2-4 Porolithon onkodes (Heydrich) Foslie, 1909:57.—Gordon, Ma- saki, and Akioka, 1976.—Lee, 1967.—Lemoine, 1966.— Taylor, 1950.—Womersley and Bailey, 1970. Litholhamnwn onkodes Heydrich, 1897a:6, pl. 1: fig. 11. Lithophyllum oncodes Heydrich, I897c:410. Goniolithon oncodes (Heydrich) Foslie, 1899:5. Lithophyllum onkodes (Heydrich) Foslie, 1900a:8, 1903a; 1907a,b.—Weber-van Bosse and Foslie, 1904. Porolithon oncodes (Heydrich) Foslie, 1909:57; 1929.—Littler, 1971b. Lithophyllum (Porolithon) oncodes (Heydrich) Foslie, 1909:38. DESCRIPTION.—Crusts well developed, lacking branches and excrescences, hemispherical in shape (Figure 2A), few mm to many cm thick, pink to yellow, with a rough appearance due to the abundance of heterocyst fields on the surface (Figure 2B,C). Epithallium a layer of rounded cells 1 to 3 cells thick (2-6 /xm long, 5-9 /xm diam.). Intercalary meristem large celled, occur- ring immediately below the epithallium (4-11 /mi long, 4-8 jum diam.) (Figure 3). Perithallium multilayered, fusions common, cells 4-13 /xm long and 4-10 ttm diam. Heterocysts, 10-30 /am long and 4-14 /im diam., throughout perithallium forming compact fields to 100 /xm diam. (Figure 2E,F). Hypothallium multilayered, plumose (Fig- ure 2F), 50-150 (350) /xm thick, may be thin with filaments oriented parallel to the substrate, cells 11-24 /xm long and 5-14 /xm diam. Tetrasporic conceptacles unipored, scattered, convex, small (240-300 /xm outside diameter (O.D.), 110-230 /xm inside diameter (I.D.), and 30-140 /xm high), with roof apex about 40 /tm above surface (Figure 2D), columella present; tetrasporangia 50-120 /xm long and 20-50 /xm diam., restricted to concep- tacle periphery. Cystocarpic conceptacles uni- pored, slightly raised above surface, 150-210 /xm I.D., 80-140 /xm high; carpospores (35-70 /xm long, 25-70 /xm wide) arising from the periphery SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • ' ... :•'- vr * -■ .» ■:'-■■:■ * ■ vT ■;>■,■ .■■■:■:-. .■ ' mf&fXi&Vfii&llSiMZlBEiaaiBP. C*V^A L-A"Ws .** '■■■ •'-i'V ^ . - FIGURE 2.—Porolithon onkodes: A, habit, X 1; B, crust with tetrasporic conceptacle (arrow) and heterocyst fields (arrowhead), X 7; c, crust with heterocyst fields, X 12; D, tetrasporic concep- tacle, X 250; E, heterocyst field at thallus surface, X 500; F, plumose hypothallium and buried heterocyst field (arrow), X 200. (Specimen nos.: A, B, 71-82-84; c, 71-82-85; D, 71-81-2; E, 71-82- 76; F, 71-59-28; micrographs reduced to 85%.) NUMBER 15 O 7 Z LU -1 6 _i _i LU U 5 o O o P. gardineri ( 4 plants ) O P. onkodes (25 plants) < Q —j i LU U ep 1 M 3 4 5 6 PERITHALLIUM FIGURE 3.—Mean epithallial and perithallial cell dimensions in Porolithon onkodes and P. gardineri. of a single discoid fusion cell. Male conceptacles unipored, slightly raised, 130-200 /xm I.D., 20-35 /tm high; spermatangial mother cells arising from the floor of the conceptacle only; spermatia at first elongate, crescent shaped, 3-5 /xm long, 1-2 /xm diam., becoming ellipsoidal. TYPE-LOCALITY.—On coral ridges, Tami Is- land, northwest edge of the Huon Gulf, New Guinea. HOLOTYPE.—Heydrich 97, March 1892, col- lected by Bambler, in herbarium of M. Foslie (TRH). DISTRIBUTION.—Borneo, Easter Island, Funa- futi Atoll, Guam, Hawaii, Marshall Islands, New Guinea, Solomon Islands, Sulu Archipelago, In- dian Ocean, Maldives and Laccadives, Red Sea. SPECIMENS STUDIED.—Hawaii: Hilo Bay, March 1971, 71-58-38, 71-59-28; Kawaihae, March 1971, 71-57-20. Midway: South Island, August 1971, 71- 82-76, 71-82-84, 71-82-85. Oahu: Kaneohe Bay, March 1971, 71-54-4; August 1971, 71-81-2; Wainae, March 1971, 71-52-9. REMARKS.—Only three sexual specimens were seen; all were monoecious. The male and female conceptacles are buried and occur at different levels in the thallus, indicating that P. onkodes inhibits self-fertilization by having different re- productive phases of the life cycle of any one plant maturing at different times. It was not clear whether there are two or four spermatia per spermatangial mother cell in P onkodes. No pro- 10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES carpic conceptacles were found in the Hawaiian specimens. In shallow water, P. onkodes was the single most abundant species collected (Figure 4); over 50 percent of intertidal collections were P. onkodes. Individual plants of this species as small as one to two square centimeters are easily identified by the pustulous heterocyst fields over the thallus. Although P. onkodes is dominant in nearly all wave-beaten intertidal or uppermost sublittoral situations, it reaches its "peak of development" on algal ridges (Doty, 1974). Here it tends to dominate with the branched Lithophyllum kot- schyanum Unger and the less abundant Porolithon gardineri (Foslie) Foslie occurring along the sides of channels and near low water. In the Caribbean, the very similar "pair species" (see "Glossary") Porolithon pachydermum (Foslie) Foslie and Litho- phyllum congestum (Foslie) Foslie occur in the same ecological niches (Adey, 1979; Steneck and Adey, 1976). Borings through a number of high ridges in St. Croix, U.S. Virgin Islands, show that Lith- ophyllum congestum has been the dominant Holo- cene builder of these ridges (Adey, 1975). On the other hand, borings through the higher energy algal ridges of the easternmost Lesser Antilles have shown that Porolithon pachydermum and Mil- lepora (Coelenterata) are the dominant builders of these more massive and higher algal ridges. Porolithon gardineri (Foslie) Foslie FIGURES 3-5 Porolithon gardineri (Foslie) Foslie, 1909:57.—Lee, 1967.—Lit- tler, 1971b.—Taylor, 1950. Lithophyllum gardineri Foslie, 1907a: 190; 1929. Lithophyllum (Porolithon) gardineri Foslie, 1907b:30; 1909. DESCRIPTION.—Crusts weakly developed, quickly producing branches initially simple, ter- ete, 1.5-3 mm diam., slightly tapering and with rounded tops (Figure 5A); branches typically be- coming complexly branched, often flattened, fused and anastomosing to produce 12 cm or larger hemispheric heads (quiet-water forms with delicate branches, 1-2 mm diam., are known and discussed below); pink to yellow, with a granular texture as in P onkodes; the heterocyst fields crowded and more pustulous than P. onkodes with a marked upturned epithallial rim. Epithallium a single layer of cells, 3-5 /xm long, 8-10 /xm diam. Perithallium multilayered, fusions common, cells 5-10 /xm long and 6-11 /xm diam (Figure 3). Heterocysts throughout perithallium, 13-26 /xm long and 7-15 /xm diam. Hypothallium not meas- ured. No conceptacles, asexual or sexual, were seen in sections of our material. TYPE-LOCALITY.—Coetivy, Seychelle Islands, Indian Ocean. LECTOTYPE.—Gardiner sn., September 1905, in herbarium of M. Foslie (TRH) (Adey, 1970). Lecto-isotype: USNC. DISTRIBUTION.—Hawaii, Indian Ocean, Mar- shall Islands. SPECIMENS STUDIED.—Oahu: Kanehoe Bay, March 1971, 71-54-5. Midway: March 1971, 71- 60-3; August 1971, 71-82-37. REMARKS.—As in P- onkodes, normal plants of Depth m 40 50 FIGURE 4.—Depth distribution of Porolithon onkodes (o) and P. gardineri (A) in Hawaii NUMBER 15 11 FIGURE 5.—Porolithon gardineri: A, habit of typical specimen, X 1; B, branch showing smooth surface from limpet grazing, X 10; c, branch showing characteristic heterocyst fields, X 10; D, habit of specimen from Midway Island lagoon, X 1. (Specimen nos.: A, B, 71-54-5(a); c, D, 71- 60-3; micrographs reduced to 96%.) 12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES P. gardineri have markedly distinct, pustulous het- erocyst fields (Figure 5c); however, micrograzing, probably by young limpets and perhaps chitons, can remove this pustulous surface (Figure 5B). Because of grazing effects on morphology, care must be taken not to misidentify this species with externally similar Lithophyllum kotschyanum. The chalky pink-yellow surface of P. gardineri, as op- posed to the red-brown, glossy surface of L. kot- schyanum, is usually a distinctive characteristic even in the absence of heterocyst fields. Steneck and Adey (1976) have shown with transplants that the marked variety of branch form in the Caribbean Lithophyllum congestum is a function of microenvironment. This plant had been separated into several species by Foslie (Ste- neck and Adey, 1976). A wide variety of branch form is also to be seen in the Hawaiian Porolithon gardineri and Lithophyllum kotschyanum (see below). At a single station in the lagoon of Midway Atoll, a fine-branched Porolithon (Figure 5D) that is ten- tatively placed in P. gardineri was found in abun- dance (it was not included in the measurements). Its surface texture is very similar to that of the P. gardineri found in the remainder of our stations, and the branching patterns are similar, though markedly smaller than P. gardineri, having diam- eters of 1-2 mm. As discussed above, P. gardineri is primarily a shallow sublittoral plant. In contrast to the situ- ation on central Pacific atolls (Lee, 1967), on Hawaiian algal ridges it is exceeded in abundance by the branching L. kotschyanum. The "peak" of abundance of P. gardineri appears at a depth of 18 m (Figure 4) and results largely from its occur- rence at that depth at a single station, on the south side of East Island, Midway Atoll. The small, secondary peak at 0 to 3 m is more char- acteristic for the species in the algal ridge envi- ronment. Paragomolithon, new genus DESCRIPTION.—Thallus crustosum ad fali- aceum, non ramosus. Epithallium ex unico strato cellularum rotundatarum constans, membranis exterioribus tangentialibus incrassatis. Perithal- lium pluristratosum, fusionibus frequentibus. Heterocystae in stratis horizontalibus plerumque laxe aggregatae e duabus tribusve cellulis, cellula exteriore protuberatione praedita compositae. Hypothallium pluristratosum, crassum, leniter ad valde coaxiale. Conceptacula tetrasporangialia uniporata; tetrasporangia adsunt, bisporangia rara, ad periferiam conceptaculi restricta. Con- ceptacula spermatialia uniporata; cellulae-ma- tricales spermatangiales duas ad quattuor excres- centias spermatiales habentes, ad pavimentum conceptaculi restrictae; spermatia ellipsoidae. Plantae carpogoniales cystocarpicaeque rarae. Thallus crustose to leafy, unbranched. Epithal- lium a single layer of rounded cells; outer tangen- tial wall thickened. Perithallium multilayered, fusions common. Heterocysts generally loosely grouped into horizontal fields composed of 2-3 cells, outer cell with protuberance. Hypothallium multilayered, thick, weakly to strongly coaxial. Tetrasporangial conceptacles uniporate; tetra- sporangia present, bisporangia rare, restricted to the periphery of the conceptacle. Spermatial con- ceptacles uniporate; spermatangial mother cells with 2-4 spermatial outgrowths; restricted to the floor of the conceptacle; spermatia ellipsoidal. Carpogonial and cystocarpic plants rare. TYPE-SPECIES.—Paragomolithon solubile (Foslie & Howe in Foslie), new combination. Basionym: Goniolithon solubile Foslie & Howe in Foslie, 1907c:21; Boyd, Kornicker, and Rezak, 1963; Howe, 1920; Taylor, 1928, 1960. Recom- binations as follows: Neogoniolithon solubile (Foslie & Howe in Foslie) Setchell & Mason, 1943:90.—Adey, 1970. Goniolithon propinquum Foslie, 1929:31. Neogoniolithon propinquum (Foslie) Lemoine, 1966:14.—Papen- fuss, 1968. TYPE-LOCALITY.—Culebra Island, Puerto Rico. LECTOTYPE.—Howe, 4375, in herbarium of M. Foslie (TRH) (Adey, 1970). Lecto-isotype: USNC. DISTRIBUTION.—Tropical seas, typically at moderate depths, 5-25 m, in reef environments. NUMBER 15 13 Paragomolithon conicum (Dawson), new combination FIGURES 6, 7 Hydrolithon conicum Dawson, 1960a:27. [Basionym.] Neogoniolithon conicum (Dawson) Gordon, Masaki, & Akioka, 1976:259.—Zhang and Zhou, 1980. DESCRIPTION.—Crusts firmly adherent, 0.5 to several mm thick, unbranched (Figure 6A), often draped over the basal parts of finger corals; fre- quently deep red but ranging from yellow-pink to maroon, margin smooth to undulating, within several mm to 1 cm from the margin very abun- dant nonpustulous heterocyst fields develop (50- 100 /xm diam) (Figure 6A,B) giving the surface a slightly rough appearance. Epithallium a single layer of rounded cells, 3-9 /xm diam. Intercalary meristem cells elongate, cells 4-9 /xm long and 5- 10 /xm diam., with some progressive elongation occurring throughout the perithallium (Figure 7). t'^\$ti&cK-i ***** ■ - > ■■"• *4 '" '■■ v ' FIGURE 6.—Paragomolithon conicum: A, habit of typical specimen, X 1; B, surface features including tetrasporangial conceptacles, X 10; c, section through thallus showing heterocyst field, X 200; D, section through vegetative thallus in region of heterocyst fields (arrow), X 40; E, section through tetrasporangial conceptacle, X 250; F, section through vegetative thallus showing coaxial hypothallium, X 35. (Specimen nos.: A, B, E, 71-62-32; c, D, 71-78-13; F, 71-55-35; micrographs reduced to 70%.) 14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ep PERITHALLIUM 2 3 4 5 6 7 £ 6 P. conicum (20 plants) 6- 5 ■ "2 20 P.conicum 20 80 30 60 90 40 50 Depth m FIGURE 7.—Paragomolithon conicum: A, mean epithallial and perithallial cell diameter and length; B, depth distribution. Perithallium multilayered, fusions abundant, cells 3-11 jum long and 4-13 /xm diam. (Figures 6c, 7). Heterocyst fields throughout the perithal- lium, loose, with scattered filaments between the heterocysts (Figure 6C,D), heterocysts 7-26 /xm long and 8-21 /xm diam. Hypothallium parallel to substrate, weakly to strongly coaxial, 30-400 /xm thick; cells 8-25 long and 4-16 /xm diam. (Figure 6F). Tetrasporic conceptacles unipored, regularly spaced, convex, conical, small (350-530 /xm O.D., 140-360 /xm I.D., 50-160 /xm high), strongly raised, roof apex 40-200 /xm above plant surface (Figure 6E), columella present; tetraspo- rangia restricted to the periphery of the concep- tacle, 80-200 /xm long, 45-65 /xm diam. No pro- carpic or cystocarpic material found in our spec- imens. Male conceptacle unipored, 190-260 /xm I.D., 40-80 /xm high, raised but more rounded than tetrasporangial conceptacle; spermatangial mother cells arising from the base of the concep- tacle only; spermatia ellipsoidal, 3-6 /xm long, 1- 3 /xm diam. TYPE-LOCALITY.—Intertidal reef at Binners Cove, Isla Socorro, Revillagigedo Archipelago, Pacific Mexico. HOLOTYPE.—Dawson, 12148, 19 November 1953, (US). DISTRIBUTION.—Xisha Islands, China; Guam; Pacific Mexico. SPECIMENS STUDIED.—French Frigate: La Pe- rouse, August 1971, 71-78-13. Hawaii: Honaunau Bay, March 1971, 71-55-34, 71-55-35. Midway: Lagoon, March 1971, 71-62-32. Oahu: Honauma Bay, March 1971, 71-53-7; Waikiki, March 1971, 71-50-45. REMARKS.—The type collection (Dawson, NUMBER 15 15 1960a) includes only bisporic specimens. Bisporic plants were not found in our collections; we did, however, have tetrasporic plants. The description of P. conicum should therefore be emended to include tetrasporic plants. Formation of the tet- rasporic conceptacles follows the "sur-type" of Johansen (1976) with the presence of a central columella (see Townsend, 1981). Only one sexually reproductive specimen, a male, was collected. Formation of the male con- ceptacle is similar to that described for Litho- thamnium by Lebednik (1978). The first indication of formation of the male conceptacle is cessation of an area of intercalary meristem cells. These cells become the prospective spermatangial mother cells (PSMC). The epithallium separates from the PSMC layer, which changes from a columnar to conical shape. The surrounding veg- etative filaments continue dividing and grow in over the fertile area forming the conceptacle roof. As the roof forms, the apex of the PSMC elongates and "cuts off a spermatium into the conceptacle cavity. It is not clear how many times this may occur per mother cell. Spermatangial production is restricted to the conceptacle floor. The sper- matia are at first ellipsoidal, then discoid, and may congregate in long mucus streams in the conceptacle cavity. The single pore of the concep- tacle remains plugged during spermatial produc- tion with a substance not stained with phospho- tungstic hemotoxylin. This species is illustrated on the "Finger Coral: Hawaii" coral reef commemorative stamp issued by the U.S. Postal Service, 26 August 1980. Paragomolithon conicum occurs commonly in mid- depths (Figure 7) at all stations from Midway to Hawaii. From 9 to 15 m, P. conicum and P. onkodes were the most frequent crustose coralline algae (19% of the specimens collected). Although P. conicum occurs on a variety of "coralgal" sub- substrate, it appears to especially prefer the sides of finger corals, and by gradually growing up from below, may eventually kill living coral. Two species oi Paragomolithon, P. solubile and P. "typica" (nomen nudum: Adey, 1979) occur in the Caribbean and occupy similar substrate and depth ranges (Adey, 1979); however, unlike many of the "pair species" discussed here, P. conicum seems to be rather distinct from its Caribbean counterparts. This would indicate that Paragomo- lithon has evolved more rapidly than most tropical coralline algae and that their habitat, "sides of finger corals," is relatively new for coralline algae. Neogoniolithon Setchell & Mason, 1943 Key to the Species 1. Plants thin crusts, rugulose; low-domed conceptacles < 200 /xm O.D. N. rugulosum, new species Plants thicker, not rugulose, conceptacle > 200 /xm O.D 2 2. Crusts adherent, knobby, bright pink to yellow brown, conceptacles 200- 400 /xm O.D N. rufum, new species Crusts leafy to sub-leafy, pink yellow to dark brown, conceptacles > 400 /xm O.D 3 3. Crusts sub-leafy, but eventually developing abundant rounded, often club- shaped irregular protuberances, conceptacles high conical, leaving scars on surface after spores dispersed N. clavacymosum, new species Crusts sub-leafy, becoming thick (to 5 mm) without protuberances, con- ceptacles low conical, leaving no scars N. fosliei 16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 8.—Neogoniolithon rugulosum, new species: A, habit of type specimen, X 1; B, hypothallium of type specimen, X 500; c, surface showing rugulose nature of crust, note conceptacles (arrowed), X 10; D, tetrasporangial conceptacle, X 600. (Specimen nos.: A, B, 71-53-2; c, 71-59- 1; D, 71-53-4.) NUMBER 15 17 Neogoniolithon rugulosum, new species FIGURE 8 DESCRIPTION.—Crustae tenues (< 200 /xm), ru- gulosae, maculas subtiles albas saepe praebentes; multa conceptacula parva tholiformia, aut eorum cavos (Figura 8A,C) haventes, colore rosacea ad flavo-rosaceam. Epithallium ex uno duobusve stratis cellularum tholiformium constans, mem- brana in membranis exterioribus tangentialibus radialibusque spissescens; cellulae 5-12 /xm long, atque 9-16/xm diam. Meristema intercalare elon- gatum, admodum infra epithallium, cellulis 10- 19 fim long, atque 6-10 /xm diam. Perithallium multis cellulis crassum (Figura 8B), fusionibus frequentibus, cellulis 4-8 /xm long atque 3-6 /xm diam. Heterocystae per perithallium dispersae, singulae, raro in coacervationibus horizontalibus repertae, 12-14 /xm long atque 7-9 /xm diam. Hypothallim e 2-8 stratis celularum constans (Figura 8B), subparallelum ad substratum, 25-75 /xm crass.; cellulae 9-30 /xm long atque 6-15 /xm diam. Conceptacula tetrasporica uniporata (160- 200 /xm O.D., 70-90 /xm I.D., 30-50 /xm alt.) tecta plana ad paululum tholiformia super crustas cir- cumdantes; tetrasporangia trans pavimentum conceptaculi disposita, 50-85 /xm long atque 25- 50 /xm diam (Figura 8D). Crusts thin (< 200 /xm), rugulose, often with fine white speckles; abundant small domed con- ceptacles or their cavities (Figure 8A,C); pink to yellow pink. Epithallium 1 to 2 layers of domed cells, wall thickening on outer tangential and radial walls; cells 5-12 /xm long and 9-16 jum diam. Intercalary meristem elongate, immedi- ately below epithallium, cells 10-19 /xm long and 6-10 /xm diam. Perithallium, many cells thick (Figure 8B), fusions common, cells 4-8 /xm long and 3-6 /xm diam. Heterocysts scattered through- out perithallium, single rarely in horizontal as- semblages, 12-14 /xm long and 7-9 /xm diam. Hypothallium 2-8 cell layers (Figure 8B), sub- parallel to substrate, 25-75 /xm thick; cells 9-30 /xm and 6-15 /xm diam. Tetrasporic conceptacles single pored, 160-200 /xm O.D., 70-90 /xm I.D., 30-50 /xm high, roofs flat to slightly domed above surrounding crusts; tetrasporangia across floor of conceptacle, 50-85 /xm long and 25-50 /xm diam. (Figure 8D). NO sexual material collected in present study. TYPE-LOCALITY.—Palea Point, Hanauma Bay, Oahu, Hawaii (21o07'N,157°50'W), on volcanic rock, intertidal zone. HOLOTYPE.—D. Child, 71-53-2, 1 April 1971 (USNC), Figure 8A,B PARATYPES.—Hawaii: Hilo Bay, March 1971, 71-59-1. Oahu: Honauma Bay, March 1971, 71- 53-22, 71-53-4. DISTRIBUTION.—Hawaii and Oahu, Hawaii. REMARKS.—The specific epithet rugulosum de- scribes the surface, a feature distinguishing our new taxon from other Hawaiian members of the genus. We have only a few specimens of this species from bedrock and from volcanic and terrigenous pebbles in the intertidal and uppermost sublit- toral. Its sparseness in our collections may, how- ever, only indicate considerable difficulty in col- lecting it from wave-beaten rocks rather than its rarity. A "pair species," Neogoniolithon caribaeum (Foslie) Adey, is known from the Caribbean (Adey, 1979). Neogoniolithon rufum, new species FIGURES 9-1 IA DESCRIPTION.—Crustae primum satis tenues, marginibus arete adhaerentibus (Figura 9A), 1-2 mm crass, factae, superficies saepe clare rosacea ad fusco-brunneam, autem, varians, levis, opales- cens (Figura 9B), conceptacula super thallum ca- tervatim aggregata. Epithallium ex uno strato cellularum rotundatarum constans, incrassa- tiones membranae esterioris tangentialis perspi- cuae; cellulis 2-6 /xm long, atque 4-9 /xm diam. Meristema intercalare parum elongatum, cellulis 3-9 /xm long, atque 3-10 /xm diam. Perithallium pluristratosum, aliquantula elongatione effecta (Figura 10), fusionibus frequentibus, abrupte zon- atum (Figura 9A), cellulis 3-11 /xm long, atque 2- 11 /xm diam. Heterocystae singulae, interdum 18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES «1,J|. *'* FIGURE 9.—Neogoniolithon rufum, new species: A, habit of type specimen, X 1; B, thallus surface, note strongly raised tetrasporic conceptacles, X 10; c, section through monoecious crust, note male conceptacle (arrow) and cystocarpic conceptacle (arrowhead), X 40; D, hypothallium, X 350. (Specimen nos.: A, B, 71-50-80; c, 71-82-57; D, 71-50-20; micrographs reduced to 89%.) NUMBER 15 19 ep M 1 PERITHALLIUM 2 3 4 5 10 O Z LU 5 D N . r uf um (31 plants) ° N. clavacymosum (24 plants) A N . fosliei (12 plants) < 6 9 U FIGURE 10.—Cell diameter and length data for Neogoniolithon rufum, new species, N clavacymosum, new species, and N. fosliei. rarae (aliqunado laxe verticaliter horizontali- terque aggregata) 7-29 /xm long, atque 4-12 /xm diam. Hypothallium simplex ad coaxiale, ad sub- stratum parallelum (Figura 9D), 25-130 (200) /xm crass., cellulis 9-25 /xm long, atque 4-10 /xm diam. Conceptacula tetrasporica uniporata, 30-150 /xm super crustam elevata (Figura 9B), 200-400 /xm O.D., 80-300 /xm I.D. atque 50-110 /xm alt., per perithallium obruta; tetrasporangia 50-175 /xm long, atque 29-65 /xm diam. super totum pavi- mentum conceptaculi disposita. Conceptaculum procarpicum uniporatum, 380/xm I.D., atque 100 /xm alt., omnis cellula sustinens duo initia carpo- gonialia ramorum habens. conceptacula cystocar- pica uniporata, 210-290 /xm I.D. atque 60-90 /xm alt., per perithallium cum sporis obruta; carpo- sporae 50-70 /xm long, atque 40-70 /xm diam., solum in periferia cellulaecoalescentis magnae 20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES discoideae dispositae (285-340 /xm diam.) (Figura 9c). Conceptaculum masculum uniporatum, 50- 130 /xm elevatum, 215-350 /xm I.D. atque 50-100 /xm alt.; spermatia ad pavimentum conceptaculi restricta (Figura 9c); 2-4 spermatia in omni cel- lulamatricalee, ellipsoidea ad discoidea (3-7 /xm long., 2-3 /xm diam.). Crusts initially quite thin with closely adherent margins (Figure 9A), becoming 1-2 mm thick, surface often bright pink though ranging to dark brown, smooth, opalescent (Figure 9B), concep- tacles distributed in groups over the thallus. Ep- ithallium a single layer of rounded cells, outer tangential wall thickenings marked; cells 2-6 /xm long and 4-9 /xm diam. Intercalary meristem weakly elongate, cells 3-9 /xm long and 3-10 /xm diam. Perithallium multilayered, some elonga- tion occurring (Figure 10), fusions common, sharply zonate (Figure 9c), cells 3-11 /xm long and 2-11 /xm diam. Heterocysts single, sometimes rare (occasionally grouped loosely vertically and horizontally) 7-29 /xm long and 4-12 /xm diam. Hypothallium simple to coaxial, parallel to sub- strate (Figure 9), 25-130 (200) /xm thick, cells 9- 25 /xm long and 4-10 /xm diam. Tetrasporic con- ceptacles unipored, raised 30-150 /am above the crust (Figure 9B), 2001-400 /xm O.D., 80-300 /xm I.D., and 50-110 /xm high, buried throughout perithallium; tetrasporangia 50-175 /xm long and 29-65 /xm diam., over entire conceptacle floor. Procarpic conceptacle unipored, 380 /xm I.D. and 100 /xm high, 2 carpogonial branch initials per supporting cell. Cystocarpic conceptacles uni- pored, 210-290 /xm I.D. and 60-90 /xm high, buried throughout perithallium with spores; car- pospores 50-70 /xm long and 40-70 /xm diam., only on the periphery of the large discoid fusion cell (285-340 /xm diam.) (Figure 9c). Male con- ceptacles unipored, raised 50-130 /xm above sur- face, 215-350 /mi I.D. and 50-100 /xm high; spermatia restricted to conceptacle floor (Figure 9c), 2-4 spermatia per mother cell, ellipsoidal to discoid 3-7 jum long, 2-3 /xm diam. TYPE-LOCALITY.—Waikiki, Oahu, Hawaii (21°15'N, 157°35'W), on coral rubble, 130 m. HOLOTYPE.—D. Child, 71-50-80, 27 March 1971 (USNC), Figure 9A. PARATYPES.—French Frigate: La Perouse, August 1971, 71-78-14. Hawaii: Hilo Bay, March 1971, 71-58-14. Oahu: Kaneohe Bay, August 1971, 71- 81-28; Waikiki, March 1971, 71-50-20, 71-50-31, 71-50-72, 71-50-80. Midway: South Island, August 1971, 71-82-57. DISTRIBUTION.—Throughout Hawaiian Archi- pelago. REMARKS.—The specific epithet refers to the unique color of N. rufum. The formation of the male conceptacles is sim- ilar to that for Paragomolithon conicum. In Neogo- niolithon rufum there are two to four spermatangial projections from each mother cell. As in P. conicum, spermatia aggregate into chains within the con- ceptacle cavity. Carpospores are restricted to the periphery of the cystocarpic conceptacle. This contrasts to the pattern seen in Neogoniolithon ac- cretum (Foslie & Howe) Setchell & Mason and Neogoniolithon pacificum (Foslie) Setchell & Mason (Masaki, 1968). Monoecious thalli with cystocarpic and sper- matangial conceptacles were seen (Figure 9c). Both conceptacle types were buried with contents in the thallus but do not show layering as in Porolithon onkodes. One hermaphroditic concepta- cle was seen. This phenomena in crustose coral- lines has only been recorded for Phymatolithon lenormandii (Areschoug) Adey (Adey, 1966) and Synarthrophyton patena (J.D. Hooker & Harvey) Townsend (Townsend, 1979). In Neogoniolithon rufum half the conceptacle was male, and the other half procarpic. Only one procarpic concep- tacle was seen. Two carpogonial branch initials arise from the support cell. One or both of these may extend and undergo division to form a car- pogonium and hypogynous cell. Therefore, N. rufum has procarps with two carpogonial branches or one carpogonial branch and one sterile cell per supporting cell. This pattern differs from that of Neogoniolithon accretum and N. pacificum (Masaki, 1968) with only one carpogonial branch per pro- carp. Neogoniolithon rufum, Paragomolithon conicum, and NUMBER 15 21 Tenarea tessellatum together dominate the shallow to mid-depth coralline flora, although P. onkodes is abundant in water less than 20 m. The former three species occur on 25% of the collected speci- mens from low water to depths of nearly 50 m. Neogoniolithon rufum tends to be a little more abun- dant at the lower part of this range than the other species (Figure 11A). Also, while the sympatric Paragomolithon conicum and Tenarea tessellatum have Caribbean "pair species," N. rufum does not. Neogoniolithon clavacymosum, new species FIGURES 10, 11B, 12 DESCRIPTION.—Crustae bene evolutae amplae factae, ramos breves simplicesque sparsos, qui irregulariter clavati atque saepe nodulosi fiunt (Figura 12A,B) efficientes; margines leves cras- sique, foliosi et ad substratum non adhaerentes facti; conceptacula magna, conica, sparsa aut in cacuminibus ramorum crebra, saepe erupta et sic depressiones in superficie relinquentia quae oram crassam elevatam interdum habent (Figura 12B). Epithallium una duabusve cellulis crassum, cel- lulis tholiformibus, sine incrassationibus mem- branae, 2-4 /xm long, atque 4-8 /xm diam. Mer- istema intercalare breve, 2-6 /xm long, atque 3-7 /xm diam. Perithallium plurisstratosum, gradatim plus elongatum factum (Figura 10), corpora tinc- tabilia et fusiones adsunt, cicatrices conceptacu- lorum anticedentium visibiles, cellulis 2-10 /xm long, et 3-11 /xm diam. Heterocystae 5-15 /xm long, atque 4-11 /xm diam. Hypothallium plur- istratosum, coaxiale (Figura 12c), 20-250 /xm crass.; cellulae 9-28 /xm long, et 3-12 /xm diam. Conceptacula tetrasporica uniporata, 100-320 /xm super superficiem circumanbientem thalli ele- Depth m 10 20 30 Depth m o ic 20 30 40 50 60 70 80 90 ative ance % N . fosl i e i OL 3 or < 0 FIGURE 11.—Depth distribution: A, Neogoniolithon rufum, new species; B, N. clavacymosum, new species; c, N. fosliei. 22 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 12.—Neogoniolithon clavacymosum, new species: A, habit of type specimen, X 1; B, surface of type specimen, note large uniporate conceptacles (arrow) and conceptacle scars (arrowhead), X 5; c, section showing thick coaxial hypothallium and thick zonate perithallium, X 200. (Specimen nos.: A, B, 71-58-78; c, 71-58-42.) vata, 450-600 /xm O.D., 200-500 /xm I.D. atque 50-170 /xm alt.; tetrasporangia ad periferiam con- ceptaculi restricta (columella adest), 40-90 /xm long, atque 15-35 /xm diam. Thalli monecii fre- quentes. Thalli procarpiales non visi. Concepta- culum cystocarpicum uniporatum 340 /xm I.D. atque 120 /xm alt.; cellula-coalescentis 230 /xm diam.; carposporae solum in periferia concepta- culi repertae, 39-60 /xm long, atque 25-58 /xm diam. Conceptacula mascula uniporata, elevata, 310-440 /xm I.D. atque 50-110 /xm alt.; sperma- tangia ad pavimentum conceptaculi restricta; spermatia discoidea 3-5 /xm diam. Crusts well developed, becoming extensive, producing scattered short simple branches that become irregularly club shaped and often nodular NUMBER 15 23 (Figure 12A,B); margins smooth and thick, becom- ing leafy and nonadherent to the substrate; con- ceptacles large, conical, scattered, or concentrated on branch tips often breaking out to leave surface depressions, sometimes having thick raised rims (Figure 12B). Epithallium 1 to 2 cells thick; cells domed, no wall thickenings, 2-4 /xm long and 4- 8 /xm diam. Intercalary meristem short, cells 2-6 /xm long and 3-7 /xm diam. Perithallium multi- layered, shows progressive elongation (Figure 10), staining bodies present, fusions present, scars from old conceptacles visible, cells 2-10 /xm long and 3-11 /xm diam. Heterocysts 5-15 /xm long and 4- 11 /xm diam. Hypothallium multilayered, coaxial (Figure 12c), 20-250 /xm thick; cells 9-28 /xm long and 3-12 /xm diam. Tetrasporic conceptacles unipored, raised 100-320 /xm above the surround- ing thallus surface, 450-600 /xm O.D., 200-500 jum I.D., and 50-110 /xm high; tetrasporangia restricted to the conceptacle periphery (columella present), 40-90 /xm long and 15-35 jum diam. Monoecious thalli common. Procarpial thalli not seen in our collections. Cystocarpic conceptacle unipored, 340 /xm I.D. and 120 /xm high; fusion cell discoid 230 /xm diam.; carpospores only on periphery of conceptacle, 39-60 /xm long and 25- 58 /xm diam. Male conceptacles unipored, raised, 310-440 /xm I.D. and 50-110 /xm high; sperma- tangia restricted to the conceptacle floor; sper- matia discoid 3-5 /xm diam. TYPE-LOCALITY.—Hilo Bay, Hawaii, Hawaii (19°45'N, 155°0'W), windward side of reef, 45 m depth. HOLOTYPE.—D. Child, 71-58-78, March 1971 (USNC), Figure 12A,B. PARATYPES.—French Frigate: La Perouse, August 1971, 71-78-17. Hawaii: Hilo Bay, March 1971, 71-58-42, 71-58-78, 71-59-36; Kawaihae, March 1971, 71-57-18. Oahu: Kaneohe, August 1971, 71- 81-12, 71-81-24; Waianae, March 1971, 71-52-1, 71-52-3. DISTRIBUTION.—Throughout southern Ha- waiian Archipelago. REMARKS.—The specific epithet refers to the branch shape in N. clavacymosum. Spermatial development is similar to that de- scribed for Paragomolithon conicum. Spermatangial and cystocarpic conceptacles are not buried in thallus as in Porolithon onkodes. Tetrasporangial conceptacle development is similar to that described for Paragomolithon conicum. Two to three tetrasporangial rows, with inter- spersed sterile paraphyses, are formed on the periphery of the conceptacle. A central columella of sterile paraphyses is present. Neogoniolithon clavacymosum occurs at a frequency of about 10% throughout the shallow mid-depth range of 6-30 m (Figure 11B). The peak at 50 m (Figure 11B) is probably not real but rather a reflection of our minimal collecting in that par- ticular zone. In the Caribbean, N. clavacymosum is represented by a very similar but as yet unnamed and poorly known "pair species," which appears only sporad- ically in USNC Caribbean collections (Adey, un- published data). Neogoniolithon fosliei (Heydrich) Setchell & Mason FIGURES 10, lie, 13 Neogoniolithon fosliei (Heydrich) Setchell & Mason, 1943:90.— Lemoine, 1965.—Womersley and Bailey, 1970.—Gordon et al., 1976.—Zhang and Zhou, 1980. Lithothammon fosliei Heydrich, 1897b:58. Lithophyllum fosliei (Heydrich) Foslie, 1897c:410. Archaeolithothamnwn fosliei (Heydrich) Foslie, 1898:4. Goniolithon fosliei (Heydrich) Foslie, 1903a:470.—Weber-van Bosse and Foslie, 1904.—Howe, 1918b.—Setchell, 1926. [See also Lemoine, 1966, for distributional notes.] DESCRIPTION.—Thick, extensive crusts, up to at least 5 mm thick (Figure 13A), surface dull and minutely rather rugulose, abundant large raised conceptacles dispersed evenly over surface, break- ing out to leave strong scars (Figure 13A,B), often consisting of white patches or rings of dead cells; light pink on vegetative surface but attaining yellowish cast on conceptacle domes. Epithallium 2 or 3 cell layers, usually only lowest cell layer has cytoplasmic contents; cell production not syn- chronous, cells domed with outer tangential wall thickening of same dimensions as cell lumen, 2- 24 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 13.—Neogoniolithon fosliei: A, habit of typical specimen, X 1; B, thallus surface with conceptacles (arrow) and conceptacle scars (arrowhead), X 10; c, section through vegetative thallus, note heterocysts (arrow) and rounded epithallial cells (arrowhead), X 300; D, section through coaxial hypothallium, X 200; E, section through tetrasporangial conceptacle, X 100. (Specimen nos.: A, B, E, 71-58-47; c, D, 71-78-20.) NUMBER 15 25 12 /xm long and 5-13 /xm diam. Intercalary mer- large cells both in the meristem and the resulting istem elongate, cells 5-29 /xm long and 4-15 /xm perithallium, a character setting it apart from diam. Perithallial tissue irregular, little cell elon- other Hawaiian members of Neogoniolithon. Some gation (Figure 10), multilayered; cells 4-18 /xm of the irregularity of cell size seen in Figure 10 is long and 4-15 /xm diam. Heterocysts, single, large due to the small number of measurements. The (Figure 13c), 8-49 fim long and 5-24 diam. Hy- ver7 large and abundant heterocysts are in part pothallium parallel to coaxial, 50-200 /xm thick responsible for the tissue irregularity. The gradual (Figure 13D); cells 12-35 /xm long and 5-19 /xm reduction of cell lumen diameter with depth in diam. Tetrasporic conceptacles unipored, raised the perithallium is rather unusual among coral- 24-300 /xm above thallus, 700-1500 /xm O.D, lines' beinS previously described only for two 240-900 /xm I.D, and 90-190 /xm high; tetraspo- crustose coralline parasites, Ezo epiyessoense Adey, rangia borne over entire floor, 115-160 /xm long Masaki> & Akioka and Kvaleya ePllaeve AdeY & and 45-100 /xm diam. (Figure 13E). No sexual Sperapani (Adey, Masaki, and Akioka 1974; conceptacles seen. Adey and SPerapam, 1971), and Mesophyllum lich- ry T p, T D , c enoides Lemoine (Adey and Adey, 1973). lYPE-LiOCALiTY.—iM lor, rCed Sea. Ar . .... ...... . . . . .. TT TT i-irrv-i i • r-i.* Neogoniolithon fosliei is a relatively shallow-water HOLOTYPE.—Heydrich, 59, in herbarium of M. . , , , , \ . , +u species being largely restricted to depths less than T* ^ 25 m (Figure 1 lc). It was not found even in small DiSTRiBUTiON.-Borneo, Guam, Maldives and quantities at depths greater than 30 m as oppoSed Laccadives, Murray and Cocos Keeling islands, to Guam? where it has been recorded to 35 m Red Sea, Solomon Islands, Tahiti, Xisha Islands. (Gordon et al, 1976). In the Caribbean, N. fosliei SPECIMENS STUDIED.—French Frigate: La Pe- is represented by the "pair species" Neogoniolithon rouse, August 1971, 71-78-20. Hawaii: Hilo Bay, "megacarpum" (nomen nudum, Adey, 1979). The March 1971, 71-58-23, 71-58-47. Midway: South latter plant is quite similar in color, anatomy, Island, August 1971, 71-82-42. Oahu: Waikiki, conceptacle size, and ecology, though it rarely March 1971, 71-50-4. seems to obtain the thallus thickness seen in N REMARKS.—Neogoniolithon fosliei has relatively fosliei. Hydrolithon (Foslie) Foslie, 1909 Key to the Species 1. Thallus branched or mammillate 2 Thallus crustose 3 2. Bluish or purplish, mammillate; large perithallial cells (mostly >12 /xm long) H. reinholdii Pink, simple irregular branches; small perithallial cells (mostly <12 /xm long) H. breviclavium 3. Crust thin (<200 /xm), pink-yellow, conceptacles, scattered, and dis- tinct H. laeve, new species kachary and Ganesan, 1966.—Littler, 1971b; 1973a,b.— Gordon et al., 1976. Lithophyllum reinboldu Weber-van Bosse & Foslie in Foslie, 1902a:5; 1903a. Lithophyllum cerebelloides Heydrich, 1901c:405. Goniolithon reinboldu (Weber-van Bosse & Foslie in Foslie) Foslie in Weber-van Bosse and Foslie, 1904:49. Hydrolithon reinholdii (Weber-van Bosse & Foslie) Foslie FIGURES 14-16A Hydrolithon reinboldu (Weber-van Bosse & Foslie in Foslie) Foslie, 1909:55.—Dawson, 1954b; 1960b; 1961a.—Desi- Crust thicker (to several mm), irregular, pink, conceptacles abundant, small and not distinct H. megacystum, new species 26 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Porolithon reinholdii (Weber-van Bosse & Foslie in Foslie) Lemoine, 1911:166. DESCRIPTION.—Thick well-developed tessellate crusts (Figure 14B), smooth to highly mammillate (2-10 mm diam.) (Figure 14A), red-brown when exposed to much light, light pink-lavender on shaded surfaces. Epithallium a single layer of rectangular cells, 3-5 /xm long, 6-11 /xm diam. Intercalary meristem slightly elongate; cells 9-14 /xm long and 5-9 /xm diam. Perithallium multi- layered, irregular, fusions massive (Figure 14c) common deeper in the tissue; cells 5-20 /xm long and 5-17 /xm diam. (Figure 15). Heterocysts not abundant, single or sometimes arranged in small groups, 12-30 /xm long, 9-21 /xm diam (Figure 14E). Hypothallium a single layer of cells (Figure 14C,F), fusions common, cells 11-30 /xm long and 5-17 /xm diam. Tetrasporic conceptacles uni- pored, grouped, pore plugged with mucilage, con- ceptacle reef sunken to slightly raised (20-50 /xm), 200-600 /xm O.D, cavities 200-340 /xm I.D. and 50-160 /xm high (Figure 14D); tetrasporangia re- stricted to the periphery of the conceptacle (col- umella present), 80-100 /xm long and 40-75 /xm diam. No sexual material sectioned. TYPE-LOCALITY.—Mocaras reef, inner side, east coast Borneo. HOLOTYPE.—Weber-van Bosse, 22 June 1899, in herbarium of M. Foslie (TRH). Isotype: USNC. DISTRIBUTION.—Borneo, El Salvador, Guam, Hawaii, India, Maldives and Laccadives, Pacific Mexico, Phillipines, Timor, Vietnam. SPECIMENS STUDIED.—Hawaii: Honaunau, March 1971, 71-55-67A; Kawaihae, March 1971, 71-57-9. Oahu: Kaneohe, March 1971, 71-54-11, 71-54-18, 71-54-19, 71-54-21; Waikiki, March 1971, 71-50-14. REMARKS.—A question of priority of the bas- ionym Lithophyllum cerebelloides Heydrich (1901c) over Lithophyllum reinboldii Foslie & Weber-van Bosse in Foslie (1902a) has arisen. Both publica- tions were available in 1901; Foslie (1902a) on 10 July 1901 (as reprints), and Heydrich (1901c) on 24 July 1901 (as a journal). Foslie 1902a was published in a volume of Det Kongelige Norske Videnskabers Selkabs Skrifter in 1902. Under the International Code of Botanical Nomenclature, article 11, L. reinboldii has priority over L. cerebelloides. This species was sporadically abundant on shallow reef flats (0-10 m) and rubbly fore reefs throughout the Hawaiian chain. Hydrolithon rein- boldii is sharply restricted to shallow water (Figure 16A), no specimens being found over 15 m depth. In Hawaiian waters, it is the only important shallow-water coralline alga involved in the for- mation of rhodoliths. See also discussion under//. megacystum, new species (below). The Caribbean Hydrolithon b0rgesenii (Foslie) Foslie is quite similar to H. reinboldu in color, surface texture, and anatomy, though it is not usually so strongly mammillate as the Hawaiian plant. Also, H. b0rgesenii has a very wide depth range and, in the Caribbean, appears to be the ecological equivalent of all of the Pacific Hydro- lithon species. Hydrolithon b0rgesenii is also impor- tant in the formation of shallow-water rhodoliths, though in that role it is not as abundant as the Caribbean species Neogoniolithon mammillare (Harvey) Setchell & Mason. Hydrolithon breviclavium (Foslie) Foslie FIGURES 15, 16B, 17 Hydrolithon breviclavium (Foslie) Foslie, 1909:56.—Littler, 1973b. Goniolithon (Hydrolithon) breviclavium Foslie, 1907b:20. DESCRIPTION.—Well-developed crusts with thick even margins adhering well to substrate, branched (Figure 17c) or mammillate (Figure 17A,B), branches short; surface similar to Hydroli- thon reinboldii but more finely tessellate (Figure 17E), conceptacles moderately large, raised, evenly but not densely scattered over surface (Figure 17E); bright to dusty pink, deep-water plants sometimes brown. Epithallium single lay- ered cells ellipsoidal, with thickened outer tan- gential wall, cells 2-10 /xm long and 5-12 /xm diam. Intercalary meristem cells 3.5-12 /xm long and 3.5-10 /xm diam. Perithallium multilayered, generally disorganized below third cell layer, fre- NUMBER 15 27 . - .; ._■ ._ D Ss^SgSgj^ •■•A \ IP* ^..- * '«i>i >. . r FIGURE 14.— Hydrolithon reinboldu: A, habit, X 1; B, surface of thallus showing conceptacles and tesselate pattern, X 10; c, transverse section through thallus, note single-layered hypothallium, X 5; D, tetrasporangial conceptacle, X 200; E, section through thallus showing heterocyst, intercalary meristem, and epithallium, X 500; F, hypothallium, X 300. (Specimen nos.: A, 71- 54-21; B, 71-54-19; c, D, 71-50-14; E, F, 71-54-11; micrographs reduced to 91%.) 28 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ep M PERITHALLIUM 12 3 4 5 z u < _i UJ 15" 14 13 12 11- 10 8- 4- 3 . 11- 10 ^ -r-H. reinboldii (7 plants) o H. breviclavium (17 plants) A H . laeve (14 plants) □ H. megacystum (5 plants) FIGURE 15.—Mean epithallial and perithallial cell dimensions in Hydrolithon reinboldii, H. breviclavium, H. laeve, new species, and H. megacystum, new species. NUMBER 15 29 Depth IT) 30 40 50 60 H. reinboldii 70 80 90 B Depth m 10 20 30 40 50 60 70 80 90 5? « 3 20 > c 'Z D -S 1 *-'. >« v i - I > < * » . -■ " r £ * - K FIGURE 18.—Hydrolithon laeve, new species: A, habit of type specimen, X V2; B, section through tetrasporic plant, note single-layered hypothallium, tetrasporangia restricted to periphery of the conceptacle, X 200; c, section through thallus, note epithallium with outer tangential wall thickening (arrow), large single heterocyst in the perithallium, and the "large-celled" meristem, X 500; D, surface of type specimen, note tetrasporangial conceptacles, X 5. (Specimen nos.: A, D, 71-82-27; B, 71-82-24; c, 71-50-123.) 32 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES brunneas; conceptacula perspicua, elevata, sparsa magnaque praebentes (Figura 18D). Epithallium ex unico strato cellularum rotundatarum con- stans (Figura 18c), membrana tangentialis exte- rior incrassata, (3-7 /xm long., 6-12 /xm diam.). Meristema intercalare paululum elongatum; cel- lulis 6-12 /xm long, atque 6-10 /xm diam. (Figura 18c). Perithallium pluristratosum, irregulare, fu- sionibus praesentibus; cellulis 4-17 /xm long, atque 4-14 /xm diam., profundiores in tela ma- iores factae (Figura 15). Heterocystae singulae (Figura 18c) (10-25 /xm long., 5-20 /xm diam.) Hypothallium ex unico strato constans (Figura 18B), cellulis 9-25 /xm long, atque 5-18 /xm diam. Conceptacula tetrasporangialia uniporata, ele- vata (100-120 /xm) aggregata (Figura 18B,D) (600-1100 /xm O.D., 220-480 /xm I.D, 90-180 /xm alt.); tetrasporangia ad periferiam pavimenti con- ceptaculi restricta, columella adest (Figura 18B), 50-120 /xm long, 30-70 /xm diam. Conceptacula procarpica uniporata, elevata (80-120 /xm) 500- 650 /xm O.D, 200-300 /xm I.D, 60-120 /xm alt.; omnis cellula sustinens unicum ramum carpogon- ialem habens. Crusts often extensive, thin (<200 /xm) (Figure 18A), surface very finely rugulose (Figure 18D), pink to brownish; scattered large, raised conspic- uous conceptacles (Figure 18D). Epithallium a single layer of rounded cells (Figure 18c), wall thickening on outer tangential wall; cells 3-7 /xm long, 6-12/an diam. Intercalary meristem slightly elongate; cells 6-12 /xm long and 6-10 /xm diam. (Figure 18c). Perithallium multilayered, irregu- lar, fusions present; cells 4-17 /xm long and 4-14 /xm diam. becoming larger deeper in the tissue (Figure 15). Heterocysts single (Figure 18c) (10- 25 /xm long, 5-20 /xm diam.). Hypothallium a single layer (Figure 18B), cells 9-25 /xm long and 5-18 /xm diam. Tetrasporangial conceptacles un- ipored raised (100-120 /xm), grouped (Figure 18B, D) (600-1100 /xm O.D, 220-480 /xm I.D, 90-180 /xm high); tetrasporangia restricted to the periph- ery of the conceptacle floor, columella present (Figure 18B), 50-120 /xm long, 30-70 /xm diam. Procarpic conceptacles uniporate raised (80-120 /xm), 500-650 /xm O.D, 200-300 /xm I.D, 60-120 /xm high; 1 carpogonial branch per support cell. No cystocarpic or male material seen. TYPE-LOCALITY.—Sand Island, Midway Atoll, Hawaii (28°13'N, 177°26'W). Outside reef, 27 m depth. HOLOTYPE.—D. Child, 71-82-27, August 1971 (USNC), Figure 18A,B,D. PARATYPES.—Hawaii: Honaunau, March 1971, 71-55-67b; Kawaihae, March 1971, 71-57-75. Midway: South Island, August 1971, 71-82-24, 71- 82-26, 71-82-27. Nihoa: west reef, August 1971, 71-75-3. Oahu: Honauma, March 1971, 71-53-9; Waikiki, March 1971, 71-50-123; 71-50-132. DISTRIBUTION.—Throughout Hawaiian Archi- pelago. REMARKS.—The specific epithet laeve refers to its thin crustose form. The outer tangential wall thickening of the epithallial cells of Hydrolithon laeve is thinner than that of H. breviclavium. Hydrolithon laeve occurs as thin crusts covering the surfaces of coral and other carbonate rubble fragments. It is common from 25-50 m (Figure 16c) but may occur at any depth. We do not know of a Caribbean "pair species" (see discus- sion under Hydrolithon reinboldii). Hydrolithon megacystum, new species FIGURES 15, 16D, 19 DESCRIPTION.—Crustae rugulosae (Figura 19c) nodulosae (Figura 19), usque aliquot mm crass, rosaceae; conceptacula lata elevata super super- ficiem sparsa, interdum dense crebra (Figura 19c). Epithallium unistratosum; cellulis 3-6 /xm long, atque 6-10 /xm diam. Meristema intercalare non manifeste elongatum, intense tinctum potens (Figura 19B); cellulae 5-15 /xm long, atque 5-10 /xm diam. Perithallium pluristratosum, fusiones frequentes (Figura 19B), cellulis satis brevibus prope superficiem, profundioribus in tela, autem, progredienter longioribus factis (Figura 15) (4-16 /xm long, 4-15 /xm diam.). Heterocystae singulae (10-27 /xm long, 7-15 /im diam.). Hypothallium unistratosum (Figura 19B); cellulis 11-29 /xm NUMBER 15 33 F^H .^■■L* iH ■I^^HIfe.* ^^B I^H W AJ£< \ B^^^Ssi K«^^^ ^P^^^^^CPPvHb^i^HMfllBbL I ■§' v « Jr\* ■ | f< r* m • ■ . L_<^ f >v *^ .^fl B < ! -"A ^ .^^fl ■FIGURE 19.—Hydrolithon megacystum, new species: A, habit of type specimen, X 2; B, section through vegetative thallus, note single-layered hypothallium and heavily staining intercalary meristem, X 50; c, surface of type specimen showing conceptacles, X 10. (Specimen nos.: A, c, 71-50-100; B, 71-82-3.) 34 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES long, atque 6-15 /xm diam. Conceptacula tetra- sporangialia uniporata, lata, elevata (150-200 /xm), 400-800 /xm O.D, cavitates 300-440 /im I.D. atque 100-240 /xm alt, paraphyses steriles adsunt; tetrasporangia trans totum pavimentum conceptaculi disposita, 75-170 /xm long, 30-100 /xm alt. Crusts rugulose (Figure 19c), nodular (Figure 19A) becoming several mm thick, pink; broad raised conceptacles scattered over surface, occa- sionally densely concentrated (Figure 19c). Epi- thallium a single layer; cells 3-6 /xm long and 6- 10 /xm diam. Intercalary meristem not markedly elongate, stains heavily (Figure 19B); cells 5-15 /xm long and 5-10 /xm diam. Perithallium multi- layered, fusions common (Figure 19B), cells quite short near surface but becoming progressively elongate with depth in the tissue (Figure 15) (4- 16 /xm long, 4-15 /xm diam.). Heterocysts single (10-27 /xm long, 7-15 /xm diam.). Hypothallium a single layer of cells (Figure 19B); cells 11-29 /xm long and 6-15 /xm diam. Tetrasporangial concep- tacles uniporate, broad, raised (150-200 /xm), 400-800 /xm O.D, cavities 300-440 /xm I.D. and 100-240 /xm high, sterile paraphyses present; tetrasporangia across entire floor of conceptacle, 75-170 /xm long, 30-100 /xm high. No sexual material seen. TYPE-LOCALITY.—Waikiki, Oahu, Hawaii (2H6'N, 157°50'W), leeward. HOLOTYPE.—D. Child, 71-50-100, March 1971 (USNC), Figure 19A. PARATYPES.—Hawaii: Hilo Bay, March 1971, 71-58-68. Maui: south-central coast, August 1971, 71-66-14. Midway: South Island, August 1971, 71- 82-3. Oahu: Waikiki, March 1971, 71-50-100. DISTRIBUTION.—Throughout Hawaiian Archi- pelago. REMARKS.—The specific epithet megacystum re- fers to the unusually large reproductive struc- tures. Hydrolithon breviclavium, H. megacystum, and H. laeve range widely in the mid-depth zones (Figure 16B,C,D), although H. breviclavium is twice as com- mon as the other two species. Hydrolithon laeve does not contribute to rhodolith formation, though H. megacystum may occasionally occur in this form. These three species are not known in the Indo- Pacific outside of Hawaii. This is probably not indicative of actual occurrence, since corallines have not been widely collected and identified from the subtidal beyond Hawaii; the Siboga ex- pedition (Weber, 1902) dredging was generally in deeper water. Some reports of H. reinboldii may actually belong to our new species. All of the four Hydrolithon species have elongate intercalary meristems and considerable addi- tional cell elongation with burial in the perithal- lium (Figure 15). Perithallial cell elongation is quite rapid in H. reinboldii but more gradual in the other three species which occupy roughly the same depth range and are probably closely re- lated; it would be of considerable interest to examine the microhabitat preferences of these species. Lithoporella (Foslie) Foslie, 1909 Lithoporella melobesioides (Foslie) Foslie FIGURE 20 Lithoporella melobesioides (Foslie) Foslie, 1909:59.—Lemoine, 1963.—Masaki, 1968.—Papenfuss, 1968.—Womersley and Bailey, 1970.—Gordon et al., 1976. Maslophora melobesioides Foslie, 1903b: 24; 1908a,b.—Weber- van Bosse and Foslie, 1904. DESCRIPTION.—Thin, usually leafy, pink to ma- roon crusts with scattered raised conceptacles (Figure 20A,B,C), growth fanlike and easily dis- cernible under the dissecting microscope (Figure 20B). Epithallium a single cell layer; cells rounded to triangular (Figure 20D), 3-5 /xm long, 7-8 /xm diam. Perithallium present only around concep- tacles, lateral fusions common, no secondary pit connections (Figure 20D); cells 4-14 /xm long and 7-11 /tm diam. Hypothallium single layered, fu- sions absent; cells 30-40 /xm long and 10-15 /xm diam. Tetrasporangial conceptacles unipored, highly raised, 600-1200 /xm O.D. No sexual ma- terial seen. NUMBER 15 35 TYPE-LOCALITY.—Maldive Islands, Nilandu Atoll, Indian Ocean. FIGURE 20.—Lithoporella melobesioides: A, surface of thallus showing tetrasporangial conceptacles and "fan-like" over- growth, X 2; B, surface of thallus showing large-celled over- growth patterns, X 10; c, habit, X 1; D, section through vegetative thallus, X 10. (Specimen nos.: A-C, 71-59; D, 71- 81-16.) HOLOTYPE.—Gardiner, 20 April 1900, 36 fathoms, in herbarium of M. Foslie, (TRH). Iso- type: USNC. DISTRIBUTION.—Borneo, Cape Verde Island, Guam, Japan, Java, Philippines, Red Sea, Sa- moa, Solomon Islands, Timor. SPECIMENS STUDIED.—Molokai: southwest coast, August 1971, 71-73-8, Oahu: Hilo, March 1971, 71-59; Honauma, March 1971, 71-53-6; Ka- neohe, March 1971, 71-81-16. REMARKS.—We found nine specimens of Lith- oporella; all were referred to L. melobesioides (Foslie) Foslie. Lithoporella is found from the intertidal to 85 m depth and from Midway Atoll to the island of Hawaii. Only three specimens were microtome- sectioned; therefore, the preceding description is minimal. Considering the wide depth range of Lithoporella it is possible that we mixed several species. A more detailed study is therefore desir- able. LITHOPHYLLOIDEAE Setchell, 1943 Tenarea Bory, 1832 Only one species is recognized in the Hawaiian flora. Tenarea tessellatum (Lemoine) Littler FIGURES 21, 22 Tenarea tessellatum (Lemoine) Littler, 197la:355. Lithophyllum (Dermatolithon) tessellatum Lemoine, 1929:68.— Taylor, 1945. Goniolithon tessellatum (Lemoine) Setchell & Mason 1943:89.— Dawson, 1960b. DESCRIPTION.—Glossy, thin to several mm thick, bright pink to reddish crusts showing a very distinctive concentric or spiral pattern of overlapping tissue layers with thin white growth margins (Figure 21A,C), "terraces" varying from 200-2000 /xm in width, scattered broad but slightly raised conceptacles that leave circular depressions or are covered by the overlapping growth (Figure 21A,D). Thallus 2 cell layers; up- 36 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 21.—Tenarea tessellatum: A, surface of thallus showing spiral "terraces'' and tetrasporangial conceptacles, X 10; B, section through thallus showing tetrasporangial conceptacles per cell layer (epithallium) triangular or rounded, cells 2-8 /xm long and 5-13 /xm diam.; lower cell layer (hypothallium), cells 30-50 /xm long and 7- 15 /xm diam. Tetrasporangial conceptacles uni- pored, initially raised (90-110 /xm) but quickly buried by overlapping growth (Figure 21B,D), conceptacles 500-700 /xm O.D, 210-430 /im I.D, and 70-180/xm high (Figure 21B); tetrasporangia restricted to the periphery of the conceptacle, columella present, 50-100 /xm long and 30-50 /xm diam. One female plant seen, cystocarpic concep- tacles, unipored, raised 400 /xm O.D, 210-250 /xm I.D, 70-80 /xm high, not buried in thallus; carposporangia restricted to periphery of large discoid fusion cell; fusion cell, 125 /xm diam.; carpospores rounded, 30-35 /xm diam. No male conceptacles seen. TYPE-LOCALITY.—Post Office Bay, Floreana (Charles) Island, Galapagos Islands. HOLOTYPE.—Crossland, s.n, August 1924 (PC). DISTRIBUTION.—Galapagos, Hawaii, Panama. SPECIMENS STUDIED.—Hawaii: Honaunau, March 1971, 71-55-26. Midway: Lagoon, March 1971, 71-62-40; South Island, August 1971, 71- 82-22, 71-82-59. Oahu: Honauma, March 1971, 71-53-18; Waikiki, March 1971, 71-50-44, 71-50- 89. REMARKS.—The specific epithet tessellatum de- scribes a mozaic pattern rather than the spiral pattern seen on the thallus surface of T. tessellatum. It is unfortunate Lemoine (1929:68) chose this name, since confusion with the "tessellate sur- face" of Hydrolithon reinboldii may occur. Tenarea tessellatum was fairly common in our collections from the intertidal to 40 m depth (Figure 22). The Caribbean alga Tenarea proto- typum (Foslie) Adey is a "pair species" of T tessel- latum. It is common in the Caribbean (Adey, 1979) and appears to differ from T. tessellatum by having much narrower "terraces" (rarely more than about 1 mm wide). (arrowed), X 50; c, habit, X 1; D, section through thallus showing overgrowth of conceptacles, X 50. (Specimen nos.: A, c, 71-62-40; B, 71-82-59; D, 71-50-89; micrographs reduced to 90%.) NUMBER 15 37 0> u c o c 10 20 30 Depth m 40 50 70 T. tessellatum 80 90 FIGURE 22.—Depth distribution of Tenarea tessellatum. Lithophyllum Philippi, 1837 Key to the Species 1. Plants branched 2 Plants crustose 3 2. Branches long, terete to flattened L. kotschyanum Branches short, terete to mushroom shaped L. pallescens 3. Crust tessellate L. insipidum, new species Crust smooth 4 4. Crust smooth glossy, pink, medium-sized conceptacles (>300 /xm O.D.) . . L. ganeopsis, new species Crust smooth dull, pink, very small conceptacles (<200 um O.D.) L. punctatum Lithophyllum kotschyanum Unger FIGURES 23-25 Lithophyllum kotschyanum Unger, 1858:22.—Foslie, 1909; 1929.—Pilger, 1919.—Gordon et al, 1976. Lithothamnium kotschyanum Unger, 1858:22. [See also synonyms listed by Foslie, 1909, 1929.] DESCRIPTION.—Massive branching plants, sometimes with an extensive basal crust (Figure 24B) but most commonly densely branched with little crust; wide range of branch morphology: short simple knobs, long thin finger-like projec- tions, vertical plates or large fused clubs (Figure 24A,B,C,D); glossy; red-purple, frequently becom- ing white and chalky on enlarged branch ends; conceptacles slightly raised with no definite bor- ders, densely grouped when present (Figure 24D) but often completely absent. Epithallium single cell layer, often absent; cells 2-4 /im long and 6- 12 /am diam. Intercalary meristem generally elon- gate, forming a distinct layer; cells 5-25 /xm long and 7-11 /xm diam. Perithallium multilayered; cells often arranged in horizontal layers of uni- form size and alignment (Figure 24E,F) elongating greatly with branching or change in direction of growth, secondary pit connections common (Fig- ure 24F), 4-24 /xm long and 4-10 /xm diam. (Figure 23). Hypothallium single layer, lower per- ithallium often curved upwards into false coaxial hypothallium; cells isodiametric to slanting pali- sade, size varying greatly, 6-21 /xm long and 7- 25 /xm diam. Tetrasporangial conceptacles uni- pored, usually level with surface or slightly raised (occasionally raised to 60 /xm), 350-450 /xm O.D, 250-370 /xm I.D, 80-210 /xm high, raised central columella present (Figure 24E); tetrasporangia 38 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 0 Z u PERITHALLIUM 3 4 5 □ L. kotschyanum (13 plants) I L. pallescens (12 plants) AL. ganeopsis (4 plants) ■ L. insipidum (5 plants) oL. punctatum (4 plants) D A ep M PERITHALLIUM 3 4 5 6 < 7 o A u FIGURE 23.—Mean epithallial and perithallial cell dimensions of Lithophyllum kotschyanum, L. pallescens, L. ganeopsis, new species, L. insipidum, new species, and L. punctatum. NUMBER 15 39 FIGURE 24.—Lithophyllum kotschyanum: A, B, habit, X 1; c, close-up of branches, X 10: D, surface with conceptacles, X 10; E, tetrasporangial conceptacle showing raised central columella, X 130; F, perithallium with secondary pit connections, X 500. (Specimen nos.: A, C, 71-57-12; B, 71-57-2; D, 71-59-10; E, 71-57-1; F, 71-59-8; micrographs reduced to 80%.) 40 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES L . kotschyan um 10 20 30 40 50 °0 Depth m FIGURE 25.—Depth distribution of Lithophyllum kotschyanum. and bisporangia restricted to the conceptacle pe- riphery, 40-100 /xm long, 20-60 /xm diam. No sexual material was found. TYPE-LOCALITY.—Gulf of Bahrain, Persian Gulf. HOLOTYPE.—Kotschy, in herbarium of M. Fos- lie (TRH). Isotype: USNC. DISTRIBUTION.—Guam, Persian Gulf, Red Sea. SPECIMENS STUDIED.—Hawaii: Hilo Bay, March 1971, 71-58-72, 71-59-8, 71-59-10, 71-59- 25; Honaunau, March 1971, 71-56-1; Kawaihae, March 1971, 71-54-1, 71-57-2, 71-57-12. Midway: lagoon, March 1971, 71-62-21; South Island, Au- gust 1971, 71-82-60. Oahu: Kaneohe, August 1971, 71-82-60. REMARKS.—Lithophyllum kotschyanum is similar in habit, surface detail, and morphology anatomy to the Caribbean alga Lithophyllum congestum (see Steneck and Adey, 1976), yet there are some differences in the ecology of these plants. In the Caribbean, L. congestum is an algal ridge builder, the peak of its abundance occurring at mean low water. At depths of several meters, the branches are much reduced, and the few plants found at 5-10 m are generally without branches. On the other hand, L. kotschyanum occurs to a considerable depth in Hawaii (Figure 25) and, at least in our data, was much less abundant at low water levels, although Littler (pers. comm.) indicates L. kot- schyanum replaces P. gardineri to some extent on the algal ridges, especially along channel and groove sides. Lithophyllum pallescens (Foslie) Foslie FIGURES 23, 26, 27 Lithophyllum pallescens (Foslie) Foslie, 1900c:20; 1901a; 1909. —Dawson, 1944; 1954a; 1960b; 1961b.—Heydrich, 1901a.—Lemoine, 1911. Lithothamnwn pallescens Foslie, 1895:4.—Heydrich, 1897b,c. Goniolithon pallescens (Foslie) Foslie, 1898:9. Lithophyllum okamurai Foslie, 1900b:4; 1904; 1906.—Weber- van Bosse and Foslie, 1904.—Howe, 1918b.—Okamura, 1936.—Masaki and Tokida, 1963.—Lemoine, 1965; 1966. —Masaki, 1968.—Papenfuss, 1968.—Womersley and Bai- ley, 1970. Lithophyllum californiense Heydrich, 1901a. Lithophyllum cephaloides Heydrich, 1901 b: 271. DESCRIPTION.—Plants initially crustose but quickly developing densely spaced small short simple club-shaped branches (Figure 27A,B); ini- tial growth usually occurs on attached substrate, but a developed head often becomes detached and occasionally forms large rhodoliths reaching to 8 cm diam. (Figure 27A); surface fairly smooth with occasional patches of small light-colored conceptacles, slightly raised or sometimes slightly depressed. Cover cells present. Epithallium usu- ally a single cell layer (rarely 2 cell layers); cells 2-7 /xm long and 7-12 /xm diam. Intercalary meristem a distinct layer; cells usually quite square but occasionally elongating individually, 5-15 /xm long and 6-11 /xm diam. Perithallium multilayered, regular with strong horizontal lay- ering secondary pit connections common between cells; cells 5-18 /xm long and 4-15 /xm diam. (Figure 23). Hypothallium single layer when pres- ent (Figure 27c); cells 6-24 /xm long and 6-10 /xm diam. Tetrasporangial conceptacles unipored, usually level with surface (may be sunken (20 /xm) or raised (5 /xm)) 140-310 /xm I.D, 50-160 /xm high; tetrasporangia restricted to the periph- ery of conceptacle (columella present), 30-45 /xm long and 15-20 /xm wide. Male conceptacle uni- porate, raised (70-100 /xm) 190-480 /xm I.D, 30- 80 /xm high; spermatangial mother cells restricted to the conceptacle floor, 2 spermatial extensions NUMBER 15 41 10 20 30 40 50 60 Depth m 70 80 90 FIGURE 26.—Depth distribution of Lithophyllum pallescens. FIGURE 27.—Lithophyllum pallescens: A, habit of typical rhodolith, X 1; B, surface showing branches and conceptacles (arrowhead), X 10; c, section through vegetative thallus, X 100. (Specimen nos.: A-C, 71-50-96; micrographs reduced to 81%.) per mother cell; spermatia discoid, wound into mucus strands in conceptacle cavity, 1-2 /xm diam. TYPE-LOCALITY.—Sublittoral bank along the west shore of Isla Espiritu Santo, near La Paz, Gulf of California, Mexico. HOLOTYPE.—Diguet Hariot #5, in the herbar- ium of M. Foslie (TRH). Isotypes: PC, UC. DISTRIBUTION.—Australia; Baja, California; Borneo, Gulf of California, Indian Ocean, Japan, New Guinea, Panama, Phillipines, Red Sea, Sol- omons, Vietnam. SPECIMENS STUDIED.—Hawaii: Hilo, March 1971, 71-58-53; Honaunau, March 1971, 71-55- 20. Midway: lagoon, March 1971, 71-62-33. Molo- kai: southwest, August 1971, 71-73-9. Oahu: Ka- neohe, March 1971, 71-54-2, August 1971, 71-81- 34; Waikiki, March 1971, 71-50-52, 71-50-96. REMARKS.—We have been unable to differen- tiate between Lithophyllum okamurai Foslie (1900), the designation usually used for Indo-Pacific spec- imens of this plant (see Womersley and Bailey, 1970), and L. pallescens. Since L. pallescens has priority by five years, we use that name here. The outside diameter of tetrasporangial con- ceptacles has not been given since such a mea- surement has little taxonomic value in Lithophyl- lum pallescens. The development of the male con- 42 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ceptacle is similar to that described for Parago- molithon conicum. Lithophyllum pallescens is a primary rhodolith- builder in depths of 15-80 m (the low value at 45 m in Figure 26 is probably more a reflection of incomplete collecting at that depth rather than any real minimum value). Like Hydrolithon brevi- clavium, L. pallescens is generally replaced by me- lobesioid species on the abundant small nodules occurring below depths of 80 m. Littler (1973b) describes in detail the occurrence of rhodolith communities off the island of Oahu at depths of 8-28 m; however, that author did not differen- tiate between the sympatric L. pallescens and H. breviclavium occurring in this study site. A morphological and ecological "pair species" of this plant has not been found in the Caribbean. Lithophyllum ganeopsis, new species FIGURES 23, 28 DESCRIPTION.—Crustae bene evolutae usque aliquot mm crassitudine, superficie maxime ni- tida (Figura 28A,B); conceptacula non perspicua, paululum elevata fortuite sparsa (Figura 28c). Epithallium unistratosum, raro duostratosum; 2- 5 /xm long, atque 6-9 /xm diam. Meristema inter- calare relative breve, cellulis 6-11 /xm long, et 5- 8 /xm diam. Perithallium pluristratosum, saepe regulariter stratosum (Figura 28D), foveo-colliga- tiones secondariae abundates; cellulis 4-16 /xm long, et 4-12 /xm diam. (Figura 23). Hypothal- lium ex unico strato cellularum plerumque iso- diametricarum (7-19 /xm long, 7-14 /xm diam.) constans. Conceptacula tetrasporangialia unipor- ata, paululum elevata (Figura 28c,E), 250-500 /xm O.D, 140-300 /xm I.D. atque 60-120 /xm alt, in perithallio obruta; tetrasporangia per pavi- mentum conceptaculi disposita, 40-65 /xm long, 20-40 /xm diam. Conceptacula procarpica non visa. Conceptacula cystocarpica uniporata, ele- vata (100 /xm), 230-280 /xm I.D, 70-80 /xm alt, carposporangia ad periferiam conceptaculi res- tricta; cellula-coalescens discoidea, 150-200 /xm diam.; carposporae 50 /xm long.; 25 /xm diam. Conceptacula mascula uniporata, elevata (60 /xm), 230-310 /xm I.D, 40-60 /xm alt.; cellulae- matricales 2 extensiones spermatiales habente; spermatia discoidea ad ellipsoidea, 3-4 /xm long, 2-3 /xm diam. intra fila mucosa reperta (Figura 28F). Well-developed crusts reaching a thickness of several mm, surface highly glossy (Figure 28A,B); indistinct, slightly raised, randomly scattered con- ceptacles (Figure 28c). Epithallium single (rarely 2) cell layers; cells 2-5 /xm long and 6-9 /xm diam. Intercalary meristem relatively short; cells 6-11 /tm long and 5-8 /xm diam. Perithallium multi- layered, often regularly layered (Figure 28D), abundant secondary pit connections; cells 4-16 /xm long and 4-12 diam. (Figure 23). Hypothal- lium a single layer of generally isodiametric cells (7-19 /xm long, 7-14 /xm diam.). Tetrasporangial conceptacles unipored, raised slightly (Figure 28c, E), 250-500 /xm O.D, 140-300 /xm I.D. and 60- 120 /xm high, buried in perithallium; tetraspo- rangia across floor of conceptacle, 40-65 /xm long, 20-40 /xm diam. No procarpic conceptacles seen. Cystocarpic conceptacles uniporate, raised (100 /xm), 230-280 /xm I.D, 70-80 /xm high; carpo- sporangia restricted to periphery of conceptacle; fusion cell discoid, 150-200 /xm diam.; carpo- spores 50 /xm long, 25 /xm diam. Male concepta- cles uniporate raised (60 /im), 230-310 /xm I.D, 40-60 /xm high; spermatangial mother cells re- stricted to the conceptacle floor, 2 spermatial extensions per mother cell; spermatia discoid to ellipsoidal, 3-4 /xm long, 2-3 /xm diam, within mucus strands (Figure 28F). TYPE-LOCALITY.—Honauma Bay, Oahu, Ha- waii (21°15'N, 157°20'W); windward, 8 m. HOLOTYPE.—D. Child 71-53-23, March 1971 (USNC), Figure 28A,B. FIGURE 28.—Lithophyllum ganeopsis, new species: A, surface of type specimen, X 10; B, habit of type specimen, X 1; c, surface view with conceptacles (arrowed), X 10; D, section through vegetative thallus, note single-layered hypothallium (arrowed), X 150; E, tetrasporangial conceptacle, X 150; F, male conceptacle, note spermatia in strands along roof of conceptacle, X 150. (Specimen nos.: A, B, 71-53-23; c, D, 71- 56-5; E, 71-81-6; F, 71-81-29.) NUMBER 15 43 ;i<'d»»-^-".> • <-■ ----- - j*^.■•■- '•£•■■■ ■ -■> ■*■ *| ,A'"^*^ Ik *&W £s 44 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES PARATYPES.—Hawaii: Honaunau, March 1971, 71-56-5; Kaneohe, August 1971, 71-81-6, 71-81- 29. Necker: August 1971, 71-76-1. REMARKS.—The specific epithet ganeopsis refers to the glossy surface common to this species. Tetrasporangial and cystocarpic conceptacles are buried in the perithallium with both tetra- sporangial and carposporangial remains within the conceptacle cavity. Overgrowth occurs from the roof of the conceptacle. Lithophyllum ganeopsis is a very distinctive but infrequent plant in our collections. Its maximum abundance for any one zone was 2% of the col- lected specimens. Although most of the plants found occurred from the intertidal to 10 m depth, a few were taken at depths of 30 m. We do not know of a Caribbean "pair species," though Lith- ophyllum nitorum that W. and P. Adey described for the temperate eastern Atlantic (Adey and Adey, 1973) is morphologically similar. Lithophyllum insipidum, new species FIGURES 23, 29 DESCRIPTION.—Crustae non amplae, satis cras- sae, autem, factae (usque ad 1 cm) rosaceae ad lavandulas, saepe maculas albas non perspicuas praebentes, superficie tessellata distincte sed ir- regulariter plerumque facta (Figura 29B,C,F). Conceptacula plerumque abundantia, parva, paululum elevata (Figura 29F). Epithallium ex unico strato cellularum (3-6 /xm long, 6-10 /xm diam.) constans (Figura 29D). Meristema inter- calare perspicuum, elongatum, cellulis 11-20 long, atque 6-9 /xm diam. (Figura 29D). Perithal- lium pluristratosum, foveo-colligationes secon- darias abundantes praebens (Figura 29D), bene stratosum, cellulis 4-22 /xm long, atque 5-10 /tm diam. (Figura 23). Hypothallium unistratosum (Figura 29A); cellulis 12-16 /xm long, atque 12- 19 /xm diam. Conceptacula tetrasporangialia un- iporata, paululum elevata (30-70 /xm), 450-750 /xm O.D, 170-230 /xm I.D, 80-140 /xm alt. (Fig- ura 29A); tetrasporangia ad periferiam concepta- culi restricta, columella adest, 40-100 /xm long, 20-50 /xm diam. Crusts not extensive but becoming quite thick (to 1 cm), pink to lavender, often with indistinct white patches, surface usually becoming dis- tinctly though irregularly tessellate (Figure 29B, C,F); conceptacles usually abundant, small, slightly raised (Figure 29F). Epithallium a single layer of cells (3-6 /xm long, 6-10 /xm diam.) (Figure 29D). Intercalary meristem distinct, elon- gate; cells 11-20 /xm long and 6-9 /xm diam. (Figure 29D). Perithallium, multilayered, with abundant secondary pit connections (Figure 29D), well layered; cells 4-22 /xm long and 5-10 /xm diam. (Figure 23). Hypothallium a single cell layer (Figure 29A); cells 12-16 /xm long and 12- 19 /xm diam. Tetrasporangial conceptacles uni- pored, slightly raised (30-70 /xm), 450-750 /xm O.D, 170-230 /xm I.D, 80-140 /xm high (Figure 29c); tetrasporangia restricted to the conceptacle periphery, columella present, 40-100 /xm long, 20-50 /xm diam. Sexual plants not found. TYPE-LOCALITY.—Waikiki, Oahu, Hawaii (21°10'N, 157°55,W), leeward, 3-5 m depth. HOLOTYPE.—D. Child, 71-50-17, March 1971 (USNC), Figure 29B. PARATYPES.—Hawaii: Hilo, March 1971, 71-58- 39; Honaunau, March 1971, 71-55-71. Midway: South Island, August 1971, 71-82-32. Oahu: Waianae, March 1971, 71-52-25; Waikiki, March 1971, 71-50-17. DISTRIBUTION.—Southern part of archipelago. REMARKS.—Specific epithet refers to the "dull" patchy appearance of this species in contrast to the typical glossy Lithophyllum surface. Although the anatomy of this plant is typically that of the genus Lithophyllum, and thus strikingly different from Hydrolithon, L. insipidium, with its lavender color and tessellate surface, may be con- fused with Hydrolithon reinboldii. Thicker plants with the color and texture of H. reinboldii that are lacking mammillons or with relatively small con- ceptacles should be investigated closely to deter- mine whether they are H. reinboldii or L. insipidum. Examination of a fractured vertical surface for the irregular, almost porous perithallium (due to extensive fusions) or H. reinboldii should easily NUMBER 15 4r) '{>:^;J,', " - ' A 9 >t'r* • FIGURE 29.—Lithophyllum insipidum, new species: A, section through thallus showing lower vegetative tissue, X 600; B, habit of type specimen, X 1; c, surface of type specimen showing tesselate pattern, X 10; D, section through thallus epithallium (arrowed), intercalary meristem (arrowhead), and perithallium, X 300; E, tetrasporangial thallus, note conceptacle and region of new growth, X 150; F, surface of type specimen with conceptacles (arrow), X 10. (Specimen nos.: A, 71-52-25; B, c, F, 71-50-17; D, 71-82-32; E, 71-58-39; micrographs reduced to 93%.) 46 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES -v ' • * ¥■ HVK 3* V *.ir+ r V;V- > ■- JV - - MfccVfy *•• ■ '■•■"" - --■ - • - * ■ ■ - I l! * ~,^^-- ■*'■" f?*&Jk* ^ " f&@$ A -4; * FIGURE 30.—Lithophyllum punctatum: A, habit, X 1; B, surface with intact conceptacles, X 10; c, surface with spent conceptacles, X 10; D, tetrasporangial conceptacle, X 250; E, section of vegetative tissue showing hypothallium, X 300. (Specimen nos.: A, 71-52-20; B, 71-54-8; c, 71- 52-30; D, 71-53-32; E, 71-50-7; micrographs reduced to 83%.) NUMBER 15 47 differentiate a crust from the even, fine-grained, and layered Lithophyllum pattern. Lithophyllum insipidum was more abundant in our collections than L. ganeopsis, but still is only occasional, reaching a maximum zonal abun- dance of 6% from 0 to 3 m. A few specimens were found from 15 to 35 m, but most occurred from low water to 10 m. Lithophyllum punctatum Foslie FIGURES 23, 30 Lithophyllum punctatum Foslie, 1906:22. DESCRIPTION.—Surface dull and appearing dry, pink-lavender color, crusts becoming fairly thick, to several mm (Figure 30A,B); conceptacles gen- erally abundant, level with surface or slightly raised, scattered randomly over the surface, leav- ing open pits after spore release (Figure 30B,C). Epithallium 1 to 3 cell layers; cells 2-6 /xm long and 6-12 /xm diam. Intercalary meristem of ran- domly elongating cells (5-9 jum long, 6-8 /xm diam.). Perithallium multilayered, secondary pit connections present, scattered large cells at sur- face (16-18 /xm long, 10-11 /xm diam.), adjacent cells varying in size and position, tissue appearing as a system of loose independent vertical strands; cells elongating greatly with depth in the tissue (3-13 /xm long, 4-10 /xm diam.) (Figure 23). Hypothallium a single cell layer; cells isodia- metric to horizontally elongate, 7-16 /xm long and 6-10/xm diam. (Figure 30E). Tetrasporangial conceptacles unipored, raised 40-50 /xm above the surrounding crust, 160-170 /xm I.D, 60-70 /xm high (Figure 30D); tetrasporangia restricted to periphery of conceptacle, columella present, 40-80 /xm long and 20-65 /xm diam. No sexual material collected. TYPE-LOCALITY.—Tricomalie, Ceylon. HOLOTYPE.—Svedelius, 17 April 1903, in her- barium of M. Foslie (TRH). Isotype: USNC. DISTRIBUTION.—Ceylon, the island of Oahu. SPECIMENS STUDIED.—Oahu: Honauma, March 1971, 71-53-32; Kaneohe, March 1971, 71-54-8; Waianae, March 1971, 71-52-20, 71-52-30; Wai- kiki, March 1971, 71-50-7. REMARKS.—We found only 10 specimens of this species, all from less than 10 m depth. Recently, a similar plant as yet undescribed (Adey, unpub- lished data) was found in the Caribbean, on the shallow algal pavements of Martinique. MELOBESIOIDEAE (J.E. Areschoug) Yendo, 1902 Archaeolithothamnium Rothpletz, 1891 The authors realize that the use of Archaeolith- othamnium over Sporolithon Heydrich (1897b) has been questioned (Papenfuss, 1968). In this paper, we shall use Archaeolithothamnium. There has been considerable concern in recent years over the taxonomic position of this genus. Adey (1970; Adey and Johansen, 1972) placed Archaeolithothamnium in the subfamily Melobesioi- deae, closely related to Lithothamnium. Cabioch (1972), because of the apparent presence of sec- ondary pits, has placed the genus in a separate subfamily, the Sporolithoideae Setchell (1943: 134). Womersley (pers. comm.), on the other hand, has suggested that, because of the occa- sional presence of cruciate tetrasporangia, per- haps the genus should be placed in a separate new family. Our position remains unchanged in this regard. Even though transmission electron microscope studies of Australian members of Archaeolitho- thamnium (as Sporolithon, Townsend, unpublished data) have shown the presence of secondary pit connections as well as lateral cell fusions, the cover cells or epithallial cells and the meristem cells of both Lithothamnium and Archaeolithotham- nium are quite similar (Adey and Macintyre, 1973). This type of cover cell is unique in the Corallinaceae and other Melobesioideae. In ad- dition, the sporangial caps that define the subfamily Melobesioideae are present in Archaeo- lithothamnium, though they are generally smaller than in other genera. While it is true that the sporangia sometimes develop cruciately, this is unusual, the zonate development being more typ- 48 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ical in Hawaiian species of Archaeolithothamnium. It is our opinion that the nature of the epithallium and the presence of sporangial caps far outweigh these irregularities in indicating the relationship of the genus. We feel that Lithothamnium is very closely related to Archaeolithothamnium, probably having evolved from it during the mid to late Mesozoic (Adey and Macintyre, 1973). Key to the Species of Archaeolithothamnium Glossy, red-brown crusts, sometimes mammillate or with short branches, with generally extensive sori (> 5 mm diam.) and narrow sporangial pores (not visible with a hand lens or barely so) A. erythraeum Glossy, red to pink crusts, sometimes having small irregular branches, with small sori (< 5 mm diam.) bearing large sporangial pores (easily visible with a hand lens) A. episoredion, new species Archaeolithothamnium erythraeum (Rothpletz) Foslie FIGURES 31-33 Archaeolithothamnium erythraeum (Rothpletz) Foslie in Weber- van Bosse and Foslie, 1904:39.—Foslie, 1900c [nomen nudum]; 1907a; 1908a.—Littler, 1971b; 1973a.—Setchell, 1924; 1926. Lithothamnium erythraeum Rothpletz, 1893:5. Sporolithon erythraeum (Rothpletz) Kylin, 1956:205.—Papen- fuss, 1968.—Womersley and Bailey, 1970. DESCRIPTION.—Crusts thick, often growing nonadherent to substrate sometimes developing mammillons, surface smooth, glossy (Figure 31A) bright bluish red to brown. Cover cells "Lithothammum-type" (Figure 31C,D) forming a single layer over the epithallium; cells 2-7 /xm long and 5-10 /xm diam. Intercalary meristematic cells 5-8 /xm long and 5-11 /xm diam. Perithal- lium multilayered regular, lateral cell fusions and secondary pit connections present (Figure 31B); cells square, tending to elongate and enlarge with depth in tissue (Figure 32), 5-12 /xm long and 4- 15 /xm diam. Hypothallium multilayered, parallel to weakly coaxial, 30-110 /xm thick; cells 13-34 /xm long and 6-9 /xm diam. Tetrasporangial sori buried in tissue (Figure 31E), surface sori not seen in collections; tetrasporangia 60-90 /xm long and 25-40 /xm diam. TYPE-LOCALITY.—Red Sea. HOLOTYPE.—Unknown (Papenfuss, 1968; pos- sibly destroyed in Berlin bombings). DISTRIBUTION.—Borneo, Celebes, Hawaii, In- dian Ocean, New Guinea, Philippines, Red Sea, Samoa, Solomon Islands, Tahiti, Timor. SPECIMENS STUDIED.—Midway: lagoon, March 1971, 71-62-8; South Island, August 1971, 71-82- 63, 71-82-69. Oahu: Honauma, March 1971, 71- 53-24; Waikiki, March 1971, 71-50-61. REMARKS.—Archaeolithothamnium erythraeum is not an abundant plant in our collections. In contrast to the abundant deep-water Archaeolith- othamnium episoredion, new species, most of the A. erythraeum were taken between low water and 15 m (Figure 33). Doty (1974, as Sporolithon) indi- cated that an Archaeolithothamnium species is a major sublittoral shade element of Pacific algal ridges, but he did not recognize the numerous Neogoniolithon species that tend to dominate such environments in both the Caribbean and Hawaii. We suggest that Doty's Hydrolithon and Sporolithon "communities" may include, or even be domi- nated by, Neogoniolithon species. NUMBER 15 49 FIGURE 31.—Archaeolithothamnium erythraeum: A, habit of typical specimen, X 2; B, scanning electron micrograph of perithallium in area of secondary pit connection; X 2500; c, scanning electron micrograph of "Lithothammum-tyoe" cover cell, X 7500; D, light micrograph of "Lithothammum-iype'1 cover cell, X 2000; E, thallus with buried sorus (arrow), X 20. (Specimen nos.: A, 71-82-69; B, c, 71-53-24; D, E, 71-82-63; micrographs reduced to 95%.) 50 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ep 6<> M PERITHALLIUM 3 4 5 6 7 0 A. erythraeum (5 plants) oA. episoredion (6 plants) FIGURE 32.—Mean epithallial and perithallial cell dimensions oi Archaeolithothamnium erythraeum and A. episoredion, new species. A. erythraeum FIGURE 33.—Depth distribution of Archaeolithothamnium erythraeum. Archaeolithothamnium erythraeum is the "pair spe- cies" of the Caribbean alga Archaeolithothamnium dimotum, which appears very similar in habit and anatomy. Archaeolithothamnium dimotum occurs in shallow areas, favoring strongly shaded situations (in holes around the bases of coral heads or under Sargassum) on reef flats. We have not seen a mammillate form of A. dimotum; however, the relationship between the crustose and mammil- late forms of A. erythraeum is not entirely clear at present. These plants are relatively minor ele- ments in both our Caribbean and Hawaiian col- lections and are in need of considerable additional study. NUMBER 15 51 Archaeolithothamnium episoredion, new species FIGURES 32, 34, 35 DESCRIPTION.—Crustae leves nitidae (Figura 35A) dilute rosaceae ad laete rubiginosus, amplae factae; sori elevati, ovales usque ad aliquot mm dimensione longissima, poris obturaculisve spor- angialibus magnis, facile visibilibus (oculo ar- mato) praediti (Figura 35B,C). Cellulae obte- gentes typi Lithothamnu (Figura 35E), 2-4/xm long, atque 2-7 /xm diam. cellulae meristematis inter- calaris 4-8 /tm long, atque 6-10 /xm diam. Peri- thallium regulare, pluristratosum, cellulis quad- ratis, progredienter longioribus profundioribus in tela (Figura 32), 4-18 /xm long, atque 5-13 /xm diam. Hypothallium pluristratosum, parallelum, 20-60 /xm crass.; cellulis 4-25 /xm long, et 5-10 /xm diam. Sori superficiales, in perithallio inter- dum obruti (Figura 35D), post liberationem spor- angialem flaviformes facti (Figura 35B,C); tetra- sporangia zonata, 70-200 /xm long, atque 35-100 /xm diam. Smooth, glossy crusts (Figure 35A) light pink to bright brownish red, becoming extensive; raised oval sori reaching several mm in longest dimension with large, easily visible (with a hand lens) sporangial pores or plugs (Figure 35B,C). Cover cells "Lithothammum-X.ype'' (Figure 35E), 2- 4 /xm long and 2-7 /tm diam. Intercalary meristem cells 4-8 /xm long and 6-10 /un diam. Perithal- lium regular, multilayered; cells square, becom- ing increasingly elongate with depth (Figure 32), 4-18 /xm long and 5-13 /xm diam. Hypothallium multilayered, parallel, 20-60 /xm thick; cells 14- 25 /xm long and 5-10 /xm diam. Sori superficial, sometimes buried in perithallium (Figure 35D), upon sporangial release becoming "honeycomb"- like (Figure 35B,C); tetrasporangia zonata, 70-200 /xm long and 35-110 /xm diam. Sexual thalli were not seen. TYPE-LOCALITY.—St. Rogatien Bank, north- west, Hawaii (24°15'N, 167°0'W), 70-95 m. HOLOTYPE.—D. Child, 71-79-(35-47f), August 1971 (USNC), Figure 35A. PARATYPES.—Hawaii: Honaunau, March 1971, 71-55-18. Maui: south-central coast, August 1971, 71-67-9. Nihoa: west coast, August 1971, 71-75-4. Oahu: Waikiki, March 1971, 71-50-77. DISTRIBUTION.—South and central archipelago. REMARKS.—The specific epithet episoredion re- fers to the superficial nature of the sori. This species is abundant in deep water and is an important builder of rhodoliths (Figure 34). It is very similar to the Caribbean Archaeolithotham- nium episporum Howe (1918a), which was described from shallow water and we have consistently found in deep water elsewhere in the Caribbean (Adey, unpublished data). We have artificially kept the species separate, even though it has meant the perhaps unnecessary description of a new taxon. In the tropical East Pacific, two spe- cies with similarly small sori have been described (A. howei Lemoine; A. pacificum Dawson), but both of these generally have smaller cells and sporangia. 10 20 30 Depth 40 m 50 60 70 80 90 Relative Abundance % o o A.episoredion FIGURE 34. — Depth distribution oi Archaeolithothamnium episoredion, new species. 52 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 35.—Archaeolithothamnium episoredion, new species: A, habit of type specimen, X 2; B, c, surface of thallus showing sori, note large sporangial pores, X 10; D, section through thallus showing buried sori, X 20; E, section through thallus showing cover cells, X 1500. (Specimen nos.: A, 71-79-(35-47f); B, c, 71-67-9; D, E, 71-55-18.) We collected two small rhodolith specimens of Archaeolithothamnium from 70-90 m, one with an almost smooth surface, the other with abundant short branches (1-2 mm long) about 1 mm diam. Both of these plants had sori that were only 400- 600 /xm in outside diam, smaller than the con- ceptacles of many species of Lithothamnium (see below, L. pulchrum Weber-van Bosse & Foslie in Foslie); however, these small sori showed the basal "honeycomb" pattern indicating partial calcifi- cation between the individual sporangia charac- teristic of Archeolithothammum. In section, narrow fusions or apparent secondary pits are present, along with the larger cell fusions, and the spor- angial cap walls are very short. These specimens have been tentatively placed with the anatomi- cally similar A. episoredion, though it may be that they belong to a separate species. In this case, it is interesting to note that the difference between an Archaeolithothamnium sorus and a Lithothamnium conceptacle is not marked, even though this char- acteristic has been used for generic, subfamily, and even family differentiation in crustose coral- lines. NUMBER 15 53 Lithothamnium Philippi, 1837 Key to the Species Extensive leafy pink-brown crusts becoming nodular and eventually devel- oping coarse branches (branches 2-5 mm diam.); large conceptacles > 500 M™, O.D L. pulchrum Thin, bright-red crusts developing abundant, narrow often irregular branches, 0.5-2 mm diam.; conceptacles < 500 /xm, O.D L. australe Lithothamnium pulchrum Weber-van Bosse & Foslie FIGURES 36-38 Lithothamnium pulchrum Weber-van Bosse & Foslie in Foslie, 1902a:3.—Weber-van Bosse and Foslie, 1904. Mesophyllum pulchrum (Weber-van Bosse & Foslie in Foslie) Lemoine, 1928:252. DESCRIPTION.—Rather leafy, medium thin plants (200-500 /xm thick) with large raised mul- tipored conceptacles; developing numerous knobs and eventually large irregular branches 2-5 mm diam. (Figure 36A,B). Cover cells "Lithothammum- type," single cell layer; cells 1-4 /xm long and 3- 10 /xm diam. Intercalary meristem medium length; cells 3-6 /xm long and 5-10 /xm diam. Perithallium multilayered, abundant fusions, scattered staining bodies (phosphotungstic hae- matoxylin) (Figure 36c); cells 3-9 /xm long and 9-22 /xm diam. (Figure 37). Hypothallium mul- tilayered, noncoaxial, 20-65 /xm thick; cells 3-11 /xm long and 4-11 /xm diam. (Figure 36c). Tet- rasporangial conceptacles multipored, strongly raised (Figure 36E) 500-1000 /xm O.D, 360-470 /xm I.D, 170-250 /xm high; tetrasporangia not seen. Sexual conceptacles seen, but not sectioned (Figure 36D). TYPE-LOCALITY.—Sailus Besar, Paternoster Is- lands, Indonesia (station 315, Siboga Expedition), 36 m depth. HOLOTYPE.—Weber-van Bosse, 19-21 Febru- ary 1900, in herbarium of M. Foslie (TRH). Isotype: USNC. DISTRIBUTION.—Celebes. SPECIMENS STUDIED.—Hawaii: Honaunau, March 1971, 71-55-25. Maui: south-central, Au- gust 1971, 71-67-1, 71-68-14. Midway: South Is- land, August 1971, 71-82-29. Oahu: Waikiki, March 1971, 71-50-128. St Rogatien Bank: August 1971, 71-79-9, 71-79-11. REMARKS.—Next to Lithothamnium australe Fos- lie in Weber-van Bosse & Foslie, L. pulchrum was the most abundant species encountered at depths greater than 60 m (Figure 38). Scattered plants were taken in shallow water, and whereas no data on microhabitat were recorded, judging by the habit of Lithothamnium on algal ridges and coral reefs in the Caribbean, these were probably from cryptic locations. Lithothamnium pulchrum is a ma- jor rhodolith former on the Hawaiian banks and was especially abundant on St. Rogatien Bank. Lithothamnium australe Foslie FIGURES 37, 39, 40 Lithothamnium australe Foslie in Weber-van Bosse & Foslie, 1904:24.—Foslie, 1907a.—Dawson, 1944; 1954a; 1960b. —Taylor, 1945. Lithothamnium australe Foslie, 1900c: 13 [nomen nudum]. Lithothamnium corallwides P. & H. Crouan f. australis Foslie, 1895:8. Lithothamnium australe Foslie f. americana Foslie in Weber-van Bosse and Foslie, 1904:25. Lithophyllum australe (Foslie) Lemoine, 1917:131. Mesophyllum australe (Foslie) Lemoine, 1928:252; 1929. DESCRIPTION.—Light pink to bright red plants, usually with sparse primary branches (2-6 mm diam.) bearing numerous small, very irregular projections 0.5-2 mm diam. (Figure 39A,D,E); con- ceptacles medium-sized, rather rounded, flat- 54 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 36.—Lithothamnium pulchrum: A, branch of thallus, X 20; B, habit, X 1; c, section through vegetative crust, X 150; D, sexual crust with conceptacles (arrow), X 10; E, tetrasporangial conceptacles (arrow), X 5. (Specimen nos.: A, B, 71—79— 9; c, 71-79-11; D, 71-68-14; E, 71-67-1; micrographs reduced to 94%.) NUMBER 15 55 ep M 1 PERITHALLIUM 2 3 4 5 6 x i— O u 9- D L. p u I c h r um (13 plants) o L . a ustra I e (10 plants) OL LU u FIGURE 37.—Perithallial cell dimensions oi Lithothamnium pulchrum and L. australe. L. pulchrum FIGURE 38.—Depth distribution of Lithothamnium pulchrum. 56 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 39.—Lithothamnium australe: A, habit, X 1; B, section through epithallium showing "Lilhothammum-type ' cover cells, X 2200; c, section through thallus, note meristem (arrow) and perithallium, X 600; D, tetrasporangial conceptacles on surface of thallus (arrow), X 5; E, branch morphology, X 10. (Specimen nos.: A, D, E, 71-73; B, c, 71-72-1; micrographs reduced to 93%.) NUMBER 15 57 Depth m u c o TJ c D _Q < 10 20 30 40 50 60 70 80 90 30 20 10 -o— —o o o L.australe Oo- FIGURE 40.—Depth distribution oi Lithothamnium australe. topped multipored domes or conspicuous single- pored, pointed cones, often found on branch tips. Cover cells "Lithothamnium-type" (Figure 39B), single layer; cells 2-3 /xm long and 3-12 /xm diam. Intercalary meristem short and wide; cells 4-6 /xm long and 5-10 /xm diam. Perithallium multi- layered, fusions frequent (Figure 39c), primary pit connections with darkly staining plugs; cells often becoming elongate with depth in tissue (Figure 37), 3-16 /xm long and 3-11 /xm diam. Hypothallium thin, parallel to substrate, 30-55 /xm thick; cells 11-24 /xm long and 4-10 /xm diam. Tetrasporic conceptacles slightly raised, 300-500 /xm O.D. (none seen in section). Male concepta- cles unipored raised (50-150 /xm), 300-450 /xm O.D, 250-350 /xm I.D, 90-110 /xm high; sper- matangial mother cells dendroid, restricted to floor of conceptacle cavity; spermatia discoid, 1- 3 /un diam. TYPE-LOCALITY.—Gulf of California: La Paz, Baja California del Sur. LECTOTYPE.—Diguet (Foslie, 1895, fig. 7), in the herbarium of M. Foslie (TRH). Lecto-isotype: USNC. DISTRIBUTION.—Borneo, Celebes, Gulf of Cali- fornia, Indian Ocean, Pacific Mexico, Panama, Philippines, Timor. SPECIMENS STUDIED.—Maui: south-central, Au- gust 1971, 71-66-14, 71-67-16, 71-71-4. Midway: South Island, August 1971, 71-82-10. Molakai: south-central, August 1971, 71-69-7, 71-72-1, 71- 72-2; southwest, August 1971, 71-73-35. Oahu: Waikiki, March 1971, 71-50-94. REMARKS.—The typification of Lithothamnium australe is unclear from the literature. Lithotham- nium australe was published as a nomen nudum in Foslie (1900c) (Taylor 1945). The name was first validly published by Foslie in Weber-van Bosse and Foslie (1904). As to the typification, Foslie states of Lithothamnium australe f. americana: "This form constitutes the basis of Lithothammon aus- trale. . The said form from the Gulf of Califor- nia is here named f americana'" (Weber-van Bosse and Foslie 1904:25). The specimens from the Gulf of California are those collected by Diguet and given to Hariot, who sent them to Foslie. Foslie (1895) places these specimens under Lithothamnium coralloides P. & H. Crouan f. australis Foslie. It is misleading of Foslie to list as a synonym of Litho- thamnium australe "L. coralloides f. australis Foslie Norw. Lith. p. 62, ex parte'" (Weber-van Bosse and Foslie 1904:24), since he apparently did not wish to include any Norwegian material as types of Lithothamnium australe. It is also unfortunate that Foslie used as the species epithet the same name he had previously used as a forma name of a different taxon. Since a holotype was not designated by Foslie (Weber-van Bosse and Foslie, 1904) and a lecto- type has not been, designated, we choose the specimen corresponding with Foslie (1895, fig. 7) as the lectotype of Lithothamnium australe Foslie. Despite the "small-celled" meristem and pro- gressive elongation of the perithallium (Figure 37) seen in this species, the cover cell shape is 58 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES characteristic of the genus Lithothamnium, and lacking reproductive details we have chosen to retain it in Lithothamnium. Lithothamnium australe is the most abundant sin- gle species occurring in our collections from depths greater than 60 m (Figure 40). It was found throughout the Hawaiian chain wherever we dredged and occurred occasionally at 35 m, the limit of our diving range. It is abundant on bank situations as a small rhodolith or maerl- former and is the ecologic and anatomic "pair species" of the widespread Caribbean Lithotham- nium occidentale (Foslie) Foslie. Mesophyllum Lemoine, 1928 REMARKS.—Mesophyllum was the dominant ge- nus on banks over 65 m and especially over 80 m. Lithothamnium followed close behind in abun- dance, with Archaeolithothamnium and finally Hy- drolithon being lesser elements. Unlike the other genera, with one or two species each, Mesophyllum is represented by at least six species. We will describe and discuss the major elements, but con- siderable additional collecting preparation and study, especially in the central part of the Indo- Pacific, is necessary to satisfactorily accomplish a basic description of these bank floras. Key to the Species of Mesophyllum 1. Branched (branches quickly developed from a crustose base) M. madagascariensis Crustose 2 2. Crusts extensive smooth, sub-leafy; conceptacles > 600 /xm O.D 3 Crusts somewhat irregular; conceptacles < 600 /xm O.D 4 3. Few, scattered conceptacles, > 800 /xm O.D M. prolifer Densely concentrated mass of conceptacles, 600-800 /xm O.D M. purpurascens 4. Crusts irregular knobby, densely coated with raised conceptacles M. syrphetodes, new species Crusts not grossly irregular, conceptacles scattered M. fluatum, new species Mesophyllum madagascariensis (Foslie) Adey FIGURES 41A-C, 42 Mesophyllum madagascariensis (Foslie) Adey, 1970:25. Lithothamnium erubescens Foslie f. madagascariensis Foslie, 1902b:3.—Masaki, 1968. Lithothamnium madagascariensis Foslie, 1906:19. DESCRIPTION.—Initially crustose plants yellow- ish pink to bluish pink in shaded areas, typically developing small dense finger-like or slightly flat- tened bifurcating branches, usually less than 2 mm diam. and 3 mm long, sometimes forming massive anastomosing branched growth (Figure 41A,B); large domed conceptables, more abun- dant in sparsely branched crusts. Epithallium single, rounded to quite angular; cells 2-4 /xm long and 4-9 /xm diam. Intercalary meristem elongate; cells oval, 4-10 /xm long and 3-9 /xm diam. Perithallium multilayered, thin when thal- lus crustose, staining bodies present, fusions fre- quent; cells 3-12 /xm long and 3-9 /xm diam. (Figure 42). Heterocysts abundant, single or in small groups, 7-19 /xm long and 7-18 /xm diam. Tetrasporangial conceptacles multipored, few ini- tially raised (80 /xm) but soon becoming buried, walls of sterile cells remaining (Figure 41c), 140- 210 /xm I.D, 60-90 /xm high; tetrasporangia and bisporangia present, 60-90 /xm long and 35-50 /xm diam. Male conceptacles unipored (1 plant sectioned) 300-400 /xm O.D, 140-208 /xm I.D, 50-100 /xm high; spermatangial mother cells sim- ple, columnar, restricted to the conceptacle floor; NUMBER 15 59 FIGURE 41.—Mesophyllum madagascariensis: A, habit of typical specimen, X 1; B, surface of thallus showing branches, X 5; c, section through tetrasporic plant, note sterile filaments in conceptacle cavity (arrow), X 100. Mesophyllum prolifer: D, habit of typical specimen, X 2; E, sexual conceptacles (arrow), X 5. (Specimen nos.: A, 71-81-35; B, 71-81-31; c, 71-81-9; D, 71-68-5; E, 71-70-(46).) 60 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES M 1 ^ M. ma dag as ca r e n s i s (5 plants) M. syrphetodes (9 plants) PERITHALLIUM 3 4 5 6 -i 1- I 7 < 5 4 u FIGURE 42.—Perithallial cell dimensions of Mesophyllum madagascariensis and M. syrphetodes, new species. spermatia, ellipsoidal, in mucus strands, 2-3 /xm long and 1-2 /xm diam. TYPE-LOCALITY.—Madagascar. DISTRIBUTION.—Ceylon, Madagascar, Japan. SPECIMENS STUDIED.—Hawaii: Kawaihae, March 1971, 71-57-17. Oahu: Honauma, March HOLOTYPE.—In herbarium of M. Foslie 1971, 71-53-21; Kaneohe, August 1971, 71-81-9, (TRH). Isotype: USNC. 71-81-31, 71-81-35. NUMBER 15 61 REMARKS.—Spermatangial mother cells in this species only occur on the floor of the conceptacle, not the roof and walls. This is at variance with the findings of Lebednik (1978) for Mesophyllum, although this is true for Synarthrophyton patena (Townsend, 1979). Heterocysts are known in a few species of Me- lobesioideae (e.g., Phymatolithon rugulosum, Adey, 1964). They are known to the authors in a species of Mesophyllum important in southern Hokkaido, Japan, and have been reported from New Zea- land (Johnson, pers. comm.) and Australia (R. Townsend). The plants we worked with in Ha- waii are characterized by abundant single heter- ocysts. Although this plant apparently occurs though- out the Indo-Pacific and has a "pair species" (or possibly the same species, i.e., Mesophyllum erubes- cens (Foslie) Lemoine) in the tropical Atlantic, we encountered only a few large and and well-devel- oped heads (about 15 cm diam.), and there is a total of only 10 specimens in our collection. Eight of the collected specimens came from the outer reef off Kaneohe Bay, Oahu, and all specimens were found from 3-12 m depth. Mesophyllum prolifer (Foslie) Adey FIGURE 41D,E Mesophyllum prolifer (Foslie in Weber-van Bosse & Foslie) Adey, 1970:25. Lithothamnium prolifer Foslie in Weber-van Bosse & Foslie, 1904:18. DESCRIPTION.—Thin, dark pink somewhat leafy crust, surface usually nonlustrous, undulate, sometimes developing random low knobs (Figure 4ID). Epithallium single layer of rounded cells, 3-4 /xm long and 4-7 /xm diam. Intercalary meri- stem elongate; cells 4-5 /xm long and 5-6 /xm diam. Perithallium multilayered, irregular; cells somewhat square, 2-10 /xm long and 3-9 /xm diam. Hypothallium weakly coaxial 20-60 /xm thick; cells 7-27 /xm long and 5-12 /xm diam. Tetrasporangial conceptacles multipored, low and rounded domes, epithallium persistent, 800- 1200 /xm O.D. (none sectioned). Sexual concep- tacles single pored sharply raised (Figure 41E), none sectioned. TYPE-LOCALITY.—Pulu Sebangkatan, Borneo Bank, Borneo; 34 m depth, coral bottom and Lithothamnium. LECTOTYPE.—Weber-van Bosse, 971, 14 June 1899, in herbarium of M. Foslie (TRH) (Adey, 1970). DISTRIBUTION.—Borneo. SPECIMENS STUDIED.—Maui: South-central, Au- gust 1971, 71-68-5, 71-70-(46). REMARKS.—Only six specimens of this plant occur in the collection, all from about 80 m on the bank on the lee side of Maui. A presently unnamed "pair species," quite similar in anatomy and morphology, is known to the authors from deep reef situations in the Caribbean. Mesophyllum purpurascens (Foslie) Adey FIGURE 43 Mesophyllum purpurascens (Foslie) Adey, 1970:26. Lithothamnium funafutiense Foslie f. purpurascens Foslie, 1901b:18. Lithothamnium purpurascens Foslie, 1907a: 182; 1929.—Le- moine, 1917; 1965.—Papenfuss, 1968. DESCRIPTION.—Crusts smooth thin and glossy (Figure 43A). Epithallium a single layer of rounded cells, 2-3 /xm long and 5-8 /xm diam. Intercalary meristem elongate; cells 6-8 /xm long and 4-6 /xm diam. Perithallium multilayered, zoned from formation of conceptacles; cells 4-8 /xm long and 3-6 /xm diam. (Figure 43B). Hypo- thallium multilayered, coaxial; cells 10-16 /xm long and 3-4 /xm diam. (Figure 43E). Tetraspor- angial conceptacles centrally crowded (Figure 43A), raised (150 /xm) (Figure 43D), becoming buried (Figure 43B), 500-900 /xm O.D, 300-700 /xm I.D, 180-270 /xm high, walls of sterile cells remaining in cavity (Figure 43B,c); tetrasporan- gia not seen. TYPE-LOCALITY.—Koh Mesan and Cape Liant, Gulf of Thailand, 18 m depth. LECTOTYPE.—Danish expedition to Thailand, 62 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 43.—Mesophyllum purpurascens: A, habit showing centrally placed tetrasporangial concep- tacles, X 1; B, section through tetrasporangial thallus showing buried conceptacle, X 150; c, section through tetrasporangial conceptacle, X 300; D, surface view of tetrasporangial concep- tacles, X 20; E, coaxial hypothallium, X 1000. (Specimen nos.: A-E, 71-66-2.) NUMBER 15 63 1900, in herbarium of M. Foslie (TRH). Lecto- isotype: USNC. DISTRIBUTION.—Indian Ocean, Red Sea, Thai- land. SPECIMENS STUDIED.—Maui: south-central, Au- gust 1971, 71-66-2; 71-67-3. REMARKS.—Only two specimens of this species were found in this study. A morphological and ecological "pair species" of this species has not been found in the Caribbean. Mesophyllum syrphetodes, new species FIGURES 42, 44 DESCRIPTION.—Crustae prinum tenues rosa- ceae, superficie sordida, margines supercrescentes interdum abundantes, quam ob rem crusta usque ad plura mm. crassitudine facta (Figura 44A,C). Epithallium 1-2 cellulis crassum, cellula exteriore cellulam-obtegentem interdum efficiente; cellu- lae rotundatae 2-4 /xm long, atque 4-8 /xm diam. Meristema intercalare non manifeste elongatum; cellulis 3-7 /xm long, atque 3-5 /xm diam. Peri- thallium pluristratosum, tenue, ordinationem in- conspicue zonatam praebens, praecipue regioni- bus auctus conceptaculorum, et fusionibus non frequentibus, una heterocysta visa; cellulae rotun- datae, 3-9 /xm long, atque 3-8 /xm diam. (Figurae 44c, 42). Hypothallium pluristratosum, paral- lelum ad coaxiale, 20-75 /xm crass.; cellulae 10- 20 /xm long, 5-9 /xm diam. Conceptacula tetra- sporangialia multiporata, crebre disposita (Figura 44B) in quibusdam plantae regionibus repetite evenientia, frequenter obtecta facta et crassitu- dinem plantae augentia (Figura 44c), elevata (80-180/xm), tholiformia (Figura 44B,C), 250-400 /xmO.D, 150-300/xm I.D, 70-120/xm alt.; tetra- sporangia 80-90 /xm long, atque 25-45 /xm diam. Initially thin, reddish pink crusts with a dull surface, sometimes overgrowing margins abun- dant and crust achieving many mm of thickness by overgrowing (Figure 44A,C). Epithallium 1-2 cells thick, outer cell may form cover cell; cells rounded, 2-4 /xm long and 4-8 /xm diam. Inter- calary meristem not markedly elongate; cells 3-7 /xm long and 3-5 /xm diam. Perithallium multi- layered, thin with faint zonate pattern, especially in areas of conceptacle development, and fusions not common, one heterocyst seen; cells rounded 3-9 /xm long and 3-8 /xm diam. (Figures 44c, 42). Hypothallium multilayered, parallel to coaxial, 20-75 /xm thick; cells 10-20 /xm long 5-9 /xm diam. Tetrasporangial conceptacles multipored, densely spaced, (Figure 44B) occurring repeatedly in certain areas of the plant, frequently becoming overgrown and adding to thickness of plant (Fig- ure 44c), raised (80-180 /xm), domed (Figure 44B, c), 250-400 /xm O.D, 150-300 /xm I.D, 70-120 /xm high; tetrasporangia 80-90 /xm long and 25- 45 /xm diam, bisporangia present (Figure 44c). Sexual plants not seen. TYPE-LOCALITY.—Southwest Molokai, Hawaii (21°0'N, 157°0'W), 70 m depth. HOLOTYPE.—D. Child, 71-72-2, August 1971 (USNC), Figure 44A. PARATYPES.—Maui: south-central, August 1971, 71-71-1. Molokai: southwest, August 1971, 71-65- 1, 71-73-6. Midway: South Island, August 1971, 71-82-20. Oahu: Kaneohe, August 1971, 71-81-15. DISTRIBUTION.—Throughout the archipelago. REMARKS.—The specific epithet syrphetodes re- fers to the appearance of the thallus. A morphological and ecological "pair species" of this species has not been found in the Carib- bean. Mesophyllum Huatum, new species FIGURE 45 DESCRIPTION.—Crustae tenues rosaceae (25- 100 /xm), super superficiem "profluens," et non praesertim rugatae neque substratum plane illus- trantes (Figura 45A,B). Epithallium ex unico strato cellularum rotundatarum, 3-4 /xm long, atque 3-5 /xm diam. constans. Meristema inter- calare non elongatum, cellulis 2-6/xm long, atque 4-7 /xm diam. Perithallium pluristratosum, cel- lulis 4-7 /xm long, atque 4-12 /xm diam. Hypo- thallium pluristratosum coaxiale, 20-25 /xm crass.; cellulis 10-17 /xm long, atque 4-5 /xm crass. 64 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES WBP&$gt\ •-. .' ./ FIGURE 44.—Mesophyllum syrphetodes, new species: A, habit of type specimen, X 2; B, surface morphology of bisporangial plant, X 10; c, section through thallus, note overgrowing (arrow), X 200. (Specimen nos.: A, 71-72-2; B, c, 71-73-6; micrographs reduced to 92%.) NUMBER 15 65 FIGURE 45.—Mesophyllum flualum, new species: A, habit of type specimen, X 2; B, surface morphology of typical specimen, note tetrasporangial conceptacles (arrow) and concep- tacle scars (arrowhead), X 150. (Specimen nos.: A, B, 71-68-2.) Conceptacula tetrasporangialia multiporata, (solo uno secto) 525 /xm O.D, 350 /xm I.D, 150 super superficiem dispersa (Figura 45B), elevata /xm alt.; carposporangia ad periferiam concepta- (50-100/xm) 300-500/xm O.D. (solum a superficie culi restricta; sporangia 45-50 /xm long, atque visa). Conceptacula carposporangialia uniporata, 60-75 /xm diam.; cellula-coalescens reticulata. 66 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Thin pink crusts (25-100 /xm), "flowing" over the surface and not particularly crumpled or strongly reflecting the substrate (Figure 45A,B). Epithallium single layer of rounded cells, 3-4 /xm long and 3-5 /xm diam. Intercalary meristem not elongate; cells 2-6 /xm long and 4-7 /xm diam. Perithallium, multilayered; cells 4-7 /xm long and 4-12 /xm diam. Hypothallium multilayered coax- ial, 20-25 /xm thick; cells 10-17 /xm long and 4-5 /xm diam. Tetrasporangial conceptacles multi- pored, scattered over surface (Figure 45B), raised (50-100 /xm), 300-500 /xm O.D. (only seen from surface). Carposporangial conceptacles unipored, (only 1 sectioned) 525 /xm O.D, 350 /xm I.D, 150 /xm high; carposporangia restricted to the concep- tacle periphery; sporangia 45-50 /xm long and 60-75 /xm diam.; fusion cell reticulate. TYPE-LOCALITY.—South-central Maui-Hawaii, 92 m. HOLOTYPE.—D. Child, 71-68-2, August 1971 (USNC), Figure 45A. PARATYPES.—Maui: south-central, August 1971, 71-70-6. Molokai: southwest, August 1971, 71-68- 13. Nihoa: West, August 1971, 71-75. DISTRIBUTION.—Throughout south and central archipelago. REMARKS.—The specific epithet fluatum refers to the "flowing" of the crust over the substrate. Mesophyllum fluatum is represented in the collection by 11 specimens. All of these were collected from 60-90 m depth. No Caribbean "pair species" is known to exist for M. fluatum. Discussion Figure 46 compares the patterns of generic depth distribution in the eastern Caribbean area and the Hawaiian Archipelago. Considerable cor- relation can be seen in the major characteristics of generic dominance: Porolithon and Neogoniolithon dominate between intertidal to over ~ 30 m; Lithothamnium, Mesophyllum, and Archaeolithotham- nium dominate at depths greater than 50 m. This correlation, along with the large number of pair species generally having similar spatial ecologies, indicates that crustose coralline evolution has been very slow since the Indo-Pacific and the Caribbean were separated in the Miocene. The possibility of detailed interpretation of Neogene paleoecology, by using the relative abundance of genera found in limestone cores and outcrops, becomes apparent. Considerable detail in environmental interpre- tation can be gained using only the easily recog- nizable generic characteristics that often occur in well-preserved coralline fossils, especially type and placement of heterocyst fields, type of hypo- thallium, and the presence of sporangial sori and multipored conceptacles. Occasionally even more precise determination is possible with the occur- rence of certain distinctive species that have clearly defined ecological niches. For instance, Porolithon onkodes becomes dominant and forms thick crusts only on the shallowest wave-beaten outer and upper faces of algal ridges and coral reefs. This species is easily separated from the profusely branched P. gardinerii, the only other member of the genus found in the area. Reef-flat areas, although as shallow as the range dominated by Porolithon onkodes, seldom have appreciable amounts of that species. Such flats are often characterized by a red boring algae (melobesioid "C" of Littler, 1973A), which is probably a destructive rather than constructive agent. Crustose corallines are present usually on the shaded sides and undersides of rubble, the most common species being Hydrolithon reinboldu and the easily distinguishable Archaeolithothamnium erythraeum. These species are very useful as eco- logical indicators; both are sharply restricted to shallow water and have distinguishing morpho- logical characteristics making easy identification possible. The large cells of H. reinboldii set if off from the other species of the genus (Figure 15); the unique sori of Archaeolithothamnium should eas- ily be seen in fossilized material, and the differ- ence in their relative size is often great enough to distinguish A. erythraeum from A. episoredion—the latter is consistently associated with deeper water. The presence of abundant Hydrolithon and Ar- chaeolithothamnium in a geological sample, even if not determinable to species, would suggest shaded NUMBER 15 67 CARIBBEAN SEA • MAST0PH0R0IDEAE- -LITHOPHYLLOIDEAE- •MELOBESIOIDEAE- FREQUENCY OF OCCURRENCE ,10%, FIGURE 46.—Comparison of distribution patterns of crustose coralline genera (except Lithoporella) in idealized littoral zones in the Hawaiian Islands and Caribbean. reef flat environments only if Lithothamnium and Mesophyllum species were not present. Doty (1974) reports abundant Hydrolithon and Archaeolitho- thamnium from the reef crest, but they were not found in that position in this study. On more protected reef flats and in shallow lagoons, the elongate and slender type of branching seen in Porolithon gardineru and Neogoniolithon frutescens may be encountered. It is not likely that either of these would be found outside the reef and at any great 68 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES depth. It is possible to find representatives of species characteristic of much deeper water in shallow areas, but these occurrences are limited to cryptic situations where light conditions are similar to those existing at depth. Abundant coarsely branched Hydrolithon, if mixed with a branching Lithophyllum, would in- dicate deeper fore-reefs or island slopes (15-20 m). Large amounts of Lithothamnium and Meso- phyllum, especially if in rhodolith (nodule) form, can only indicate bank depths of greater than 50- 60 m. Although members of both of these genera do not occur in shallow water, they are not nodule formers in these environments but are represented by thin crusts in cryptic reef and ridge environ- ments. Shade-loving Neogoniolithon species usually dominate in those areas. Glossary Branch. Outgrowth of plant in which the height above the thallus surface is greater than the largest diameter of the growth (Figure 17c). Columella. One of several paraphyses situated directly be- low the pore in a uniporate tetrasporangial conceptacle (Figures 6E, 24E). Coralgal. A calcareous substrate consisting of coral and corallines or other calcareous algae. FIGURE 47.—Schematic representation of the conceptacle. A. Cover cell. More or less specialized cells (e.g., thick walls), which form the outer layer of epithallium and form a protective layer over the surface of the thallus (Figures 18c, 29D, 13c). "Lithothamnium-type'1 cover cells. Characteristic oi Lithothamnium (Figure 31c, D). High (conceptacle). Measurement C, Figure 47. I.D. (inside diameter, conceptacle). Internal widest diam- eter of a conceptacle when sectioned through the true median plane, measurement B, Figure 47. Intercalary meristem. Layer of meristematic cells giving rise to the epithallium in one direction and the perithal- lium in the other direction (Figures 18c, 29D). Layered (perithallium). Tissue whose cells form regular horizontal rows (Figures 24F, 27c). Leafy. Crust that is partially loose, especially at the grow- ing margins, and appearing as a "leaf1 upon the substrate. Mammillon. Outgrowth of the crust in which the height above the thallus surface is less than the largest diameter of the structure (Figure 17A,B). O.D. (outside diameter, conceptacle). External widest di- ameter of a conceptacle when viewed from the surface, measurement A, Figure 47. Parallel (hypothallium). Tissue where the filaments run in line with the substrate and are not coaxial. See "Simple'- (Figure 8B). Pair species. Similar taxa separated by land masses or other barriers. The taxa exhibit similar morphology (morpho- logical p.s.) and/or exist in similar niches (ecological p.s.). Plumose (hypothallium). Ascending and descending fila- ments from a single-or few-layered central plane, which give the hypothallium the look of a fountain in section (Figure 2F). Rugose (surface). A surface with an irregular low-mounded appearance as though covered with wrinkles, not corru- gations as Foslie (1906) used for Porolithon antillarum (Foslie & Howe) Foslie. Rugulose (surface). Diminutive of rugose. Simple (hypothallium). Noncoaxial. Spermatangial mother cell. Cell giving rise to spermatia. Tessellate (surface). Mosaic surface of irregular patterns, sometimes polygonal, not the spiral whirls of Tenarea tes- selatum that Lemoine (1929) incorrectly considered tesse- late (cf. to surface oi Hydrolithon reinboldii). Zonate, zoned (perithallium). 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Standing Stock Measurements of Crustose Coral- line Algae and Other Saxicolous Organisms. Jour- nal of Experimental Marine Biology and Ecology, 6:91- 99. 1973a. The Population and Community Structure of Ha- waiian Fringing Reef Crustose Corallinaceae. Jour- nal of Experimental Marine Biology and Ecology, 11:103-120. 1973b. The Distribution, Abundance, and Communities of Deep-Water Hawaiian Crustose Corallinaceae. Pacific Science, 27(3):381-390. Masaki, T. 1968. Studies on the Melobesioideae of Japan. Memoirs of the Faculty of Fisheries, Hokkaido University, 16(1/2): 80 pages. Masaki, T., and J. Tokida 1963. Studies on the Melobesioideae of Japan, VI. Bul- letin of the Faculty of Fisheries, Hokkaido University, 14(1): 6 pages. Okamura, K. 1936. Nippon Kaiso-shi. Frontispiece + [9 + 6] + 964 + [11] pages. Tokyo. Papenfuss, GF. 1968. A History, Catalogue, and Bibliography of Red Sea Benthic Algae. Israel Journal of Botany, 17(1-2): 118 pages. Philippi, R.A. 1837. 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Index (Synonyms and page numbers of principal accounts in italics) accretum, Neogoniolithon, 20 americana, Lithothamnium australe f., 53, 57 Archaeolithothamnium, 6, 7, 47, 48, 52, 58, 66, 67 dimotum, 48 episoredion, 48, 50, 51, 52, 66 episporum, 51 erythraeum, 48, 49, 50, 66 fosliei, 23 howei, 51 pacificum, 51 australe, Lithophyllum, 53 Lithothamnium, 53, 55-58 Mesophyllum, 53 australis, Lithothamnium coralloides fi, 53, 57 b0rgensenii, Hydrolithon, 26 breviclavium, Goniolithon, 26 Hydrolithon, 26, 28-32, 34, 42 californiense, Lithophyllum, 40 caribaeum, Neogoniolithon, 16 cephaloides, Lithophyllum, 40 cerebelloides, Lithophyllum, 25, 26 clavacymosum, Neogoniolithon, 19, 21, 22, 23 congestum, Lithophyllum, 10, 12, 40 conicum, Hydrolithon, 13, 34 Neogoniolithon, 13 Paragoniolithon, 13, 14, 15, 20, 21, 23,42 dimotum, Archaeolithothamnium, 48 epilaeve, Kvaleya, 25 episoredion, Archaeolithothamnium, 48, 50, 51, 52, 66 episorum, Archaeolithothamnium, 51 epiyessoense, Ezo, 25 erubescens, Mesophyllum, 61 erythraeum, Archaeolithothamnium, 48, 49, 50, 66 Lithothamnium, 48 Sporolithon, 48 Ezo epipyessoense, 25 fluatum, Mesophyllum, 58, 63, 65, 66 fosliei, Archaeolithothamnium, 23 Goniolithon, 23 Lithophyllum, 23 Lithothamnium, 23 Neogoniolithon, 15, 19, 21, 23, 24, 25 frutescens, Neogoniolithon, 5, 67 ganeopsis, Lithophyllum, 37, 38, 42, 44, 47 gardineri, Lithophyllum, 10 Porolithon, 9, 10, 11, 12, 40, 66, 67 Goniolithon breviclavium, 26 fosliei, 23 oncodes, 7 pallescens, 40 propinguum, 12 reinboldu, 25 solubile, 12 tessellatum, 35 howei, Archaeolithothamnium, 51 Hydrolithon, 6, 25, 26, 30, 34, 44, 48, 58, 67, 68 b0rgensenii, 26 breviclavium, 25, 26, 28-32, 34, 42 conicum, 13 laeve, 25, 28, 29, 31, 32, 34 megacystum, 25, 28, 29, 32, 33, 34 reinboldii, 25, 26-29, 31, 32, 34, 36, 44,66 insipidum, Lithophyllum, 37, 38, 44, 45,47 kotschyanum, Lithophyllum, 10, 12, 37, 38-40 Lithothamnium, 37 Kvaleye epilaeve, 25 laeve, Hydrolithon, 28, 29, 31, 32, 34 lenormandii, Phymatolithon, 20 lichenoides, Mesophyllum, 25 Lithophyllum, 6, 7, 37, 44, 47, 67, 68 australe, 53 californiense, 40 cephaloides, 40 cerebelloides, 25, 26 congestum, 10, 12, 40 fosliei, 23 ganeopsis, 37, 38, 42, 44, 47 gardineri, 10 insipidum, 37, 38, 44, 45, 47 kotschyanum, 10, 12, 37, 38-40 nitorum, 44 okamurai, 40, 41 oncodes, 7 onkodes, 7 pallescens, 31, 37, 38, 40, 41, 42 punctatum, 37, 38, 45, 47 reinboldu, 25, 26 tessellatum, 35 Lithoporella, 6, 34, 35, 67 melobesioides, 6, 34, 35 Lithothamnium, 6, 7, 15, 47, 48, 52, 53, 58, 66-68 australe, 53, 55-58 australe i. americana, 53, 57 coralloides fi australis, 53, 57 erubescens i. madagascariensis, 58 erythraeum, 48 fosliei, 23 funafutiense i. purpurascens, 61 kotschyanum, 37 madagascariensis, 58 occidentale, 58 onkodes, 7 pallescens, 40 prolifer, 61 pulchrum, 52, 53, 54, 55 purpurascens, 61 madagascariensis, Lithothamnium, 58 Mesophyllum, 55, 59, 60 Lithothamnium erubescens fi, 58 mammillare, Neogoniolithon, 26 Mastophora melobesioides, 34 "megacarpum," Neogoniolithon, 25 megacystum, Hydrolithon, 28, 29, 32, 33, 34 melobesioides, Lithoporella, 6, 34, 35 Mastophora, 34 73 74 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Mesophyllum, 6, 7, 58, 61, 66-68 australe, 53 erubescens, 61 fluatum, 58, 63, 65, 66 lichenoides, 25 madagascariensis, 58, 59, 60 prolifer, 58, 59, 61 pulchrum, 53 purpurascens, 58, 61, 64 siamense, 5 syrphetodes, 58, 60, 63, 64 Millepora, 10 Neogoniolithon, 6, 15, 25, 30, 48, 66-68 accretum, 20 caribaeum, 16 clavacymosum, 15, 19, 21, 22, 23 conicum, 13 fosliei, 15, 19, 21, 23, 24, 25 frutescens, 5, 67 mammillare, 26 "megacarpum," 25 pacificum, 20 propinguum, 12 rufum, 15, /6, 18-21 rugulosum, 15, 16, 17 solubile, 12 nitorum, Lithophyllum, 44 occidentale, Lithothamnium, 58 okamurai, Lithophyllum, 40, 41 oncodes, Goniolithon, 1 Lithophyllum, 1 Porolithon, 7 onkodes, Lithophyllum, 7 Lithothamnium, 7 Porolithon, 3, 7, 8-10, 20, 21, 23, 66 pachydermum, Porolithon, 10 pacificum, Archaeolithothamnium, 51 Neogoniolithon, 20 pallescens, Goniolithon, 40 Lithophyllum, 31, 37, 38, 40, 41, 42 Lithothamnium, 40 patena, Synarthrophyton, 20, 61 Paragoniolithon, 6, 12, 15, 67 conicum, 13, 14, 15, 20, 21, 23, 42 solubile, 12, 15 "typica," 15 Phymatolithon lenormandii, 20 rugulosum, 61 Porolithon, 6, 7, 12, 66 gardineri, 7, 9, 10, 11, 12, 40, 66, 67 oncodes, 7, 23 onkodes, 3, 7, 8-10, 20, 21, 23, 66 pachydermum, 10 reinboldu, 26 prolifer, Lithothamnium, 61 Mesophyllum, 58, 59, 61 propinquum, Goniolithon, 12 Neogoniolithon, 12 prototypum, Tenarea, 36 pulchrum, Lithothamnium, 52, 53, 54, 55 Mesophyllum, 53 punctatum, Lithophyllum, 37, 38, 45, 47 purpurascens, Lithothamnium, 61 Lithothamnium funafutiense fi, 61 Mesophyllum, 58, 61, 62 reinboldii, Goniolithon, 25 Hydrolithon, 25, 26-29, 31, 32, 34, 36, 44, 66 Lithophyllum, 25, 26 Porolithon, 26 rufum, Neogoniolithon, 15, 16, 18-21 rugulosum, Neogoniolithon, 15, 16, 17 Phymatolithon, 61 siamense, Mesophyllum, 5 solubile, Goniolithon, 12 Neogoniolithon, 12 Paragoniolithon, 12, 15 Sporolithon, 47, 48 erythraeum, 48 Synarthrophyton patena, 20, 61 syrphetodes, Mesophyllum, 58, 60, 63, 64 Tenarea, 6, 35, 67 prototypum, 36 tessellatum, 6, 20, 21, 35, 36 tessellatum, Lithophyllum, 35 Goniolithon, 35 Tenarea, 6, 20, 35, 36 "typica," Paragoniolithon, 15 REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION Manuscripts intended for series publication receive substantive review within their originating Smithsonian museums or offices and are submitted to the Smithsonian Institution Press with approval of the appropriate museum authority on Form SI-36. 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