The Crustose Coralline Algae (Rhodophyta: Corallinaceae) of the Hawanian Islands = WALTER_H. -ADEY, ROBERTA A. TOWNSEND, Leer oe — . ln cee psec os: iu -and - Rein, nl ae meer — eer rat aR cs Og ane tiga ~y i ty sa Y rat a“ “WILLIAM ' T. BOYKINS ~ a SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of “diffusing knowledge’ was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: ‘‘It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.’”’ This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution | SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES © NUMBER 15 The Crustose Coralline Algae (Rhodophyta: Corallinaceae) of the Hawaiian Islands Walter H. Adey, Roberta A. Townsend, and William T. Boykins ISSUED DEC 17 1992 SMITHSONIAN PUBLICATIONS —_~ 2 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 of 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 handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Srithsonian 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. IL. Boykins, William T. III. Title. IV. Series. QOK569.C8A33 589.4/1 81-21247 AACR2 Contents Meno CC hlOnvye Maye 2 ec, tos Ween ac FACKNOWICdeINENtS eres ee Materials and Methods ........... MIR AKOMOMIV AM te acres sts. eae Eresentation.of Material ~).0. 5... (CORALIMINVNGIOND “Joscccusesseecesens Kevatonenensubtamulies andi General: iene oe ge es MASTOPHOROIDEAE (Svedelius) Setchell, 1943'00....... ......... Porolithon (Foslie) Foshe, 1909 . INeyatonthe: Species) a... eovoletnonmonikodesn (Idey.duccly)shOslie tn sant ore ee Joona gorenngp (ROSIE) SOS" 34 oh eae a vade see aera ne eee Paragoniolithon, new genus ..... Paragoniolithon conicum (Dawson), new combination ............ NeosomolithonssetchiellySaNiasony NOLS) ee ea Ney RtOnEMe SPECIES hays a) INeoconvolithonmmuowloscumssINeWASPECIES! =. oF es ee 2 a oe INeoconvolithonmftum ene Was PECs Ysa eee ek ee Neogoniolithon clavacymosum, new species ..................-..--: Neogoniolithon fosliec (Heydrich) Setchell & Mason .............. FHydrolithon (Foshe) Foslie, 1909 IKevatontne Species) 3) 4-1: HHydrolithon reinboldu (Weber-van Bosse & Foslie) Foslie ......... Heyanoleihon) oneuiclaviuma(boslieFoslie 4a) 256.9. 42 4). 2) Hydrolithon laeve, new species ily drolthoneumegacy StunPanNeWaSPeCles 2s -mas4 ae eee ae 3 ae Lithoporella (Foslie) Foslie, 1909 ninoponellamelobesvowdess(Poslic)Foslie =) 9.22524. ee a LITHOPHYLLOIDEAE Setchell, 1943 HicnarcamDOGyplOS2 ee eae? Nicwarcautesselatium«(lkemoine)pleittlem ye Lithophyllum Philippi, 1837... INS TO WS S)NSHES Soe es eg JLaivanoplintin (SOUSA. OTN? o.oo ocboddsoespaccbacaucsanoue Lithophyllum pallescens (Foslie) OSE eer ete ome oe rn a ligihopyllmscancopstsaine Was pe Clesh ene er ae ane ithopiayliumncnsypidtumeine WaSCClES=. yee a ee ak Lithophyllum punctatum Foslie il 15 Www iwoh NM bh rh PP MOK Doo ol Wwiwoiwo ow oOo NINN OOO Pe sa NM © SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES MELOBESIOIDEAE (J.E. Areschoug) Yendo, 1902 .................... ArchacolithothamniumiNothpletz aloe pe ern ae Lithothamnum Philippi, 1837 Key to the Species of Archacolithothammum 10 en eee Archaeolithothamnium erythraeum (Rothpletz) Foslie .............. Archaeolithothamnium episoredion, new species Key to the: Species! ii: 2235 ake nce ae ec ee Lithothamnium pulchrum Weber-van Bosse & Foslie .............. Lithothamnium australe Koshiey= 4) ee ee Mesophylium Wemoimes 923 ae eee oe Key toithe Species of Mesoplyllun v= )) ane Mesophyllum madagascariensis (Foslie) Adey ...................... Mesophyllum: prolifer (Boslie) Adey. a, sn) ae an eee Mesophyllum: purpurascens (Koslie)-Adey (2) 752447) eee Mesophylium syrphetodes, new Species: = nn ea) eee Mesophylium fiiatum, new Speciess 5) 1. see eee Discussion: 22a ea ers ee ee Ie a Glossary) sce coicge ee dee tea dio ener een a Mi neh hee re tre rns SIC er Literature Cited: yc ee e Index 47 47 48 48 51 a3 oy) a3 53 58 58 58 61 61 63 63 66 68 69 13 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 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- formation. 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, Australva. 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 2 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 Teritu; 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- lum 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 1s a complex three-dimensional shape with several SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 20¢4 009! “Spueysy ULIIPMBFY IY) UT SOzIS uOTI[[OF)— | ANNO 9! OL! _. _ STWOHS “ery FLvOlds ! Ail, HONS ! eoN, ~ 2 ol AVMOIW N3ILVDOY 1S! 1a-Ag STIOVNNIG _ wANaBVvd ounve d E NO.LdWWVHHLYON pie dA ‘ YNVE WLIVe ouW a 1 oNVSAWT"” wnve 2! YASINOld 61k | SLI _LMSNVISIT YWNVG NOWIVS SWu3H ¥ Vad (cp) T¥OHS YNYG OWIN viIaWvo = NUMBER 15 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 stamense (Fos- lie) Adey, although present, have not been in- cluded. The following abbreviations are used in this study. PC Muséum d’Histoire Naturelle, Labora- toire de Cryptogamie, Paris TRH herbarium of M. Fosle, Trondheim, Norway UC herbarium of the University of Califor- nia US U.S. National Herbarium, Smithsonian Institution USNC 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, 1.e., tetrasporangial conceptacles single DORE TU erate! Cer ete care ek 3. 9 6 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Tetrasporangia with caps, 1.e., tetrasporangial conceptacles multipored (Melobesioidéae)iish 5 ea ty ec, er 7 2. Perithallial secondary pit connections present, cell fusions more or less rare or lacking (Lithophylloideae)™ 2) nee re ee 3 Perithallial secondary pit connections absent, cell fusions present (Mastophoroideae) 2.3 Mee eh 4 3. Perithallium palisade, single-layered hypothallium .............. Tenarea Perithallium nonpalisade, single-layered or multilayered coaxial hypothalliwin 3.4 os! se wee ee i acre ate ets ee ae Lithophyllum 4.) FHeterocysts;absemt o).27 0 einen an eae ot ey ae Lithoporella Heterocysts present: 220i, os aga Sate ee ey) 5. iy pothallium#singlejlayeredyerrs ssi e er eee Hydrolithon Hypothallium: multilayered’ 2°. a) tee oie ee 6 6. Heterocysts vertical rows or single, hypothallium coaxial or simple-parallel OES Willys taecaitie eee ee meee ee Bei Res arn rs ee ok Neogoniolithon Heterocysts loosely grouped into horizontal fields, hypothallium COA xTalllt tice et ie eae inn ane or sh AeA MIRE Paragoniolithon, new genus Heterocysts grouped into tight horizontal pustulous fields, hypothallium simple-parallelror plumose my) eee ee Porolithon 7. “Lithothamnium-type” cover cells not present, hypothallium COAXIAL 5:45 ts ag sic oda cee enya ea thr ee Mesophyllum “Lithothamnium-type” cover cells present, hypothallium noncoaxial ..... 8 8. Tetrasporangia in broad sori, partially calcified walls between adjacent SPOrangilals 7 yt ene saree nin: vaeeiot Wane Archaeolithothamnium Tetrasporangia in conceptacles, hypothallium simple-parallel 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- Fete Meese EN tee Lithothamnium lium, delineates the four genera of Mastophoro- ideae: Porolithon, Paragoniolithon, Neogoniolithon, and FHydrolithon. 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) Foshe (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- othammum 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 1f 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 Porolithon onkodes (Heydrich) Foslie Ficures 2—4 Porolithon onkodes (Heydrich) Foslie, 1909:57.—Gordon, Ma- saki, and Akioka, 1976.—Lee, 1967.—Lemoine, 1966.— Taylor, 1950.—Womersley and Bailey, 1970. Lithothamnion onkodes Heydrich, 1897a:6, pl. 1: fig. 11. Lithophyllum oncodes Heydrich, 1897c: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 2p,c). Epithallium a layer of rounded cells 1 to 3 cells thick (2-6 wm long, 5-9 wm diam.). Intercalary meristem large celled, occur- Ned tn eb, Nl Go or Seca aya AL CYS P. onkodes P. gardineri ring immediately below the epithallium (4-11 wm long, 4-8 wm diam.) (Figure 3). Perithallium multilayered, fusions common, cells 4—13 wm long and 4-10 um diam. Heterocysts, 10-30 um long and 4-14 wm diam., throughout perithallium forming compact fields to 100 wm diam. (Figure 2r,F). Hypothallium multilayered, plumose (Fig- ure 2F), 50-150 (350) ym thick, may be thin with filaments oriented parallel to the substrate, cells 11-24 um long and 5-14 wm diam. Tetrasporic conceptacles unipored, scattered, convex, small (240-300 pum outside diameter (O.D.), 110-230 jm inside diameter (I.D.), and 30-140 um high), with roof apex about 40 um above surface (Figure 2p), columella present; tetrasporangia 50-120 um long and 20-50 wm diam., restricted to concep- tacle periphery. Cystocarpic conceptacles uni- pored, slightly raised above surface, 150-210 um I.D., 80-140 um high; carpospores (35-70 fm long, 25-70 fm wide) arising from the periphery SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES crust with tetrasporic conceptacle (arrow) and > habit, X 1; B heterocyst fields (arrowhead), X 7; > Figure 2.—Porolithon onkodes: A X 12; p, tetrasporic concep- > plumose hypothallium and buried x 500; F, 82- > crust with heterocyst fields C, 3 E, heterocyst field at thallus surface heterocyst field (a 76; , X 250; tacle -E, 71-82- > « eS 82-85; p, 71-81 mile Cc 84; , /1- B > A 200. (Specimen nos:.: x ), 8; micrographs reduced to 85%.) rrow 71-59-2 F, 3 | | | NUMBER 15 9 e = 18 a Ie = eo Ze Lu =n —_! —_t Lu NS) © P. gardineri (4 plants ) 4 2 © P. onkodes (25 plants) co] E iB a —=- —_—_——.—— —— - oP ene —= Lu = < Z ° a s+ 6 —_! Lu UO 5 ep ] 2 3 5 $ 7 8 M 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 pm I.D., 20-35 um high; spermatangial mother cells arising from the floor of the conceptacle only; spermatia at first elongate, crescent shaped, 3-5 ym long, 1-2 um diam., becoming ellipsoidal. Type-Loca.ity.—On coral ridges, Tami Is- land, northwest edge of the Huon Gulf, New Guinea. Ho.otypee.—Heydrich 97, March 1892, col- lected by Bambler, in herbarium of M. Foslie (TRH). DistrisuTion.—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.—Hawaz: 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 gardinert (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) Foshe 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 Muil- lepora (Coelenterata) are the dominant builders of these more massive and higher algal ridges. Porolithon gardineri (Foslie) Foslie Ficures 3-5 Porolithon gardinert (Foslie) Foslie, 1909:57.—Lee, 1967.—Lit- tler, 1971b.—Taylor, 1950. Relative Abundance % Lithophyllum gardinert Foshie, 1907a:190; 1929. Lithophyllum (Porolithon) gardineri Foslie, 1907b:30; 1909. DeEscripTion.—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, |—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 wm long, 8-10 pm diam. Perithallium multilayered, fusions common, cells 5-10 pm long and 6-11 pm diam (Figure 3). Heterocysts throughout perithallium, 13-26 ym long and 7-15 ym diam. Hypothallium not meas- ured. No conceptacles, asexual or sexual, were seen in sections of our material. Type-Locaity.—Coetivy, Seychelle Islands, Indian Ocean. Lectrotype.—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 50 60 70 80 90 Ficure 4.—Depth distribution of Porolithon onkodes (0) and P. gardineri (A) in Hawaii. NUMBER 15 Ficure 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%.) iba 12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES P. gardinert have markedly distinct, pustulous het- erocyst fields (Figure 5c); however, micrograzing, probably by young limpets and perhaps chitons, can remove this pustulous surface (Figure 5p). 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 gardinert and Lithophyllum kotschyanum (see below). At a single station in the lagoon of Midway Atoll, a fine-branched Porolithon (Figure 5p) that is ten- tatively placed in P. gardinert was found in abun- dance (it was not included in the measurements). Its surface texture is very similar to that of the P. gardinert found in the remainder of our stations, and the branching patterns are similar, though markedly smaller than P. gardinerr, 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. Paragoniolithon, 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.—Paragoniolithon solubile (Foslie & Howe in Foslie), new combination. Basionym: Gonvolithon 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. Typr-Loca.ity.—Culebra Island, Puerto Rico. Lecrotype.—Howe, 4375, in herbarium of M. Foslie (TRH) (Adey, 1970). Lecto-isotype: USNC. DistrrpuTion.—Tropical seas, typically at moderate depths, 5-25 m, in reef environments. NUMBER 15 Paragoniolithon conicum (Dawson), new combination Ficures 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- a ¢ =, CL Le See he ek ie We SO Tore > oN es Bee \~ Nee a 13 quently deep red but ranging from yellow-pink to maroon, margin smooth to undulating, within several mm to | cm from the margin very abun- dant nonpustulous heterocyst fields develop (50- 100 wm diam) (Figure 64,B) giving the surface a slightly rough appearance. Epithallium a single layer of rounded cells, 3-9 um diam. Intercalary meristem cells elongate, cells 4-9 um long and 5- 10 wm diam., with some progressive elongation occurring throughout the perithallium (Figure 7). Figure 6.—Paragoniolithon 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, p, 71-78-13; F, 71-55-35; micrographs reduced to 70%.) 14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES M PERITHALLIUM A ep 1 2 3 4 5) 6 If 8 els lig 7 O Le me 6 L2 a fo) P.conicum (20 plants) Oy 2 wo CELL DIAMETER ron Relative Abundance % 0 10 20 30 40 P.conicum 50 60 70 80 90 Depthm Figure 7.—Paragoniolithon conicum: a, mean epithallial and perithallial cell diameter and length; B, depth distribution. Perithallium multilayered, fusions abundant, cells 3-11 wm long and 4-13 wm diam. (Figures 6c, 7). Heterocyst fields throughout the perithal- lium, loose, with scattered filaments between the heterocysts (Figure 6c,p), heterocysts 7-26 fm long and 8-21 wm diam. Hypothallium parallel to substrate, weakly to strongly coaxial, 30-400 fam thick; cells 8-25 long and 4-16 wm diam. (Figure 6F). Tetrasporic conceptacles unipored, regularly spaced, convex, conical, small (350-530 pm O.D., 140-360 pm I.D., 50-160 um high), strongly raised, roof apex 40—200 um above plant surface (Figure 6£), columella present; tetraspo- rangia restricted to the periphery of the concep- tacle, 80-200 um long, 45-65 wm diam. No pro- carpic or cystocarpic material found in our spec- imens. Male conceptacle unipored, 190-260 um I.D., 40-80 pm high, raised but more rounded than tetrasporangial conceptacle; spermatangial mother cells arising from the base of the concep- tacle only; spermatia ellipsoidal, 3-6 zm long, 1 3 wm diam. Type-Loca.ity.—Intertidal reef at Binners Cove, Isla Socorro, Revillagigedo Archipelago, Pacific Mexico. Ho.otypee.— Dawson, 1953; (US). DistTrRIBUTION.—Xisha Islands, China; Guam; Pacific Mexico. SPECIMENS STUDIED.—French Frgate: La Pe- rouse, August 1971, 71-78-13. Hawai: 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, 12148, 19 November NUMBER 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- thammum 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 15 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. Paragoniolithon conicum occurs commonly in mid- depths (Figure 7) at all stations from Midway to Hawai. 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 of Paragoniolithon, 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 Paragonio- 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 |. Plants thin crusts, rugulose; low-domed conceptacies < 200 um O.D. ite AR eae N. rugulosum, new species Piantsacnicker not nuguloseconceptacles= 9200 jim OD) 272222222. 2 2. Crusts adherent, knobby, bright pink to yellow brown, conceptacles 200- AX O} sari OY De Sas Ae te eae SRO Sere ae. N. rufum, new species Crusts leafy to sub-leafy, pink yellow to dark brown, conceptacles > 400 (itiae ORD) Reo newt Leet teh 5 Baa! Goa ne GNA ae aOR 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 See nee N. clavacymosum, new species Crusts sub-leafy, becoming thick (to 5 mm) without protuberances, con- ceptacles low conical, leaving no scars 5 ee ia AS ee ee ee N. fosliet 16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Ficure 8.—Neogoniolithon rugulosum, new species: A, habit of type specimen, X 1; 8, 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 Neogoniolithon rugulosum, new species Ficure 8 DescriIPpTION.—Crustae tenues (< 200 pm), 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 um long. atque 9-16 ym diam. Meristema intercalare elon- gatum, admodum infra epithallium, cellulis 10- 19 um long. atque 6-10 wm diam. Perithallium multis cellulis crassum (Figura 88), fusionibus frequentibus, cellulis 4-8 wm long atque 3-6 um diam. Heterocystae per perithallium dispersae, singulae, raro in coacervationibus horizontalibus repertae, 12-14 wm long atque 7-9 wm diam. Hypothallim e 2-8 stratis celularum constans (Figura 8B), subparallelum ad substratum, 25-75 uum crass.; cellulae 9-30 wm long atque 6-15 wm diam. Conceptacula tetrasporica uniporata (160- 200 um O.D., 70-90 um I1.D., 30-50 pm alt.) tecta plana ad paululum tholiformia super crustas cir- cumdantes; tetrasporangia trans pavimentum conceptaculi disposita, 50-85 wm long atque 25- 50 pm diam (Figura 8p). Crusts thin (< 200 wm), 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 um long and 9-16 pm diam. Intercalary meristem elongate, immedi- ately below epithallium, cells 10-19 um long and 6-10 pm diam. Perithallium, many cells thick (Figure 8B), fusions common, cells 4-8 um long and 3-6 ym diam. Heterocysts scattered through- out perithallium, single rarely in horizontal as- semblages, 12-14 um long and 7-9 wm diam. Hypothallium 2-8 cell layers (Figure 8B), sub- parallel to substrate, 25-75 pm thick; cells 9-30 fm and 6-15 ym diam. Tetrasporic conceptacles single pored, 160-200 um O.D., 70-90 um L.D., 30-50 um high, roofs flat to slightly domed above 17 surrounding crusts; tetrasporangia across floor of conceptacle, 50-85 wm long and 25-50 wm diam. (Figure 8p). No sexual material collected in present study. Type-Loca.ity.—Palea Point, Hanauma Bay, Oahu, Hawaii (21°07’N,157°50’W), on volcanic rock, intertidal zone. Ho.otyre.—D. Child, 71-53-2, 1 April 1971 (USNC), Figure 84,3 ParaATyPES.—Hawau: 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 Ficures 9-114 Description.—Crustae primum satis tenues, marginibus arcte 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, tiones membranae esterioris tangentialis perspi- cuae; cellulis 2-6 um long. atque 4-9 um diam. incrassa- Meristema intercalare parum elongatum, cellulis 3-9 um long. atque 3-10 wm diam. Perithallium pluristratosum, aliquantula elongatione effecta (Figura 10), fusionibus frequentibus, abrupte zon- atum (Figura 9a), cellulis 3-11 wm long. atque 2- 11 pm diam. Heterocystae singulae, interdum 18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 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; p, hypothallium, X 350. (Specimen nos.: A, B, 71-50-80; c, 71-82-57; p, 71-50-20; micrographs reduced to 89%.) | NUMBER 15 19 M PERITHALLIUM eal Te TO (um) aE = © Z LL — | —d — Li UO 4 a] 3 (o) = A 58 eZ ner 7 Llu = << SG A = —d Fe UO 5 6 if. 8 o N. rufum (31 plants) © N. clavacymosum (24 plants) ON. fosliei (12 plants) Ficure 10.—Cell diameter and length data for Neogoniolithon rufum, new species, N. clavacymosum, new species, and N. foslzez. rarae (aliqunado laxe verticaliter horizontali- terque aggregata) 7-29 um long. atque 4-12 um diam. Hypothallium simplex ad coaxiale, ad sub- stratum parallelum (Figura 9p), 25-130 (200) um crass., cellulis 9-25 um long. atque 4-10 wm diam. Conceptacula tetrasporica uniporata, 30-150 um super crustam elevata (Figura 9B), 200-400 um O.D., 80-300 ym I.D. atque 50-110 pm alt., per perithallium obruta; tetrasporangia 50-175 um long. atque 29-65 wm diam. super totum pavi- mentum conceptaculi disposita. Conceptaculum procarpicum uniporatum, 380 um I.D., atque 100 pum alt., omnis cellula sustinens duo initia carpo- gonialia ramorum habens. conceptacula cystocar- pica uniporata, 210-290 um I.D. atque 60-90 fm alt., per perithallium cum sporis obruta; carpo- sporae 50-70 um long. atque 40-70 um diam., solum in periferia cellulaecoalescentis magnae 20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES discoideae dispositae (285-340 wm diam.) (Figura 9c). Conceptaculum masculum uniporatum, 50- 130 pm elevatum, 215-350 um I.D. atque 50-100 pum alt.; spermatia ad pavimentum conceptaculi restricta (Figura 9c); 2-4 spermatia in omni cel- lulamatricalee, ellipsoidea ad discoidea (3-7 um long., 2-3 wm 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 4m long and 4-9 wm diam. Intercalary meristem weakly elongate, cells 3-9 um long and 3-10 4m diam. Perithallium multilayered, some elonga- tion occurring (Figure 10), fusions common, sharply zonate (Figure 9c), cells 3-11 um long and 2-11 wm diam. Heterocysts single, sometimes rare (occasionally grouped loosely vertically and horizontally) 7-29 um long and 4-12 wm diam. Hypothallium simple to coaxial, parallel to sub- strate (Figure 9), 25-130 (200) um thick, cells 9- 25 pm long and 4—10 wm diam. Tetrasporic con- ceptacles unipored, raised 30-150 wm above the crust (Figure 9p), 2001-400 wm O.D., 80-300 pm I.D., and 50-110 wm high, buried throughout perithallium; tetrasporangia 50-175 wm long and 29-65 wm diam., over entire conceptacle floor. Procarpic conceptacle unipored, 380 fm I.D. and 100 wm high, 2 carpogonial branch initials per supporting cell. Cystocarpic conceptacles uni- pored, 210-290 um I.D. and 60-90 pm high, buried throughout perithallium with spores; car- pospores 50-70 wm long and 40-70 pm diam., only on the periphery of the large discoid fusion cell (285-340 wm diam.) (Figure 9c). Male con- ceptacles unipored, raised 50-130 um above sur- face, 215-350 wm I.D. and 50-100 yum high; spermatia restricted to conceptacle floor (Figure 9c), 2-4 spermatia per mother cell, ellipsoidal to discoid 3-7 wm long, 2-3 wm diam. Type-Locarity.—Waikiki, Oahu, Hawaii (21°15’N, 157°35’W), on coral rubble, 130 m. Hototype.—D. Child, 71-50-80, 27 March 1971 (USNC), Figure 9a. PaRATYPES.—French Frigate: La Perouse, August 1971, 71-78-14. Hawai: 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. DistripuTION.— 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 Paragoniolithon 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 Neogontolithon 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 lenormandu (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, Paragoniolithon conicum, and NUMBER 15 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 Paragoniolithon conicum and Tenarea tessellatum have Caribbean “pair species,” N. rufum does not. Neogoniolithon clavacymosum, new species Ficures 10, 11s, 12 DescriptTion.—Crustae bene evolutae amplae factae, ramos breves simplicesque sparsos, qui irregulariter clavati atque saepe nodulosi fiunt (Figura 12a,B) efficientes; margines leves cras- 21 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 12s). Epithallium una duabusve cellulis crassum, cel- lulis tholiformibus, sine incrassationibus mem- branae, 2—4 wm long. atque 4-8 um diam. Mer- istema intercalare breve, 2-6 um long. atque 3-7 pum diam. Perithallium plurisstratosum, gradatim plus elongatum factum (Figura 10), corpora tinc- tabilia et fusiones adsunt, cicatrices conceptacu- lorum anticedentium visibiles, cellulis 2-10 um long. et 3-11 wm diam. Heterocystae 5-15 wm long. atque 4-11 wm diam. Hypothallium plur- istratosum, coaxiale (Figura 12c), 20-250 wm crass.; cellulae 9-28 um long. et 3-12 wm diam. Conceptacula tetrasporica uniporata, 100-320 fam super superficiem circumanbientem thalli ele- A I Depth m 0 10 20 30 40 50 60 70 80 90 xe 20 5 ; — — —_ : 5 © o . rufum > 0 ets 3 oe es >) 2 Lt B | Depth m 0 10 20 30 40 50 60 70 80 90 5 20 ec . clavacymosum 2D $ 510 2 § 4 <0 € Depth m 20 0 10 20 30 40 50 60 70 80 90 32 7 ; a T T T v T a ah N. fosliei a w = ie 3 5 oe a 5 me) 12 um LOIN) eemeeee ry eer Ares Sita ee i ea ciec ee NRE ict cee een H. reinboldiu Pink, simple irregular branches; small perithallial cells (mostly <12 fm OTC) Ae ee Sect A ewe See Ste iin ee ae ea aL H. breviclavium 3. Crust thin (<200 pm), pink-yellow, conceptacles, scattered, and dis- tinct ae ep ae ene eee H. laeve, new species Crust thicker (to several mm), irregular, pink, conceptacles abundant, small ANGENOt-GIStINCtE sayen eee oe. Hydrolithon reinboldii (Weber-van Bosse & Foslie) Foslie Ficures 14—16a Aydrolithon reinboldi (Weber-van Bosse & Foslie in Foslie) Foslie, 1909:55.—Dawson, 1954b; 1960b; 1961a.—Desi- oe ae aha H. megacystum, new species kachary and Ganesan, 1966.—Littler, 1971b; 1973a,b.— Gordon et al., 1976. Lithophyllum reinboldi Weber-van Bosse & Foslie in Foslie, 1902a:5; 1903a. Lithophyllum cerebellordes Heydrich, 1901c:405. Goniolithon reinboldii (Weber-van Bosse & Foslie in Foslie) Foslie in Weber-van Bosse and Foslie, 1904:49. 26 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Porolithon reinboldii (Weber-van Bosse & Foslie in Foslie) Lemoine, 1911:166. Description.— Thick well-developed tessellate crusts (Figure 148), 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 wm long, 6-11 wm diam. Intercalary meristem slightly elongate; cells 9-14 pm long and 5-9 pm diam. Perithallium multi- layered, irregular, fusions massive (Figure 14c) common deeper in the tissue; cells 5-20 um long and 5-17 wm diam. (Figure 15). Heterocysts not abundant, single or sometimes arranged in small groups, 12-30 pm long, 9-21 fm diam (Figure 14). Hypothallium a single layer of cells (Figure 14c,F), fusions common, cells 11-30 um long and 5-17 wm diam. Tetrasporic conceptacles uni- pored, grouped, pore plugged with mucilage, con- ceptacle reef sunken to slightly raised (20-50 pm), 200-600 pm O.D., cavities 200-340 wm I.D. and 50-160 wm high (Figure 14p); tetrasporangia re- stricted to the periphery of the conceptacle (col- umella present), 80-100 fm long and 40-75 wm diam. No sexual material sectioned. ‘Type-Loca.tity.—Mocaras reef, inner side, east coast Borneo. Ho.otypr.—Weber-van Bosse, 22 June 1899, in herbarium of M. Foslie (TRH). Isotype: USNC. DistrinuTion.—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. RemMarkKs.—A question of priority of the bas- ionym Lithophyllum cerebelloides Heydrich (190 1c) over Lithophyllum reinboldi 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. reinboldu 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- boldi 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 bgrgeseni (Foslie) Foslie is quite similar to H. reimboldi in color, surface texture, and anatomy, though it is not usually so strongly mammillate as the Hawaiian plant. Also, H. bgrgesenit 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 bgrgesenit 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 Ficures 15, 16B, 17 Hydrolithon breviclavium (Foshie) Foslie, 1909:56.—Littler, 1973b. Gonvolithon ( Hydrolithon) breviclavium Foslie, 1907b:20. Description.—Well-developed crusts with thick even margins adhering well to substrate, branched (Figure 17c) or mammillate (Figure 174,B), branches short; surface similar to Hydroli- thon reinbold:i but more finely tessellate (Figure 17), conceptacles moderately large, raised, evenly but not densely scattered over surface (Figure 17); bright to dusty pink, deep-water plants sometimes brown. Epithallium single lay- ered cells ellipsoidal, with thickened outer tan- gential wall, cells 2-10 4m long and 5-12 um diam. Intercalary meristem cells 3.5-12 zm long and 3.5-10 um diam. Perithallium multilayered, generally disorganized below third cell layer, fre- NUMBER 15 ee ‘ ae Ficure 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; FE, 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, p, 71-50-14; BE, F, 71-54-11; micrographs reduced to 91%.) CELL LENGTH (um) (um) CELL DIAMETER SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES M PERITHALLIUM ep 2 3 4 5 6 7 (oo) 14 13 10 reinboldii (7 plants) © H. breviclavium (17 plants) Ou. laeve (14 plants) o H. megacystum (5 plants) Ficure 15.—Mean epithallial and perithallial cell dimensions in H)drolithon reinboldu, H. breviclavium, H. laeve, new species, and H. megacystum, new species. NUMBER 15 29 A Depth m 10 20 30 40 50 60 70 80 90 RS — ° — = 1 = . — : eae 3 '0 H. reinboldii ae | B Depth m 10 20 30 40 50 60 70 80 90 ino Relative Abundance % breviclavium Relative Abundance % Oo 50 60 70 80 90 [o) H. Relative Abundance % ro) megacystum Ficure 16.—Depth distribution: a, Hydrolithon reinboldi; B, H. breviclavium; c, H. laeve, new species; p, H. megacystum, new species. quent wide cell fusions (Figures 15, 17F); cells 3- 18 um long and 4-13 wm diam. Heterocysts not abundant, occurring singly or in loose groups of 2 or 3, 15-36 wm long and 10-19 pm diam. Hypothallium single layer of cells (Figure 178), 6-29 um long and 5-18 pm diam. Tetrasporangial conceptacles unipored raised 375-1100 um O.D., 65-180 um high, 190-450 pm I.D.; tetrasporangia restricted to the periphery of the conceptacle, distinct columella present (Figure 17cG), 70-100 um long and 35-80 fm diam. Procarpic concep- tacles single pored, raised (80-100 ym), 250-500 um I.D., 60-130 um high, one carpogonial branch per support cell. No cystocarpic conceptacles seen. Male conceptacles uniporate, raised (50-175 zm), 300-450 um O.D., 150-200 um I.D., 25-75 um high; spermatangial branches simple, restricted to conceptacle floor; spermatia discoid, 2-4 wm diam. Type-Locauity.—Honolulu, Hawaii. Ho.otype.—Collected during Eugenie expedi- tion, in herbarium of M. Foslie (TRH). Isotype: USNC. DistripuTion.—Throughout Hawaiian Archi- pelago. SPECIMENS STUDIED.— Mau: central coast, Au- gust 1971, 71-67-4. Molokai: south-central, August 1971, 71-69-1. Midway: South Island, August 1971, 71-82-6b, 71-82-9. Oahu: Waikiki, March 1971, 71-50-26, 71-50-76, 71-50-78, 71-50-130. St. Rogatiens Bank: northwest reef, August 1971, 71- 79-14, 71-79-(35-47f). 30 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Ficure 17.—Hpydrolithon breviclavium: a, B, habit of mammillate thallus, X 1; c, habit of branching thallus, X 1; p, habit of nonmammillate thallus, X 1; £, surface of thallus showing tesselate appearance and conceptacles (arrow), X 5; F, section of male thallus, note single-layered hypothallium (arrow), X 300; G, section through a tetrasporangial conceptacle, X 200. (Speci-. men nos.: A, B, 71-50-76; c, D, 71-79-(35—47f); £, 71-50-78; F, 71-50-130; G, 71-82-6b.) RemARKS.—Procarpic conceptacles are found from those found in Neogoniolithon, where tangen- buried in the thallus unfertilized; overgrowth of tial wall projections occur in the cell lumen. In the conceptacles occurs from the roof of the con- —__-Hydrolithon thickenings occur on the outer tangen- ceptacle. tial wall, but there are no obvious projections. The epithallial cells in Hydrolithon are different Hydrolithon breviclavium is an abundant species NUMBER 15 (Figure 168) in mid to deep water (30-60 m). A few plants were found in some of the deeper dredgings at 90 m. Hydrolithon breviclavium is an important rhodolith former, but it tends to be overshadowed by Lithophyllum pallescens (Foslie) Foslie in the shallow part of its range. Hydrolithon breviclavium is strongly dominated by the melobe- sioid rhodolith formers (see ““Melobesioideae’’) on the deep banks (50-100 m). 31 We do not know of a comparable Caribbean “pair species” (see discussion under H. reinboldiz). Hydrolithon laeve, new species Ficures 15, 16c, 18 DeEscriPTION.—Crustae saepe ampla, tenues (<200 wm) (Figura 18a), superficies subtilissime rugulosa (Figura 18p), colore rosaceae ad sub- Ficure 18.—H)ydrolithon laeve, new species: a, habit of type specimen, X 12; 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, Xx 500; p, surface of type specimen, note tetrasporangial conceptacles, X 5. (Specimen nos.: A, b, 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 18p). Epithallium ex unico strato cellularum rotundatarum con- stans (Figura 18c), membrana tangentialis exte- rior incrassata, (3-7 wm long., 6-12 um diam.). Meristema intercalare paululum elongatum; cel- lulis 6-12 pm long. atque 6-10 wm diam. (Figura 18c). Perithallium pluristratosum, irregulare, fu- sionibus praesentibus; cellulis 4-17 sm long. atque 4-14 wm diam., profundiores in tela ma- lores factae (Figura 15). Heterocystae singulae (Figura 18c) (10-25 wm long., 5-20 um diam.) Hypothallium ex unico strato constans (Figura 18x), cellulis 9-25 um long. atque 5-18 wm diam. Conceptacula tetrasporangialia uniporata, ele- vata (100-120 wm) aggregata (Figura 18p,p) (600-1100 wm O.D., 220-480 wm I.D., 90-180 wm alt.); tetrasporangia ad periferiam pavimenti con- ceptaculi restricta, columella adest (Figura 18s), 50-120 wm long., 30-70 wm diam. Conceptacula procarpica uniporata, elevata (80-120 pum) 500- 650 wm O.D., 200-300 wm I.D., 60-120 um alt.; omnis cellula sustinens unicum ramum carpogon- ialem habens. Crusts often extensive, thin (<200 um) (Figure 18a), surface very finely rugulose (Figure 18p), pink to brownish; scattered large, raised conspic- uous conceptacles (Figure 18p). Epithallium a single layer of rounded cells (Figure 18c), wall thickening on outer tangential wall; cells 3-7 um long, 6-12 um diam. Intercalary meristem slightly elongate; cells 6-12 um long and 6-10 um diam. (Figure 18c). Perithallium multilayered, irregu- lar, fusions present; cells 4-17 um long and 4-14 fim diam. becoming larger deeper in the tissue (Figure 15). Heterocysts single (Figure 18c) (10- 25 wm long, 5-20 wm diam.). Hypothallium a single layer (Figure 188), cells 9-25 um long and 5-18 fm diam. Tetrasporangial conceptacles un- ipored raised (100-120 um), grouped (Figure 188, pb) (600-1100 ym O.D., 220-480 um I.D., 90-180 um high); tetrasporangia restricted to the periph- ery of the conceptacle floor, columella present (Figure 18B), 50-120 um long, 30-70 um diam. Procarpic conceptacles uniporate raised (80-120 fim), 500-650 um O.D., 200-300 wm I1.D., 60-120 uum high; | carpogonial branch per support cell. No cystocarpic or male material seen. Type-Loca.ity.—Sand Island, Midway Atoll, Hawaii (28°13’N, 177°26’W). Outside reef, 27 m depth. Ho.otyrpe.—D. Child, 71-82-27, August 1971 (USNC), Figure 18a,B,p. ParAtyPEs.— Hawai: 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. DistripuTion.— 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 reinboldit). Hydrolithon megacystum, new species Ficures 15, 16p, 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 wm long. atque 6-10 zm diam. Meristema intercalare non manifeste elongatum, intense tinctum potens (Figura 198); cellulae 5-15 wm long. atque 5-10 jum diam. Perithallium pluristratosum, fusiones frequentes (Figura 198), cellulis satis brevibus prope superficiem, profundioribus in tela, autem, progredienter longioribus factis (Figura 15) (4-16 um long., 4-15 wm diam.). Heterocystae singulae (10-27 um long., 7-15 wm diam.). Hypothallium unistratosum (Figura 198); cellulis 11-29 um | NUMBER 15 | Ficure 19.—H)ydrolithon 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.) 33 34 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES long. atque 6-15 pm diam. Conceptacula tetra- sporangialia uniporata, lata, elevata (150-200 pum), 400-800 ppm O.D., cavitates 300-440 pm I.D. atque 100-240 pm alt., paraphyses steriles adsunt; tetrasporangia trans totum pavimentum conceptaculi disposita, 75-170 fm long., 30-100 pm 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 wm long and 6- 10 wm diam. Intercalary meristem not markedly elongate, stains heavily (Figure 19B); cells 5-15 um long and 5-10 pm diam. Perithallium multi- layered, fusions common (Figure 198), cells quite short near surface but becoming progressively elongate with depth in the tissue (Figure 15) (4— 16 pm long, 4-15 wm diam.). Heterocysts single (10-27 wm long, 7-15 wm diam.). Hypothallium a single layer of cells (Figure 198); cells 11-29 pm long and 6-15 pm diam. Tetrasporangial concep- tacles uniporate, broad, raised (150-200 pm), 400-800 pm O.D., cavities 300-440 wm I.D. and 100-240 um high, sterile paraphyses present; tetrasporangia across entire floor of conceptacle, 75-170 wm long, 30-100 pm high. No sexual material seen. Type-Locauiry.— Waikiki, (21°16’N, 157°50’W), leeward. Horotype.—D. Child, 71-50-100, March 1971 (USNC), Figure 19a. ParATYPES.—Hawai: Hilo Bay, March 1971, 71-58-68. Mauz: south-central coast, August 1971, 71-66-14. Midway: South Island, August 1971, 71- 82-3. Oahu: Waikiki, March 1971, 71-50-100. DistripsuTion.— Throughout Hawaiian Archi- pelago. Remarks.—The specific epithet megacystum re- fers to the unusually large reproductive struc- tures. Hawall Oahu, Hydrolithon breviclavium, H. megacystum, and H. laeve range widely in the mid-depth zones (Figure 16B,c,p), although H/. breviclavium is twice as com- mon as the other two species. Hydrolithon laeve does not contribute to rhodolith formation, though /7. 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 Szboga ex- pedition (Weber, 1902) dredging was generally in deeper water. Some reports of HZ. 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. reiboldi 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 Ficure 20 Lithoporella melobesioides (Foslie) Foslie, 1909:59.—Lemoine, 1963.—Masaki, 1968.—Papenfuss, 1968.—Womersley and Bailey, 1970.—Gordon et al., 1976. Mastophora melobeswoides 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 20s). Epithallium a single cell layer; cells rounded to triangular (Figure 20p), 3-5 wm long, 7-8 wm diam. Perithallium present only around concep- tacles, lateral fusions common, no secondary pit connections (Figure 20p); cells 4-14 wm long and 7-11 um diam. Hypothallium single layered, fu- sions absent; cells 30-40 um long and 10-15 um diam. Tetrasporangial conceptacles unipored, highly raised, 600-1200 wm O.D. No sexual ma- terial seen. NUMBER 15 Type-Locatiry.—Maldive Islands, Nilandu Atoll, Indian Ocean. Ficure 20.—Lithoporella melobesioides: a, surface of thallus showing tetrasporangial conceptacles and “fan-like” over- growth, X 2; s, surface of thallus showing large-celled over- growth patterns, X 10; c, habit, X 1; p, section through vegetative thallus, x 10. (Specimen nos.: A—c, 71-59; p, 71- 81-16.) 30 Hovotyre.—Gardiner, 20 April 1900, 36 fathoms, in herbarium of M. Foslie, (TRH). Iso- type: USNC. DistrisuTIon.—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) Foshe. 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 Ficures 21, 22 Tenarea tessellatum (Lemoine) Littler, 1971a: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 wm in width, scattered broad but slightly raised conceptacles that leave circular depressions or are covered by the overlapping growth (Figure 21a,p). Thallus 2 cell layers; up- Figure 21.—Tenarea tessellatum: a, surface of thallus showing spiral “terraces” and tetrasporangial conceptacles, X 10; B, section through thallus showing tetrasporangial conceptacles SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES per cell layer (epithallium) triangular or rounded, cells 2-8 ym long and 5-13 ym diam.; lower cell layer (hypothallium), cells 30-50 wm long and 7- 15 um diam. Tetrasporangial conceptacles uni- pored, initially raised (90-110 wm) but quickly buried by overlapping growth (Figure 21B,pD), conceptacles 500-700 ym O.D., 210-430 um I.D., and 70-180 um high (Figure 218); tetrasporangia restricted to the periphery of the conceptacle, columella present, 50-100 pm long and 30-50 um diam. One female plant seen, cystocarpic concep- tacles, unipored, raised 400 ym O.D., 210-250 um I.D., 70-80 wm high, not buried in thallus; carposporangia restricted to periphery of large discoid fusion cell; fusion cell, 125 wm diam.; carpospores rounded, 30-35 wm diam. No male conceptacles seen. Type-Loca.ity.—Post Office Bay, Floreana (Charles) Island, Galapagos Islands. Ho.totypre.—Crossland, s.n., August 1924 (PC). DistriBuTION.—Galapagos, Hawaii, Panama. SPECIMENS StTuDIED.—Hawai: 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 7° tessellatum. It is unfortunate Lemoine (1929:68) chose this name, since confusion with the “‘tessellate sur- face” of Hydrolithon reinboldi 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 | 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; p, 71-50-89; micrographs reduced to 90%.) NUMBER 15 Relative [Petdantsoranchned: 3... 2]. 4.22 Si) 50 60 70 80 90 T. tessellatum Abundance % o_o Ficure 22.—Depth distribution of Tenarea tessellatum. Lithophyllum Philippi, 1837 Key to the Species 5 eid ercia bio ip 6.3, Nad aN aR As ie aaa 2 oy oy Seige So WE Be OG FOR Ee Oe eee 3 Plants crustose 2. Branches long, terete to flattened Urea Seeds L. kotschyanum Branches short, terete to mushroom shaped ................ L. pallescens en @nustatessellates sy on GWrustesmMoothane ee ee Soke See L. insipidum, new species 4. Crust smooth glossy, pink, medium-sized conceptacles (>300 um O.D.) .. Lithophyllum kotschyanum Unger Ficures 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.] DescriIpTIon.—Massive branching plants, sometimes with an extensive basal crust (Figure 248) 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 La een as L. ganeopsis, new species Crust smooth dull, pink, very small conceptacles (<200 um O.D.) a a RU sen ea ee L. punctatum cell layer, often absent; cells 2-4 wm long and 6- 12 um diam. Intercalary meristem generally elon- gate, forming a distinct layer; cells 5-25 um long and 7-11 pm diam. Perithallium multilayered; cells often arranged in horizontal layers of uni- form size and alignment (Figure 24£,F) elongating greatly with branching or change in direction of growth, secondary pit connections common (Fig- ure 24F), 4-24 um long and 4-10 pm 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 um long and 7- 25 um diam. Tetrasporangial conceptacles uni- pored, usually level with surface or slightly raised (occasionally raised to 60 wm), 350-450 zm O.D., 250-370 wm I.D., 80-210 um high, raised central columella present (Figure 24£); tetrasporangia SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES M PERITHALLIUM ep ] Sears a 255 Wie 10 Goel) = ac 8 ke O ay, Lu —l a 6 o L. kotschyanum (13 plants) G el. pallescens (12 plants) 5 AL. ganeopsis (4 plants) fe) BL. insipidum (5 plants) 4 2 oL. punctatum (4 plants) ® oO 3 A M PERITHALLIUM eon DEEN cin A B71 Ne | eae => $ 5 9 ae o ] uu = | A uu = << 7+ (a) —) ai 6 | Lu O 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 Ficure 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, ¢, 71-57-12; B, 71-57-2; p, 71-59-10; ©, 71-57-1; F, 71-59-8; micrographs reduced to 80%.) aN i= So Relative Abundance % (=) 10 20 30 40 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES L. kotschyanum 60 50 70 80 90 Depth m Ficure 25.—Depth distribution of Lithophyllum kotschyanum. and bisporangia restricted to the conceptacle pe- riphery, 40-100 pm long, 20-60 wm diam. No sexual material was found. Type-Locauity.—Gulf of Bahrain, Gulf. Hototype.—Kotschy, in herbarium of M. Fos- lie (TRH). Isotype: USNC. DistrisuTion.—Guam, Persian Gulf, Red Sea. SPECIMENS StTupIED.—Hawai: 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. gardinern to some extent on the algal ridges, especially along channel and groove sides. Persian Lithophyllum pallescens (Foslie) Foslie Ficures 23, 26, 27 Lithophyllum pallescens (Foslie) Foslie, 1900c:20; 1901a; 1909. —Dawson, 1944; 1954a; 1960b; 1961b.—Heydrich, 1901a.—Lemoine, 1911. Lithothamnion 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, 190 1a. Lithophyllum cephalordes Heydrich, 1901b:271. Description.—Plants initially crustose but quickly developing densely spaced small short | simple club-shaped branches (Figure 27,8); 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 wm long and 7-12 wm diam. Intercalary meristem a distinct layer; cells usually quite square but occasionally elongating individually, — 5-15 pm long and 6-11 wm diam. Perithallium — multilayered, regular with strong horizontal lay- _ ering secondary pit connections common between cells; cells 5-18 wm long and 4-15 wm diam. (Figure 23). Hypothallium single layer when pres- ent (Figure 27c); cells 6-24 um long and 6-10 wm | diam. Tetrasporangial conceptacles unipored, | usually level with surface (may be sunken (20 | um) or raised (5 wm)) 140-310 um I.D., 50-160 um high; tetrasporangia restricted to the periph- | ery of conceptacle (columella present), 30-45 um long and 15-20 wm wide. Male conceptacle uni- porate, raised (70-100 wm) 190-480 um I.D., 30- 80 wm high; spermatangial mother cells restricted _ to the conceptacle floor, 2 spermatial extensions — NUMBER 15 jo) 41 L. pallescens j=) Relative Abundance % 54 N co) w co) KR ro) 50 60 70 80 90 Depth m Ficure 26.—Depth distribution of Lithophyllum pallescens. FicureE 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 wm diam. Type-Locaity.—Sublittoral bank along the west shore of Isla Espiritu Santo, near La Paz, Gulf of California, Mexico. Hototype.—Diguet Hariot #5, in the herbar- ium of M. Foslie (TRH). Isotypes: PC, UC. DistripuTION.—Australia; Baja, California; Borneo, Gulf of California, Indian Ocean, Japan, New Guinea, Panama, Phillipines, Red Sea, Sol- omons, Vietnam. SPECIMENS StupiED.—Hawaii: Hilo, March 1971, 71-58-53; Honaunau, March 1971, 71-55- 20. Midway: lagoon, March 1971, 71-62-33. Molo- kau: 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. RemarkKs.—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- 49 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ceptacle is similar to that described for Parago- niolithon 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- clavum, 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 DescripTtion.—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 pm long. atque 6-9 zm diam. Meristema inter- calare relative breve, cellulis 6-11 wm long. et 5-— 8 wm diam. Perithallium pluristratosum, saepe regulariter stratosum (Figura 28p), foveo-colliga- tiones secondariae abundates; cellulis 4-16 wm long. et 4-12 um diam. (Figura 23). Hypothal- lium ex unico strato cellularum plerumque iso- diametricarum (7-19 ppm long., 7-14 wm diam.) constans. Conceptacula tetrasporangialia unipor- ata, paululum elevata (Figura 28c,E), 250-500 pm O.D., 140-300 pm I.D. atque 60-120 um alt., in perithallio obruta; tetrasporangia per pavi- mentum conceptaculi disposita, 40-65 um long., 20-40 pm diam. Conceptacula procarpica non visa. Conceptacula cystocarpica uniporata, ele- vata (100 wm), 230-280 pm I.D., 70-80 um alt., carposporangia ad periferiam conceptaculi res- tricta; cellula-coalescens discoidea, 150-200 um diam.; carposporae 50 fm long.; 25 wm diam. Conceptacula mascula uniporata, elevata (60 pum), 230-310 pm I.D., 40-60 pm alt.; cellulae- matricales 2 extensiones spermatiales habente; spermatia discoidea ad ellipsoidea, 3-4 zm long., 2-3 wm diam. intra fila mucosa reperta (Figura 28F). Well-developed crusts reaching a thickness of several mm, surface highly glossy (Figure 28,8); indistinct, slightly raised, randomly scattered con- ceptacles (Figure 28c). Epithallium single (rarely 2) cell layers; cells 2-5 um long and 6-9 ym diam. Intercalary meristem relatively short; cells 6-11 pum long and 5-8 wm diam. Perithallium multi- layered, often regularly layered (Figure 28p), abundant secondary pit connections; cells 4—16 um long and 4-12 diam. (Figure 23). Hypothal- lium a single layer of generally isodiametric cells (7-19 pm long, 7-14 wm diam.). Tetrasporangial conceptacles unipored, raised slightly (Figure 28c, E), 250-500 um O.D., 140-300 um I.D. and 60- 120 um high, buried in perithallium; tetraspo- rangia across floor of conceptacle, 40-65 pm long, 20-40 um diam. No procarpic conceptacles seen. Cystocarpic conceptacles uniporate, raised (100 pum), 230-280 wm I.D., 70-80 wm high; carpo- sporangia restricted to periphery of conceptacle; fusion cell discoid, 150-200 ym diam.: carpo- spores 50 wm long, 25 fm diam. Male concepta- cles uniporate raised (60 wm), 230-310 ym I.D., 40-60 um high; spermatangial mother cells re- stricted to the conceptacle floor, 2 spermatial extensions per mother cell; spermatia discoid to ellipsoidal, 3-4 wm long, 2-3 wm diam., within mucus strands (Figure 28F). Type-Locatiry.—Honauma Bay, Oahu, Ha- wail (21°15’N, 157°20’W); windward, 8 m. Ho.otyre.—D. Child 71-53-23, March 1971 (USNC), Figure 28a,.. Ficure 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, MNeTB=232 Gs ws 7il= 56-5; E, 71-81-6; F, 71-81-29.) NUMBER 15 43 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 Ficures 23, 29 DescripTion.—Crustae non amplae, satis cras- sae, autem, factae (usque ad | cm) rosaceae ad lavandulas, saepe maculas albas non perspicuas praebentes, superficie tessellata distincte sed ir- regulariter plerumque facta (Figura 298,c,F). Conceptacula plerumque abundantia, parva, paululum elevata (Figura 29F). Epithallium ex unico strato cellularum (3-6 fm long., 6-10 wm diam.) constans (Figura 29p). Meristema inter- calare perspicuum, elongatum, cellulis 11-20 long. atque 6-9 ym diam. (Figura 29p). Perithal- lium pluristratosum, foveo-colligationes secon- darias abundantes praebens (Figura 29p), bene stratosum, cellulis 4-22 wm long. atque 5-10 um diam. (Figura 23). Hypothallium unistratosum (Figura 29a); cellulis 12-16 pm long. atque 12- 19 wm diam. Conceptacula tetrasporangialia un- iporata, paululum elevata (30-70 wm), 450-750 pum O.D., 170-230 wm I.D., 80-140 wm alt. (Fig- ura 29a); tetrasporangia ad periferiam concepta- culi restricta, columella adest, 40-100 um long., 20-50 wm 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 298, c,F); conceptacles usually abundant, small, slightly raised (Figure 29F). Epithallium a single layer of cells (3-6 wm long, 6-10 pm diam.) (Figure 29p). Intercalary meristem distinct, elon- gate; cells 11-20 pm long and 6-9 pm diam. (Figure 29p). Perithallium, multilayered, with abundant secondary pit connections (Figure 29p), well layered; cells 4-22 um long and 5-10 pm diam. (Figure 23). Hypothallium a single cell layer (Figure 29a); cells 12-16 wm long and 12- 19 wm diam. Tetrasporangial conceptacles uni- | pored, slightly raised (30-70 wm), 450-750 ym O.D., 170-230 pm I.D., 80-140 um high (Figure | 29c); tetrasporangia restricted to the conceptacle periphery, columella present, 40-100 um long, | 20-50 pm diam. Sexual plants not found. | Type-Locatity.—Waikiki, Oahu, Hawaii (21°10’N, 157°55’W), leeward, 3-5 m depth. Hototype.—D. Child, 71-50-17, March 1971 (USNC), Figure 29s. 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 H)drolithon, L. insipidium, with its | lavender cclor and tessellate surface, may be con- | fused with Hydrolithon reinboldi. Thicker plants with the color and texture of H. reinboldi that are lacking mammillons or with relatively small con- | ceptacles should be investigated closely to deter- | mine whether they are H. reznboldw or L. instpidum. Examination of a fractured vertical surface for | the irregular, almost porous perithallium (due to | extensive fusions) or H. reiboldi should easily NUMBER Jat sa ” eee ft y ts Paes apr onte te By FicurE 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; £, 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; p, 71-82-32; ©, 71-58-39; micrographs reduced to 93%.) 46 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES a EH Ficure 30.—Lithophyllum punctatum: a, habit, X 1; 8, 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; p, 71-53-32; £, 71-50-7; micrographs reduced to 83%.) NUMBER 15 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 Ficures 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 um long and 6-12 pm diam. Intercalary meristem of ran- domly elongating cells (5-9 um long, 6-8 wm diam.). Perithallium multilayered, secondary pit connections present, scattered large cells at sur- face (16-18 um long, 10-11 wm 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 wm long, 4-10 wm diam.) (Figure 23). Hypothallium a single cell layer; cells isodia- metric to horizontally elongate, 7-16 um long and 6-10 wm diam. (Figure 30E). Tetrasporangial conceptacles unipored, raised 40-50 wm above the surrounding crust, 160-170 wm I.D., 60-70 um high (Figure 30p); tetrasporangia restricted to periphery of conceptacle, columella present, 40-80 ppm long and 20-65 pum diam. No sexual material collected. Type-Loca.ity.—Tricomalie, Ceylon. Ho.otype.—Svedelius, 17 April 1903, in her- barium of M. Foslie (TRH). Isotype: USNC. DisrripuTIon.—Ceylon, the island of Oahu. SPECIMENS STUDIED.— Oahu: Honauma, March 1971, 71-53-32; Kaneohe, March 1971, 71-54-8; 47 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- othammum 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 Archaeolithothammum, 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) aig fondle She ae ee 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). Archaeolithothamnium erythraeum (Rothpletz) Foslie Ficures 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 314) bright bluish red Cover cells “Lithothamnum-type” (Figure 3lc,p) forming a single layer over the epithallium; cells 2-7 um long and 5-10 um diam. Intercalary meristematic cells 5-8 um long and 5-11 wm diam. Perithal- lium multilayered regular, lateral cell fusions and secondary pit connections present (Figure 318); cells square, tending to elongate and enlarge with depth in tissue (Figure 32), 5-12 um long and 4- 15 wm diam. Hypothallium multilayered, parallel to weakly coaxial, 30-110 um thick; cells 13-34 tum long and 6-9 um diam. Tetrasporangial sori to brown. Sine el ecalegs A. episoredion, new species buried in tissue (Figure 31£), surface sori not seen in collections; tetrasporangia 60-90 wm long and 25-40 wm diam. Type-Locatity.—Red Sea. Ho.otrype.—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 Hpdrolithon and Sporolithon “communities” may include, or even be domi- if nated by, Neogoniolithon species. NUMBER 15 ») Ficure 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 “Lithothamnium-type” cover cell, X 7500; pb, light micrograph of “Lithothamnium-type” cover cell, X 2000; £, thallus with buried sorus (arrow), X 20. (Specimen nos.: A, 71-82-69; B, c, 71-53-24; p, £, 71-82-63; micrographs reduced to 95%.) 49 50 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES PERITHALLIUM ep 1 2 3 4 5 (um) ag (i 2) Ze Ww ed | —_ uJ (S) § 9 a & _ Ww Loe 8 UO = 500 pm, O.D. ae hl aera aN cae Le L. pulchrum Thin, bright-red crusts developing abundant, narrow often irregular branches, 0.5-2 mm diam.; conceptacles < 500 um, O.D. Lithothamnium pulchrum Weber-van Bosse & Foslie Ficures 36-38 Lithothamnum 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 pm thick) with large raised mul- tipored conceptacles; developing numerous knobs and eventually large irregular branches 2-5 mm diam. (Figure 36a,B). Cover cells “Lithothamnium- type,” single cell layer; cells 1-4 um long and 3- 10 wm diam. Intercalary meristem medium length; cells 3-6 wm long and 5-10 pm diam. Perithallium multilayered, abundant fusions, scattered staining bodies (phosphotungstic hae- matoxylin) (Figure 36c); cells 3-9 um long and 9-22 um diam. (Figure 37). Hypothallium mul- tilayered, noncoaxial, 20-65 um thick; cells 3-11 um long and 4-11 pm diam. (Figure 36c). Tet- rasporangial conceptacles multipored, strongly raised (Figure 36£) 500-1000 um O.D., 360-470 um I.D., 170-250 um high; tetrasporangia not seen. Sexual conceptacles seen, but not sectioned (Figure 36D). Type-Loca.ity.—Sailus Besar, Paternoster Is- lands, Indonesia (station 315, Szboga Expedition), 36 m depth. Ho.otype.—Weber-van Bosse, 19-21 Febru- ary 1900, in herbarium of M. Foslie (TRH). Isotype: USNC. DistRiBUTION.—Celebes. SPECIMENS StTuDIED.—Hawai: Honaunau, Cea Aa ee aaa nia ta L. australe 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 IT, Wilea(@-8), leak, REeMARKS.—Next to Lithothammum 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 Lzthothamnium 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 Ficures 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 corallioides 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,p,£); con- ceptacles medium-sized, rather rounded, flat- 54 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES | Ficure 36.—Lithothamnium pulchrum: a, branch of thallus, X 20; B, habit, X 1; c, section through vegetative crust, X 150; p, sexual crust with conceptacles (arrow), x 10; £, tetrasporangial conceptacles (arrow), X 5. (Specimen nos.: A, B, 71—79- 9; c, 71-79-11; p, 71-68-14; E, 71-67-1; micrographs reduced to 94%.) | | NUMBER 15 55 PERT ASE OI ep 1 2 8 4 5 6 Uf 8 9 eS 5) 8 2S — oN Ww aa ee ee WwW UO ae 5 Da OL. pulchrum (13 plants) ‘ o L. australe (10 plants) a — fav WwW (—. LW = < oe Aone oe ae = Ww Si 5) Ficure 37.—Perithallial cell dimensions of Lithothamnium pulchrum and L. australe. Depthm 10 20 30 40 50 60 70 80 90 SR | L. pulchrum Relative Abundance Ficure 38.—Depth distribution of Lithothamnium pulchrum. on D SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Ficure 39.—Lithothamnium australe: a, habit, X 1; B, section through epithallium showing “Lithothamnium-type”’ cover cells, X 2200; c, section through thallus; note meristem (arrow) and perithallium, X 600; p, tetrasporangial conceptacles on surface of thallus (arrow), X 5; £, branch morphology, X 10. (Specimen nos.: A, D, E, 71-73; B, c, 71-72-1; micrographs reduced to 93%.) angen NUMBER 15 Relative Abundance % ND oO L.australe Ficure 40.—Depth distribution of Lithothamnium australe. topped multipored domes or conspicuous single- pored, pointed cones, often found on branch tips. Cover cells “Lithothamnium-type” (Figure 39s), single layer; cells 2-3 um long and 3-12 pm diam. Intercalary meristem short and wide; cells 4-6 uum long and 5-10 wm 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 wm long and 3-11 pm diam. Hypothallium thin, parallel to substrate, 30-55 um thick; cells 11-24 um long and 4-10 pm diam. Tetrasporic conceptacles slightly raised, 300-500 um O.D. (none seen in section). Male concepta- cles unipored raised (50-150 pm), 300-450 ym O.D., 250-350 pm I.D., 90-110 pm high; sper- matangial mother cells dendroid, restricted to floor of conceptacle cavity; spermatia discoid, 1- 3 pm diam. Type-Loca.ity.—Gulf of California: La Paz, Baja California del Sur. Lecrotype.—Diguet (Foslie, 1895, fig. 7), in the herbarium of M. Foslie (TRH). Lecto-isotype: USNC. DistrrBuTION.—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 Lithothamniwm australe f. americana: ~“Vhis form constitutes the basis of Lithothamnion 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. Lithothamnum 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 RemMARKS.— 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) ............. cbyo cgay 3 Beato Bee cy ge Ue 108 Nee Ae M. madagascariensis Crustose yo. sei eo 6) Saag lan ee ae 2 2. Crusts extensive smooth, sub-leafy; conceptacles > 600 wm O.D. ....... 3 Crusts somewhat irregular; conceptacles < 600 um O.D. ............... 4 3. Few, scattered'conceptacles)=.800)uimml@ 1D eee M. prolifer Densely concentrated mass of conceptacles, 600-800 um O.D. re es Ake Si M. purpurascens 4. Crusts irregular knobby, densely coated with raised conceptacles ......... fee Fee M. syrphetodes, new species Crusts not grossly irregular, conceptacles scattered ....................... Mesophyllum madagascariensis (Foslie) Adey Ficures 41a—c, 42 Mesophyllum madagascariensis (Foslie) Adey, 1970:25. Lithothamnium erubescens Foslie f. madagascariensis Foslie, 1902b:3.—Masaki, 1968. Lithothammum 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 41,8); large domed conceptables, more abun- dant in sparsely branched crusts. Epithallium single, rounded to quite angular; cells 2-4 um aye Cue ere: M. fluatum, new species long and 4-9 wm diam. Intercalary meristem elongate; cells oval, 4-10 wm long and 3-9 um diam. Perithallium multilayered, thin when thal- lus crustose, staining bodies present, fusions fre- quent; cells 3-12 pm long and 3-9 um diam. (Figure 42). Heterocysts abundant, single or in small groups, 7-19 wm long and 7-18 um diam. Tetrasporangial conceptacles multipored, few ini- tially raised (80 wm) but soon becoming buried, walls of sterile cells remaining (Figure 41c), 140- 210 um I.D., 60-90 pm high; tetrasporangia and bisporangia present, 60-90 um long and 35-50 jum diam. Male conceptacles unipored (1 plant sectioned) 300-400 um O.D., 140-208 um L.D., 50-100 um high; spermatangial mother cells sim- ple, columnar, restricted to the conceptacle floor; NUMBER 15 Ficure 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: p, habit of typical specimen, X 2; EF, 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).) 2) 60 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES M PERITHALLIUM ees AG 7 8 GEEE- EE N'G71 a (um) 2 O M. madagascarensis (5 plants) o M. syrphetodes (9 plants) 1S) CELL DIAMETER (um) Ficure 42.—Perithallial cell dimensions of Mesophyllum madagascariensis and M. syrphetodes, new species. spermatia, ellipsoidal, in mucus strands, 2-3 fm Distripution.—Ceylon, Madagascar, Japan. long and 1-2 ym diam. SPECIMENS StupiED.—Hawai: Kawaihae, Type-Locatitry.—Madagascar. March 1971, 71-57-17. Oahu: Honauma, March Hototype.—In herbarium of M. Foslie 1971, 71-53-21; Kaneohe, August 1971, 71-81-9, (TRH). Isotype: USNC. 71-81-31, 7128 1-35: NUMBER 15 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- wail 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, 1.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 FicureE 41D,E Mesophyllum prolifer (Foslie in Weber-van Bosse & Foslie) Adey, 1970:25. Lithothamnum 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 41p). Epithallium single layer of rounded cells, 3—4 um long and 4-7 pm diam. Intercalary meri- stem elongate; cells 4-5 wm long and 5-6 pm diam. Perithallium multilayered, irregular; cells somewhat square, 2-10 um long and 3-9 pm diam. Hypothallium weakly coaxial 20-60 fm thick; cells 7-27 wm long and 5-12 pm diam. Tetrasporangial conceptacles multipored, low and rounded domes, epithallium persistent, 800- 61 1200 um O.D. (none sectioned). Sexual concep- tacles single pored sharply raised (Figure 41e), none sectioned. Type-Loca.tity.—Pulu Sebangkatan, Borneo Bank, Borneo; 34 m depth, coral bottom and Lithothamnium. Lectrotype.—Weber-van Bosse, 971, 14 June 1899, in herbarium of M. Foslie (TRH) (Adey, 1970). DistrrBuTION.— Borneo. SPECIMENS STUDIED.— Mawi: 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 Ficure 43 Mesophyllum purpurascens (Foslie) Adey, 1970:26. Lithothamnium funafutiense Foslie f. purpurascens Foslie, 1901b:18. Lithothamnum purpurascens Foslie, 1907a:182; moine, 1917; 1965.—Papenfuss, 1968. 1929.—Le- DescripTion.—Crusts smooth thin and glossy (Figure 43a). Epithallium a single layer of rounded cells, 2-3 wm long and 5-8 fm diam. Intercalary meristem elongate; cells 6-8 zm long and 4-6 pm diam. Perithallium multilayered, zoned from formation of conceptacles; cells 4-8 um long and 3-6 wm diam. (Figure 438). Hypo- thallium multilayered, coaxial; cells 10-16 um long and 3-4 wm diam. (Figure 43£). Tetraspor- angial conceptacles centrally crowded (Figure 43a), raised (150 wm) (Figure 43p), becoming buried (Figure 438), 500-900 wm O.D., 300-700 um 1.D., 180-270 um high, walls of sterile cells remaining in cavity (Figure 43B,c); tetrasporan- gia not seen. Type-Locatity.—Koh Mesan and Cape Liant, Gulf of Thailand, 18 m depth. Lectotype.—Danish expedition to Thailand, 62 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ihe Tel Ficure 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; p, surface view of tetrasporangial concep- tacles, X 20; £, coaxial hypothallium, X 1000. (Specimen nos.: A—E, 71-66-2.) NUMBER 15 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 Ficures 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 44<,c). Epithallium 1-2 cellulis crassum, cellula exteriore cellulam-obtegentem interdum efficiente; cellu- lae rotundatae 2—4 um long. atque 4-8 wm diam. Meristema intercalare non manifeste elongatum; cellulis 3-7 zm long. atque 3-5 fm 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 um long. atque 3-8 um diam. (Figurae 44c, 42). Hypothallium pluristratosum, paral- lelum ad coaxiale, 20-75 wm crass.; cellulae 10- 20 wm long., 5-9 wm diam. Conceptacula tetra- sporangialia multiporata, crebre disposita (Figura 448) in quibusdam plantae regionibus repetite evenientia, frequenter obtecta facta et crassitu- dinem plantae augentia (Figura 44c), elevata (80-180 wm), tholiformia (Figura 44B,c), 250-400 pum O.D., 150-300 pm 1.D., 70-120 pm alt.; tetra- sporangia 80-90 tum long. atque 25-45 um 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 wm long and 4-8 wm diam. Inter- calary meristem not markedly elongate; cells 3-7 63 pum long and 3-5 wm 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 um long and 3-8 wm diam. (Figures 44c, 42). Hypothallium multilayered, parallel to coaxial, 20-75 pm thick; cells 10-20 4m long 5-9 pm diam. Tetrasporangial conceptacles multipored, densely spaced, (Figure 448) occurring repeatedly in certain areas of the plant, frequently becoming overgrown and adding to thickness of plant (Fig- ure 44c), raised (80-180 zm), domed (Figure 44.8, c), 250-400 wm O.D., 150-300 wm I1.D., 70-120 tum high; tetrasporangia 80-90 4m long and 25- 45 um diam., bisporangia present (Figure 44c). Sexual plants not seen. Type-Loca.iry.—Southwest Molokai, Hawaii (21°0O’N, 157°0’W), 70 m depth. Hovotype.—D. Child, 71-72-2, August 1971 (USNC), Figure 44a. Paratypes.— Maw: 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. Distripution.— 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 fluatum, new species Ficure 45 DEscRIPTION.—Crustae tenues rosaceae (25- 100 wm), super superficiem “profluens,” et non praesertim rugatae neque substratum plane illus- trantes (Figura 45a,B). Epithallium ex unico strato cellularum rotundatarum, 3-4 fm long. atque 3-5 ppm diam. constans. Meristema inter- calare non elongatum, cellulis 2-6 wm long. atque 4-7 um diam. Perithallium pluristratosum, cel- lulis 4-7 wm long. atque 4-12 fm diam. Hypo- coaxiale, 20-25 pm crass.; cellulis 10-17 wm long. atque 4—5 um crass. thallium pluristratosum 64 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Ficure 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 Ficure 45.—Mesophyllum fluatum, 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, super superficiem dispersa (Figura 45), elevata (50-100 um) 300-500 wm O.D. (solum a superficie visa). Conceptacula carposporangialia uniporata, (solo uno secto) 525 wm O.D., 350 wm I.D., 150 um alt.; carposporangia ad periferiam concepta- culi restricta; sporangia 45-50 wm long. atque 60-75 pm diam.; cellula-coalescens reticulata. 66 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Thin pink crusts (25-100 pm), “flowing” over the surface and not particularly crumpled or strongly reflecting the substrate (Figure 45a,B). Epithallium single layer of rounded cells, 3-4 um long and 3-5 4m diam. Intercalary meristem not elongate; cells 2-6 fm long and 4-7 pm diam. Perithallium, multilayered; cells 4-7 um long and 4—12 um diam. Hypothallium multilayered coax- lal, 20-25 um thick; cells 10-17 wm long and 4-5 uum diam. Tetrasporangial conceptacles multi- pored, scattered over surface (Figure 45s), raised (50-100 zm), 300-500 um O.D. (only seen from surface). Carposporangial conceptacles unipored, (only | sectioned) 525 wm O.D., 350 ym 1.D., 150 um high; carposporangia restricted to the concep- tacle periphery; sporangia 45-50 wm long and 60-75 ym diam.; fusion cell reticulate. Type-Locatity.—South-central Maui-Hawaii, 92 m. Horotype.—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. Nthoa: 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; Lithothammum, Mesophyllum, and Archaeolithotham- num 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. gardinern, 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, 1973), 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 HZ. reinboldi 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 Paragoniolithonf Porolithon e Hydrolithon CARIBBEAN SEA (ee) oO (METERS) MASTOPHOROIDEAE DEPTH oO Porolithon§ 20: Neogoniolithon Hydrolithon 60 c fe) aS sz & iS ° D ie) = 5 a HAWAIIAN ARCHIPELAGO 80 67 E 5 = te om ° ae s= —_! Mesophyllum Archaeolithothamnium Lithothamnium -LITHOPHYLLOIDEAE= ———MELOBESIOIDEAE INTERTIDAL gf Lithophyllum Archaeolithothamnium Mesophy!lum Lithothamnium BREGIWENIGY Ol OG GURNEE NIGE: 028 Ficure 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 Lzthothamnium and Mesophyllum species were not present. Doty (1974) reports abundant Hydrolithon and Archaeolitho- thammum 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 gardinerui and Neogonvolithon 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 Fydrolithon, 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 6£, 246). Coralgal. A calcareous substrate consisting of coral and corallines or other calcareous algae. Ficure 47.—Schematic representation of the conceptacle. 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, 29p, 13c). “Lithothammum-type” cover cells. of Lithothamnium (Figure 31c, D). High (conceptacle). Measurement C, Figure 47. I.D. (inside diameter, conceptacle). Characteristic 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, 29p). 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 “leaf” 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). External widest di- ameter of a conceptacle when viewed from the surface, measurement A, Figure 47. O.D. (outside diameter, conceptacle). Parallel (hypothallium). Tissue where the filaments run in line with the substrate and are not coaxial. See “Simple” (Figure 8s). 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). 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University of Glas- gow: International Phycological Society (19-22 Aug 1980). ne te 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, 5/, 52, 66 episporum, 51 erythraeum, #6, 49, 50, 66 foslier, 23 howel, 51 pacificum, 51 australe, Lithophyllum, 53 Lithothamnium, 33, 55-58 Mesophyllum, 53 australis, Lithothamnium coralloides f., 53, Si ' bgrgensenti, Hydrolithon, 26 breviclavium, Gonzolithon, 26 Hydrolithon, 26, 28-32, 34, 42 californiense, Lithophyllum, 40 caribaeum, Neogoniolithon, 16 cephaloides, Lithophyllum, 40 cerebellordes, Lithophyllum, 25, 26 clavacymosum, Neogoniolithon, 19, 2/, ds 28) congestum, Lithophyllum, 10, 12, 40 conicum, Hydrolithon, 13, 34 Neogoniolithon, 13 Paragoniolithon, /3, 14, 15, 20, 21, ey AP dimotum, Archaeolithothamnium, 48 epilaeve, Kvaleya, 25 episoredion, Archaeolithothamnium, 48, 50, 5/, 52, 66 episorum, Archaeolithothamnium, 51 eplyessoense, Ezo, 25 erubescens, Mesophyllum, 61 erythraeum, 48, 49, 50, 66 Lithothamnium, 48 Sporolithon, 48 Ezo epipyessoense, 25 Archaeolithothamnium, 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, /0, 11, 12, 40, 66, 67 Goniolithon breviclavium, 26 fosler, 23 oncodes, 7 pallescens, 40 propinguum, 12 reinboldi, 25 solubile, 12 tessellatum, 35 howei, Archaeolithothamnium, 51 Hydrolithon, 6, 25, 26, 30, 34, 44, 48, 58, 67, 68 beérgensenil, 26 breviclavium, 25, 26, 28-32, 34, 42 conicum, 13 laeve, 25, 28, 29, 3/, 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, 3/, 32, 34 lenormandii, Phymatolithon, 20 lichenoides, Mesophyllum, 25 Lithophyllum, 6, 7, 37, 44, 47, 67, 68 australe, 53 californiense, 40 73 cephalordes, 40 cerebelloides, 25, 26 congestum, 10, 12, 40 fosler, 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 reinboldi, 25, 26 tessellatum, 35 Lithoporella, 6, 34, 35, 67 melobesioides, 6, 34, 35 Lithothamniun,, 6, 7, 15, 47, 48, 52, 53, 58, 66-68 australe, 53, 55-58 australe f. americana, 33, 57 coralloides £. australis, 33, 57 erubescens {. madagascariensis, 38 erythraeum, 48 foslier, 23 funafutiense f. purpurascens, 61 kotschyanum, 37 madagascariensis, 38 occidentale, 58 onkodes, 7 pallescens, 40 prolvfer, 61 pulchrum, 52, 53, 54, 55 purpurascens, 61 madagascariensis, Lithothamnium, 58 Mesophyllum, 58, 59, 60 Lithothamnium erubescens f., 58 mammillare, Neogoniolithon, 26 Mastophora melobeswoides, 34 “megacarpum, Neogoniolithon, 25 megacystum, Hydrolithon, 28, 29, 32, 335 34 melobesioides, Lithoporella, 6, 34, 35 Mastophora, 34 74 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, 6/ pulchrum, 53 purpurascens, 58, 6/, 64 siamense, 5 syrphetodes, 58, 60, 63, 64 Millepora, 10 Neogoniolithon, 6, /5, 25, 30, 48, 66-68 accretum, 20 caribaeum, 16 clavacymosum, 15, 19, 2/, 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, /6, 17 solubile, 12 nitorum, Lithophyllum, 44 occidentale, Lithothamnium, 58° okamurat, Lithophyllum, 40, 41 oncodes, Goniolithon, 7 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Lithophyllum, 7 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, Gonzolithon, 40 Lithophyllum, 31, 37, 38, 40, 41, 42 Lithothamnium, 40 patena, Synarthrophyton, 20, 61 Paragoniolithon, 6, /2, 15, 67 conicum, /3, 14, 15, 20, 21, 23, 42 solubile, 12, 15 “typica,” 15 Phymatolithon lenormandi, 20 rugulosum, 61 Porolithon, 6, 7, 12, 66 gardineri, 7, 9, /0, 11, 12, 40, 66, 67 oncodes, 7, 23 onkodes, 3, 7, 8-10, 20, 21, 23, 66 pachydermum, 10 reinboldir, 26 prolifer, Lzthothamnium, 61 Mesophyllum, 58, 59, 6/ 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 f., 61 Mesophyllum, 58, 6/7, 62 reinboldii, Gonzolithon, 25 Hydrolithon, 25, 26-29, 31, 32, 34, 36, 44, 66 ; Lithophyllum, 25, 26 Porolithon, 26 rufum, Neogoniolithon, 15, /6, 18-21 rugulosum, Neogoniolithon, 15, /6, 17 Phymatolithon, 61 siamense, Mesophyllum, 5 solubile, Gonzolithon, 12 Neogonvolithon, 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 Gonvolithon, 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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