occasional papers of the Farlow. Herbarium of cryptogamic botany No. 12 April, 1977 Harvard University, Cambridge, Massachusetts Gayle |. Hansen Cirrulicarpus Carolinensis, A New Species in the Kallymeniaceae (Rhodophyta) A Comparison of the Species of Cirrulicarpus (Kallymeniaceae, Rhodophyta) Monte G. Manuel | Studies in Cryphaeaceae III. Sphaerotheciella Fleisch. New to the Americas Edited by: Reed C. Rollins Kathryn Roby occasional papers of the Farlow. HerbDariumM of cryptogamic botany No. 1. No. 2 No. 3 No. 4. No. 5 No. 6 No. 7 No. 8 No. 9 No. 10. Sylvia A. Earle: Hummbrella, a New Red Alga of Uncertain Tax- onomic Position from the Juan Fernandez Islands (June 1969). . I. Mackenzie Lamb: Stereocau/on arenarium (Sav.) M. Lamb, a Hitherto Overlooked Boreal-Arctic Lichen (June 1972). . Sylvia A. Earle and Joyce Redemsky Young: S/phonoclathrus, a New Genus of Chlorophyta (Siphonales: Codiaceae) from Panama (July 1972). 1. Mackenzie Lamb, William A. Weber, H. Martin Jahns, Siegfried Huneck: Calathaspis, a New Genus of the Lichen Family Cladonia- ceae (July 1972). . |. Mackenzie Lamb: Stereocau/on sterile (Sav.) M. Lamb and Stereocaulon groenlandicum (Dahl) M. Lamb, Two More Hitherto Overlooked Lichen Species (March 1973). . I. Mackenzie Lamb: Further Observations on Verrucaria serpu- loides M. Lamb, the Only Known Permanently Submerged Marine Lichen (April 1973). . Bruce H. Tiffney and Elso S. Barghoorn: The Fossil Record of the Fungi (June 1974). . Donald H. Pfister: The Genus Acervus (Ascomycetes, Pezizales). |. An Emendation. II. The Apothecial Ontogeny of Acervus flavidus with Comments on A. epispartius (May 1975). . Donald H. Pfister: A Synopsis of the Genus Pulvinula. A New Combination in the Genus Gymnomyces. Norton G. Miller: Studies on North American Quaternary Bryophyte Subfossils. |. A New Moss Assemblage from the Two Creeks Forest Bed of Wisconsin (July 1976). Emmanuel Sérusiaux: Some Foliicolous Lichens from the Farlow Herbarium (August 1976). Continued on back cover CIRRULICARPUS CAROLINENSIS, A NEW SPECIES IN THE KALLYMENIACEAE (RHODOPHYTA ) GAYLE I, HANSEN! ABSTRACT Cirrulicarpus carolinensis sp. noy. is a cartilaginous, complanate, subdichotomously branched, red alga which inhabits the submerged rocky reefs off the North Carolina coast. The plant is perennial and grows by means of a marginal, surface meristem. In section, the blades consist of a medulla of primary stellate and secondary filiform cells and a cortex of large, subglobose inner cells that grade into smaller, nearly globose outer cells. The carpogonial branch apparatus consists of one or two carpo- gonial branches of three cells and occasionally up to two subsidiary cells borne on a large, lobed supporting cell, Carposporophyte development is distinctive in that the gonimoblast filaments appear to form from vegetative instead of reproductive cells. The life history consists of an alternation of gametophyte and carposporophyte generations. INTRODUCTION A new species of Cirrulicarpus was found growing in crevices of the rocky reefs off the New River Inlet in North Carolina. In habit the alga resembles Cryptonemia crenulata (J. Ag.) J. Ag., a species to which it has been incorrectly referred in carlier papers (Hoyt, 1920; Pearse and Williams, 1951). However, unlike Cryptonemia, the new alga has carpogonial branches of three cells borne on a large, lobed supporting cell and very large cystocarps. It is a member of the family Kallymeni- aceae and most closely resembles other species of the genus Cirrulicarpus, a genus that has not previously been reported from the Atlantic Ocean. A detailed examination of the North Carolina species was undertaken in order to confirm its taxonomic affinities and to understand its biology. The new name Cirrulicarpus carolinensis is proposed. Cirrulicarpus carolinensis Hansen, sp. nov. Plantae perennes, caespitosac, ad 16 cm altae. Laminae juniores ob- ovoideae integrae, maturae subdichotome ramosae ramis inferis anguste cuneatis, ramis superis obovoideis lobatis. Thallus ex cellulis corticeis parvis (4-8 pm lat.), ex cellulis subcorticeis majoribus (12-40 pm lat. ), et medulla cellularum stellatarum primariarum, secundariarum filiform- ium sonstans. Dioeciae. Plantae femineae fasciculis ramulorum carpo- gonialium praeditae; omnis fasciculus ex 1 vel 2 ramulis tricellularibus carpogonalibus et 0-2 cellulis subsidiariis e cellula magna exorientibus constans. Carposporophyta laxa; filamenta gonimoblastorum ex filamentis cellulorum brevium constantia; filamenta ec cellulis vegetativis exorientia, in cellulis vegetativas panetrantia, et cum cellulis per foveas conjugantia. Cystocarpi non ostiolati, magni (ad 3 mm lat.), fasciculati in superficie frondis, in aetate provecta verrucosi. Plantae masculinae soros sperma- 1Present address: Farlow Library and Herbarium of Cryptogamic Botany, 20 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. 1 2 GAYLE I. HANSEN tangiorum in pagina thalli irregulatin dispersos efferentes. Spermatangia ex cellulis elongatis vallum supra corticum facientibus evoluta. Tetra- sporangia ignota, type. North Carolina. Onslow County: ca. 3.2 km east of New River Inlet at Mile Hammock Rock, depth ca. 5 m, Hansen, 10 October, 1974. HOLOTYPE: NCU. Isotypes to be distributed to: AHFH,BM,DUKE,FH,LD,MEL,PC,UC,WTU, herb. C, Schneider (Trinity College, Connecticut), herb. T. Masaki (Hokkaido University, Japan). Plants caespitose. Blades entire and obovoid when young, branching subdichotomous when mature; lower branches narrowly cuneate and upper branches obovoid-lobed. Perennial, reaching 16 cm in height. Thallus composed of small cortical cells (4-8 »m in diameter ), larger subcortical cells (12-40 jam in diameter), and a medulla of primary stellate and secondary filiform cells. Dioecious. In female plants, the carpogonial branch systems consisting of one or two three-celled carpo- gonial branches and zero to two subsidiary cells on a large supporting cell. Carposporophytes diffuse; gonimoblast filaments consisting of short- celled filaments which originate from, form pit connections with, and penetrate vegetative cells. Cystocarps non-ostiolate, large (to 3 mm in diameter), arranged in clusters on the upper surfaces of the fronds, and becoming verrucose when old. In male plants spermatangial sori scattered irregularly over the thallus surface. Elongate spermatangial mother cells forming a palisade-like layer above the cortex. Tetrasporangial plants unknown. Material and Methods. Collections of Cirrulicarpus carolinensis were made about every other month over a three year period from 1972 to 1974. Drift material was collected from the south side of the New River Inlet, North Carolina, and attached material was collected by SCUBA from Mile Hammock Rock, a submerged reef located near the Inlet. Freshly collected material was preserved in 5% formalin in seawater or Carnoy’s fixative containing chloroform (Johansen, 1940). The material was prepared for staining by the following methods: (1) hand sectioning with a razor blade; (2) sectioning with a freezing microtome; (3) squash- ing small pieces under a coverslip; or (4) peeling the upper and lower cortex of the thallus apart with fine needles and placing the inner faces of the cortex upwards on a slide. Material fixed in formalin required no g in alkali (Norris, 1957) or acid (Papenfuss, 1937) before squashing or peeling. These preparations were stained with 1% aniline blue according to the procedure of Papenfuss (1937), except that a 5% hydrochloric acid solution was used to bind the stain. The material was then mounted in Karo syrup containing phenol (Womersley and Norris, 1971). A 35 mm Nikon camera attached to either a Wild M-5 dissecting microscope or a Zeiss RA light microscope was used for microphotog- presoftenin raphy, and a Pentax Spotmatic camera was used for macrophotography. CIRRULICARPUS CAROLINENSIS 3 Habitat. Cirrulicarpus carolinensis inhabits a submerged rocky outcrop located about 3.2 km east of the New River Inlet in Worth Carolina. This site, called Mile Hammock Rock by local fishermen, appears to be a part of the “Black Rock” reefs studied by Pearse and Williams (1951). These reefs are an outcropping of Lower Miocene Trent Marl. Covered with algae, the rocks attract the blackfish, Centropristes striates (L.) Mile Hammock Rock rises abruptly from the sandy sea Hone off the Inlet to about 4.0 m below the surface. The rocks are traversed by many deep, narrow fissures, and are undercut with shallow caves. Cirrulicarpus grows in irregularly scattered clumps along the vertical surfaces of the sheltered and shaded walls of the fissures. The fronds are frequently found attached to sea squirts, but growth on the porous rock alone is also quite common. The alga grows at a depth of from 4.5 to 9.0 m below the surface and is perennii al. Juvenile plants are seen infrequently throughout the year and are never found in abundance in any particular season. Based on bimonthly observations, the major growing season for C. carolinensis appears to be from late spring to late fall. Growth is arrested in the winter. During the summer, C. carolinensis dominates the red algal flora at Mile Hammock Rock because most other species have died back to their holdfasts. The fall and spring blooms grade into one another over the winter, and during this time C. carolinensis is a minor component of the flora. Vegetative Structure. Young thalli of Cirrulicarpus carolinensis are light pink, cartilaginous, complanate, and obovate with margins that are entire and somewhat undulate (Fig. 1). The blades are attached to the substratum by a short terete stipe and a small discoid holdfast. As the fronds grow, they branch subdichotomously (Fig. 2). As many as ten Fic. 1, 2. Young and mature plants of Cirrulicarpus carolinensis. Notice the thinner, more trans- parent quality of the current year’s growth; scale, 3 cm. 4 GAYLE I, HANSEN dichotomies were counted in mature plants 16 cm in height. Dichotomies in the older portions of the plant are acute; those near the apex are somewhat rounded. The thallus is broadest near the rounded tips of the ultimate branches and may reach a width of 2 cm. Mature plants are widely foliose and exceedingly undulate and appear in habit like tufts of crowded ruffles. The alga is perennial and, on the basis of morphological observations, appears to live for from two to four years. During the winter the fronds deepen in color, thicken, and become lacerated and worn. At this stage the plants are often narrowly cuneate in shape. In the spring new growth occurs by a regeneration of the meristematic m margins of the bed in certain areas near the apex. These areas are sometimes so narrow that the new bladelets appear attached to the parent plant by a stipe-like region. The bladelets are obovate-lobed like young blades and appear thinner and lighter in color than older parts of the thallus. By these characteristics, the beginning and end of each year’s growth can easily be distinguished. Cross sections of the mature blade show that it is composed of a one- or two-layered outer cortex of small subglobose cells about 4-8 »m_ in diameter, a subcortex of one or two layers of larger subglobose cells, 12-40 pm in diameter, and a medulla consisting of interlacing stellate and filiform cells (Fig. 3). The stellate cells constitute the primary medulla; the filiform cells are derived secondarily. In the blade, the primary meristematic areas are located along the margins. The margin at the apex is more active meristematically than the lateral margins where only limited or regenerative g erowth occurs. In the meristems, growth is initiated through the division a surface cells. Each surface cell may cut off from one to three daughter cells. The first two cells are cut off anticlinally or obliquely, and the last one is cut off apically from the parent cell (Fi ig. 5). In surface view, these cells appear in groups of one to three (Fig. 4). These groups consist of the parent cell alone, the parent cell in combination with its oblique daughter cells, or the daughter cells alone. As new cells are produced at the meristem, each previous generation of cells is displaced beneath the surface, creating a series of dichotomously or trichotomously branched filaments below the meristem (Fig. 7). Cells of the upper two layers of these fila- ments are undifferentiated. In the third and fourth layers, the cells begin to form secondary pit connections with neighboring filaments (Fig. 6, 7) but retain their globose, meristematic shape. At this stage, the cells Fic. 3-10. Anatomy of the blade.—3, cross section showing medullary region containing both filiform (fil) and stellate (st) cells; scale, 50 pm. .—4, surface view near the meristematic margin; scale, 20 pm.—5, oblique and periclinal divisions of the terminal meristematic cells; scale, 20 um. .—6, peel preparation of a young cortical region showing the small cells (arrows) involved in the formation of secondary pit connections; scale, 20 ym. .—7, peridermal section through the growing tip; scale, 20 ym.—8, section showing the origin of a filiform medullary cell from the subcortex; scale, 30 pum.—9, sane preparation of subcortical cells; scale, 20 pm.—10, squash preparation of stellate cells; scale, 20 pam. CIRRULICARPUS CAROLINENSIS oF%@esese O92 eDdar®" 2c SER ae e @ 47 @ Bases alte Oe: foremene® 5,0 6 9" y é ye --3 ; ai ¢ * P . , oe 6 GAYLE I. HANSEN Fic. 11-15. Anatomy of the stipe and holdfast.—11, cross section of the stipe; scale, 100 pm.—12, longitudinal section through the stipe and holdfast; scale, 500 pm.—13, 14, longitudinal sections of holdfast showing the random orientation of the medullary filaments and the outward growth of fila- ments at the margin; scale, 30 pm.—15, surface view of the base of the holdfast showing outer cortical cells elongated into rhizoidal filaments (arrow); scale, 50 pm. formed by the meristem may develop into either cortical or medullary cells, depending on the location of their filaments in the blade margin. The cortex develops from the upper four layers of cells in filaments that are displaced from the meristematic margin. Since the number of cells in the meristem increases by a factor of up to three with each new generation, it would seem that the meristematic surface area should CIRRULICARPUS CAROLINENSIS 7 likewise increase. This does not occur. Only those cells located in the central part of the margin retain their meristematic activity. Cells that are pushed to the sides by the expanding surface cease meristematic activity. These displaced surface cells and the three layers of cells in the filaments behind them differentiate to form the cortex. Cells in the upper two layers round up slightly and form the outer cortex; they do not expand in size and do not form secondary pit connections with neighbor- ing cells, Cells in the third and fourth layers of the displaced filaments form the subcortex, slowly expanding in size and readily forming sec- ondary pit connections with neighboring cells. Subcortical cells have a subglobose to slightly stellate shape when mature (F ig. 9). The primary medulla develops from cells which are located in filaments remaining behind the meristem. Differentiation of these cells begins when they are located about five layers below the surface. Lateral tension created by the expanding meristematic surface is great enough to cause the branching filaments to become separated except where they are attached by pit connections. Initially, the young cells easily extend to form protrusions in the direction of their pit connections. Secondary pit connections continue to form between the cells by the production of elongated protuberances which extend outward across the necessary distance to connect with neighboring cells. As the expansion continues, the cells appear to be stretched tautly between their pit connections and become stellate in appearance. Young stellate cells are about 10 pm in diameter and, upon maturing, expand to 20-40 pm in size, excluding the stellate arms (Fig. 10). There are usually five or six arms per cell. Secondary growth occurs mainly in the medullary region of the blade. The space between the stretched primary medullary filaments is filled by numerous, secondarily produced, filiform medullary filaments. These, which may be multicellular, are produced by any of the cells in the subcortex or medulla (Fig. 8). They may extend transversely across the medulla from cortex to cortex or peridermally and parallel to the expanding primary medulla. The filaments frequently terminate their growth by forming secondary pit connections with other vegetative cells. Secondarily derived medullary filaments are responsible for the increase in the thickness of the thallus. They may form anywhere in the plant but are particularly abundant in the basal part of the blade and in the stipe and _ holdfast. The structure of the mature stipe is variable. Nearest the blade, the stipe consists of a highly organized cortex and medulla similar to that in the blade (Fig. 11). At the holdfast, the anatomical structure appears to be totally disorganized (Fig. 12). This is caused by the abundant production of secondarily derived medullary filaments which grow downward and intertwine (Fig. 13). At the holdfast, the secondary filaments grow out- ward around stones and other small objects, aiding attachment (Fig. 14). 8 GAYLE I. HANSEN Near the edges of the holdfast the normally globose cells of the outer cortex elongate and grow outward like rhizoidal filaments over the sub- stratum (Fig. 15). These rhizoidal cortical cells also appear to assist in anchoring the plant. Reproductive Structure. Cirrulicarpus carolinensis is dioecious. Male and female plants are found throughout the year, but young sexual fronds occur only in Jate summer and fall. In male plants, spermatangia are produced in large soral areas scat- tered over the surface of the thallus. In young bladelets, spermatangial mother cells and spermatangia begin forming in August, and spermatia are produced from August through November. After this time the sperma- tangia degenerate, although senescent mother cells remain distinguish- able on the fronds throughout the year. Spermatangia are formed from the cells of the outer cortex. Sterile outer cortical cells are usually subglobose in shape (Fig. 16, 20). On becoming fertile, they clongate slightly and cut off spermatangial mother cells obliquel ly (Fig. 17, 21), Each outer cortical cell may cleave off up to four spermatangial mother cells (Fig. 22), Each spermatangial mother cell elongates, swells at the tip, and cleaves obliquely to produce one sperma- tangium which measures 4 x 5 pm in diameter (Fig. 18). The mother cell may then elongate again and produce a second spermatangium. Spermatia are released through an apical slit directly into the water (Fig. 23), On release, spermatia are 2.0 x 2.2 »m in size, appear colorless, and are covered only by a mucilage layer (Fig. 19). After spermatium release, only the sheath of the spermatangium is evident. In the female plants young carpogonial branches develop in late September or early October. Carpogonial branch systems consist of a large supporting cell, one or two three-celled carpogonial branches, and occasionally up to two subsidiary cells (Fig. 28-30). The branch systems develop from initials which are cut off by a subcortical cell inward toward the medulla. Each initial then cuts off one or two other cells (Fig. 24), each of which may divide to form a three-celled carpogonial branch. Often before all of the cells of the carpogonial branches are formed, the cells begin to elongate and become lobed (Fig. 25, 26) and assume the shape characteristic of female reproductive cells of the genus. The initial cell becomes the dumbbell-shaped supporting cell. The first cell of each branch is elongated and lobed, and develops into either a subsidiary cell or the basal cell of a carpogonial branch, The second cell becomes the hypogenous cell of the carpogonial branch; it is usually Fic. 16-23. Spermatangial development.—16, cross section through the cortex in a sterile area of the blade.—17, cross section through an immature sorus showing the initial stages in the formation of spermatangial mother cells (spme).—18, cross section through a mature sorus showing the formation of spermatangia (sp).—19, section with a spermatium (s) resting on the thallus surface.—20, surface view of the cortex in a sterile portion of the blade.—21, 22, surface views of spermatangial mother cells formed from outer cortical cells.—23, surface of a fertile area showing the slits (sl) through which spermatia have been released; scale, all 10 mm. CIRRULICARPUS CAROLINENSIS 9 10 GAYLE I. HANSEN Fic. 24—28. Camera lucida drawings of carpogonial branch system development: sc = supporting cell, cb 1, cb 2, and cp = cells of the carpogonial branch, sub c subsidiary cells; scale, 30 ym. CIRRULICARPUS CAROLINENSIS 11 slightly smaller but just as elongated and lobed as the basal cell of the carpogonial branch. The third cell forms the carpogonium (Fig. 26-28). A trichogyne is initiated almost immediately but does not elongate until all of the cells of the carpogonial branch are mature (Fig. 26, 27). When two carpogonial branches are present, they form opposite one another on the supporting cell (Fig. 26, 30), and subsidiary cells are rare. This seems to indicate that in C irrulicarpus carolinensis subsidiary cells may correspond to the basal cell of a carpogonial branch which has stopped growth. This has been noted to occur in other members of the Kally- meniaceae by Norris (1957). Mature carpogonial branch systems are covered by papilla-like outgrowths of the outer cortex (Fig. 31). Actual fertilization or spermatization has not been observed to occur in Cirrulicarpus carolinensis, and the process of carposporophyte develop- ment appears to be quite unusual. Cells of the carpogonial branch systems do fuse (Fig. 32, 33) but then appear to degenerate. These fusion cells have never been seen producing either primary or secondary gonimoblast filaments. Instead, filaments that resemble secondary gonimoblast fila- of. “ad » I a : > @e * * »\, 8 Oe. he Py -. Sef : . 4 Fic. 29-33. Female reproductive structures.—29, monocarpogonial branch system; scale, 20 pum.— 30, dicarpogonial branch systems; scale, 20 pm.- 31, surface view of papilla covering a carpogonial branch system; scale, 20 pm.—32, 33, fusion cells formed from the cells of the carpogonial branch system; scale, 30 pm. 12 GAYLE I. HANSEN Fic. 34-38. Cell penetration.—34, peel preparation showing the spread of penetrating short-celled filaments (sf) through the subcortex.—35, infested cortical cells below a scratch on the surface of the thallus.—36, 37, single subcortical cells which have been penetrated and infested by short-celled filaments.—38, a carpogonial branch system penetrated and infested by short-celled filaments; scale, all 20 wm. ments develop from vegetative cells. These filaments, which are lightly staining and short-celled, appear to perform two functions: (1) they penetrate and infest vegetative and reproductive cells in the thallus, leading to the disintegration of these cells; or (2) they form an extensive network of cells which eventually form carposporangia. Filaments pene- trating other cells may not necessarily be able to form carposporangia, but filaments forming carposporangia can always penetrate and infest nearby cells. When the surface of the alga is scratched or in some way damaged, the cortical and subcortical cells beneath are immediately penetrated by short-celled filaments which arise from neighboring cells (Fig. 34). Frequently only the cells directly under the line of pressure are invaded (Fig. 35). At times, particular subcortical cells will be invaded by fila- ments produced by neighboring cells for no obvious reason (Fig. 36, 37). CIRRULICARPUS CAROLINENSIS 13 The same filaments also penetrate and appear to digest nutrient-rich reproductive cells (Fig. 38). Short-celled filaments that form carposporangia usually develop from cells of the subcortex. F requently they arise from subcortical cells that are located next to a carpogonial branch system and then proliferate in the area around it (Fig. 39-41). However, just as often, filaments develop from vegetative cells which are removed by numerous pit connections from the carpogonial branch systems (Fig. 42). Once initiated, the fila- ments wind their way between the vegetative cells of the cortex for a considerable distance (Fig. 43). The presence of these filaments seems to stimulate the initiation of new filaments by the surrounding vegetative cells. Eventually the density of the short-celled filaments reaches a satura- tion level in both the cortex and medulla. When this point is reached, short, almost spherical cells are produced both terminally and laterally by the filaments (Fig. 44). These cells then initiate chains of cells that rapidly increase in pigment content and in size and mature into carpo- sporangia (Fig. 45). The swelling of the carposporangia causes the cystocarp to protrude on both sides of the thallus and the carposporangial chains to appear as clusters separated by gonimoblast filaments and stretched vegetative cells (Fig. 46). Young cystocarps first appear as a gray discoloration in the thallus surface, At this stage the cortex and medulla are infested by the lightly pigmented, short-celled filaments (Fig. 47). Such regions contrast sharply with the clear cortical and medullary tissues of sterile areas of the thallus (Fig. 48). When deeply-pigmented carposporangia develop, the cysto- carp becomes deep red in color. Young cystocarps develop in the mid-to-outer portions of the fronds. They are large (up to 3 mm in diameter) and oval in shape (Fig. 49). Normally cystocarps are solitary or grouped irregularly on the fronds (Fig. 50, 51) and occasionally a ring of cystocarps is formed (Fig. 52). A central cystocarp is usually present, and one or two rings of smaller, individual or confluent cystocarps surround it. The rings appear to be initiated by some influence of the central cystocarp and are preceded in development by a gray region formed in a halo around the central cysto- carp (Fig. 50). These gray areas are identical to immature cystocarps and are caused by the outward growth of short-celled filaments from the central cystocarp. When the surrounding area is saturated with filaments, carposporangia are initiated and the area deepens in color as the ring of cystocarps forms. A second ring of cystocarps may then develop in an identical manner. Carpospores are released from mature cystocarps through cracks in the surface layers. Before release, portions of the outer cortex around the developing cystocarp degenerate, and as the carposporangia enlarge, the weakened areas in the cortex split open (Fig. 53). Each cystocarp 14 GAYLE I, HANSEN CIRRULICARPUS CAROLINENSIS 15 may release spores over a period of several weeks, after which the cysto- carps continue to grow outward and become wart-like in appearance (Fig. 54). Senescent cystocarps are often present on the lower parts of female plants long after the first period of sexual maturity. Life History. Because tetrasporophytes of Cirrulicarpus carolinensis were not found in the field, carpospores of this alga were cultured in the laboratory to determine if a tetrasporophyte phase existed in the life history. The details of these studies will be reported in a later paper, and only a brief summary is included here. Carpospores, obtained from plants of Cirrulicarpus carolinensis col- lected between October and December, were released from cystocarps in uniform round sprays (Fig. 55). The spores measured 9 x 11 pm in diameter. Germination was of the “typus filamentosus mediatus” (Inoh, 1947) and of the “persistent-spore type’ (West, 1968) and proceeded in both unipolar and bipolar fashion (Fig. 56). A heterotrichous system consisting of closely branched prostrate filaments and sparsely branched upright filaments soon developed (Fig. 57). As the germlings grew older, the prostrate system became very exten- sive. It consisted of axial filaments with closely branched lateral filaments (Fig. 58) and was similar in appearance to Hymenoclonium serpens (Crouan & Crouan) Batters. Lateral branches continued to divide until a network of very closely appressed cells was produced. The elongated shape of the axial cells contrasted with the short cells of the lateral filaments (Fig. 59). Close examination of these prostrate networks revealed that the cells of neighboring filaments frequently fuse with one another (Fig. 60). Similar fusions have been reported to occur in many members of the Corallinaceae (Rosenvinge, 1917; Cabioch, 1970, 1971), in Cruoriopsis and in Rhododermis (Rhodophysema) elegans (Rosenvinge, 1917), and in Kallymenia (Codomier, 1972, 1973). The prostrate system was initially monostromatic, but as maturation proceeded, filaments of the lower layer occasionally branched upward and formed a second layer on top of the first (Fig. 61). This process continued until a thick layered crust about four cells deep developed. These thicken- ings were usually restricted to small areas on the original monostromatic network. Occasionally cells in the upper layers of the thickenings initiated short, erect filaments which were terminated by cap-like or hair-like cells (Fig. 62). Most often these erect filaments remained as crusts, but occa- sionally, after long periods in culture, certain filaments grew upward and formed erect, multiaxial, gametophytic blades, identical to small, field- collected plants of Cirrulicarpus carolinensis (Fig. 63). Thus, the life history appears to have no tetrasporophyte generation, and germlings from carpospores may be considered protonemal in nature. Fic. 39-41. A carpogonial branch system viewed at three focal planes. A short-celled filament has originated from the neighboring subcortical cell and is actively proliferating in the vicinity of the carpogonial branch; scale, 20 pm. 16 GAYLE I. HANSEN DISCUSSION AND CONCLUSIONS Because of its vegetative structure, the number and arrangement of cells in the carpogonial branch system, and cystocarp anatomy, Cirruli- carpus carolinensis clearly be longs in the family Kallymeniaceae. The new species is placed in the genus Cirrulicarpus because it is complanate and cartilaginous and has a repeatedly branched thallus which is nar- rowly cuneate below and rounded at the apices. The plant has a fila- mentous medullary system, a carpogoni al branch system that consists of elongated or lobed cells, and cystocarps that occasion: ally form in rings. The vege tative ani itomy of C irrulic arpus carolinensis corresponds with the second of three kinds of thallus structure in the K Kallymeniaceae Fic. 42—45. Gonimoblast development.—42, the initiation of short-celled filaments from subcorti- cal cells near and far from the carpogonial branch system (epbr); scale, 20 ym.—43, a peel prepara- tion showing short-celled filaments originating near a carpogonial branch system and winding their way between the preexisting vegetative cells; scale, 30° jm. 14, short-celled filaments originating from subcortical cells and growing thickly between the vegetative cells. The small, rounded, deeply staining cells (csi) initiate carposporangial chains; scale, 20 pm. 45, short-celled filaments forming a carposporangia (cs) in chains both terminally and laterally; scale, 20 pm. CIRRULICARPUS CAROLINENSIS 17 recognized by Womersley and Norris (1971) wherein the primary medullary cells, while remaining recognizable, become separated by a “profuse and usually loose development of filaments,” and the medulla appears essentially filamentous. In C. carolinensis stellate cells form the primary medulla and filiform cells the secondary medulla. Primary growth is initiated at the surface of the margin of the flattened frond, and rows of three to four undifferentiated cells lie behind each apical cell before differentiating into stellate cells. Norris (1957, 1964) described the axial filaments as extending to the surface of the growing margin in Callophyllis, Thamnophyllis, Kallymenia, and Pugetia, seeming to imply the existence of a surface meristem. In Glaphrymenia, another member of the Kallymeniaceae, Norris (1961) described an intercalary meristem Fic. 46-48. Cystocarp structure and fertile and sterile subcortical regions.—46, a mature cystocarp with carposporangia in clusters separated by stretched vegetative cells and short-celled gonimoblast filaments; scale, 0.2 mm.—47, 48, peel preparations of the subcortex in fertile and sterile areas of the thallus; scale, 20 pm. 18 GAYLE I. HANSEN Fic. 49-54. Cystocarp morphology.—49, a female plant with young cystocarps scattered in the upper portion of the blade; scale, 3 cm.—50, a solitary young cystocarp with a gray halo (arrow ) around it; scale, 1 mm.—51, young cystocarps in a scattered arrangement; scale, 1 mm,—52, a mature cystocarp with two concentric cystocarpic rings situated around the lower side; scale, 1 mm. —53, the surface of a mature cystocarp showing the cracks through which carpospores will be released; scale, 1 mm.—54, an aging cystocarp that has already released carpospores (white area) and is becoming wart-like; scale, 1 mm. located in the terminal row of cells at the growing margin. Secondary growth in Cirrulicarpus carolinensis occurs through the production of filiform medullary filaments from subcortical and stellate cells throughout the thallus, and these filaments are responsible for its thickening. In the holdfast the secondary filaments are extremely dense and grow downward aiding it 1 attachment of the blade as was also describe d for Glaphry- menia pis das by Norris (1961). Moreover, in C. carolinensis cells of the outer cortex grow downward in a rhizoidal fashion also aiding in the attachment of the blade to the substrate. The production of sexual structures in Cirrulicarpus carolinensis occurs in the fall on the North Carolina coast. Development of both sperma- tangia and carpogonia appears to be normal. Carpogonial branch systems CIRRULICARPUS CAROLINENSIS 19 develop in a pattern similar to those described for other members of the Kallymeniaceae (Norris, 1957). Spermatangia, which have seldom been studied in this family, follow a pattern of development somewhat similar to that described for Cystoclonium purpurascens in the Gigartinales by Kylin (1923). The release of spermatia in Cirrulicarpus carolinensis occurs from August to December and brackets the time of carpogonial branch maturation in October. This timing would seem to insure fertiliza- tion, particularly since numerous individuals of both sexes are found in the field. However, no conclusive evidence of fertilization was found. Fusion cells, similar to those described by Norris (1957) as post- fertilization stages for many members of the Kallymeniaceae, are present but appear to be degenerate. Carposporophyte development appears to be initiated by vegetative cells. The production of gonimoblast filaments from vegetative cells might indicate that fertilization has taken place, but filaments are frequently initiated by cells that are quite distant from a carpogonial branch system. The only visible connection between the two is through preexisting vegetative cells and their pit connections—an unlikely path for the transfer of a diploid nucleus. In Cirrulicarpus carolinensis short-celled gonimoblast filaments may either form carposporangia or penctrate and infest vegetative or repro- ductive cells apparently to digest their contents. Since similar penetrating filaments are produced by vegetative cells in wounded areas of the thallus, one might speculate that both carposporophyte development and wounding produce a similar kind of stress on the alga. The production of gonimoblast filaments from vegetative cells has rarely been reported in the Florideophycidae and, then, only in the Cryptonemiales and Gigartinales. In Kallymenia reniformis, Hommersand and Ott (1970) found that gonimoblast filaments arose from vegetative cells that had been contacted by connecting filaments. Kraft (1973) reported similar findings in Cubiculosporium koronicarpus. In Mycodea carnosa and several species of Neurophyllis, Kraft (1974) reported the production of carposporangia from vegetative cells that (1) appeared to be in direct contact with gonimoblast filaments, (2) were “adjacent to cells contacted by gonimoblast filaments,” or (3) were “removed by a distance of many pit connections” from cells initially in contact with gonimoblast filaments. In all of the species mentioned a true auxiliary cell was also reported. Cirrulicarpus carolinensis has the distinctive characteristic of having both gonimoblast filaments borne on vegetative cells and also of having no true auxiliary cells. Mature cystocarps of Cirrulicarpus carolinensis are composed of clusters of carposporangia in chains separated by vegetative and gonimo- blast filaments very much like the cystocarps of many other members of the Kallymeniaceae. However, cystocarps of C. carolinensis may also occur in small clusters or in rings, as is characteristic of other members 20, GAYLE I. HANSEN 254) { PENG “ nat: TES Poe 1 t RS o/s SN SG ‘Ene Ny Rs Me Ke Vs E ‘ aa a > Lie th) . ~ CIRRULICARPUS CAROLINENSIS eA of this genus. Cystocarpic rings in C. carolinensis are usually formed around a central cystocarp and are produced by the outward growth of gonimoblast filaments from it. Using the terminology of Feldman (1952), the life history of Cirruli- carpus carolinensis may be considered to be digenetic, with an alternation of the gametophyte and carposporophyte generations. The absence of a tetrasporophyte generation in the life history can be explained if fertiliza- tion is not successfully achieved. The entire life history, including the development of the carposporophyte generation from vegetative cells, would then be haploid, and meiosis and tetraspore formation would be climinated. ACKNOWLEDGEMENTS This paper is based on a portion of a dissertation submitted to the faculty of the University of North Carolina in partial fulfillment of the requirements for a Ph.D. degree in botany. I wish to express my appreciation to Dr. Max Hommersand for advice during the entire course of this investigation. Dr. Elizabeth Shaw prepared the Latin diagnosis. David Montezienos and Dr. Kathryn Edwards prepared the speci- mens photographed in Figs. 8 and 62 respectively. I'am grateful to Dr. Norton Miller for critically reviewing the paper and to Dr. Donald Pfister for helpful suggestions. Preparation of the manuscript was supported by the Farlow Herbarium of Harvard University, LITERATURE CITED CABIOCH, J. 1970. Sur Pimportance des phénoménes cytologiques pour la systématique et la phylogénie des Corallinacées (Rhodophycées, Cryptonémiales ). Compt. Rend. Hebd. Séances Acad. Sci. 271D:296~—299. Sa ati . 1971. Etude sur les Corallinacées I. Charactéres généraux de la cytologie. Cahiers de Biol. Mar. 12:121-186: 2 pls. Copomiuer, L. 1972. Sur le développement des spores et sur l’origene des cellules étoilées medullaires des Kallymenia (Rhodophycées, Cryptonémiales ). Compt. Rend. Hebd. Séances Acad. Sci. 271D:369-371. a . 1973. Sur le développement des spores et la formation du thalle rampant de Kallymenia microphylla J. Ag. (Rhodophyceae, Cryptonémiales ). Giorn. Bot. Ital. 107:269-280. FELDMANN, J. 1952. Les cycles de reproduction des algues et leurs rapports avec la phylogénie. Rev. Cytol. 13:1-49., HommMteRsAND, M. H., and D. W. Orr, 1970, Development of the carposporophyte of Kallymenia reniformis (Turner) ]. Agardh. |. Phycol. 6:322-331. Hoyr, W. D. 1920. Marine algae of Beaufort, N.C., and adjacent regions. Bull. Bur. Fisheries 36:367—556; pls. 84-109. INon, S. 1947. Germination of seaweed species. Tokyo (in Japanese). 255 pp: Jouansen, D. A, 1940. Plant microtechnique. McGraw Hill, New York. 523 pp. Fic. 55-63. Growth of carpospores in culture.—55, a newly released spray of carpospores; scale, 200 um.—56, carpospores germinating in unipolar (uni) and bipolar (bi) patterns; scale, 30 pm.— 57, young germlings showing a heterotrichous pattern of development with branching prostrate filaments and unbranched upright filaments; scale, 50 um.—98, a young crust similar in morphology to Hymenoclonium serpens; scale, 50 um.—59, a maturing crustose phase with main axial filaments distinguishable by their elongated cell shape; scale, 50 um.—60, cell fusions (fu) in the crustose phase; scale, 50 um.—61, cross section through a distromatic area of the crust; scale, 20 pum.—62, cross section through a thickening with upright filaments; scale, 20 um.—63, a young erect blade; scale, 1 cm. 22 GAYLE I. HANSEN Krart, G. T. 1973. The morphology of Cubiculosporum koronicarpis gen. et sp. nov. representing a new family in the Gigartinales (Rhodophyta). Amer. J. Bot. 60:872-882. ————— . 1974. Monograph of the Mychodeaceae, Dicranemaceae and Acrotyla- ceae (Gigartinales, Rhotophyta). Ph.D. Dissertation, Univ. of Adelaide. 225 pp. Kyun, H. 1923. Studien iber die Entwicklungsgeschichte der Florideen. Kongl. Svenska Vetenskapsakad. Handl. 63: 1-139. Norris, R. E. 1957. Morphological studies on the Kallymeniaceae. Univ. Calif. Publ. Bot. 28:251-334. ——————— _ 1961. The structure and reproduction of Glaphyrymenia pustulosa. Amer. J. Bot. 48: 262-268. ——— . 1964. The morphology and taxonomy of South African Kallymeniaceae. Bot. Mar. 7:90-129. Papenruss, G. F, 1937. The structure and reproduction of Claudea multifida, Van- voorstia spectabilis, and Vanvoorstia coccinea. Symb. Bot. Upsal. 2:1-66. Pearse, A. S., AND L. G. Witxiams. 1951. The biota of the reefs of the Carolinas. J. Elisha Mitchell Sci. Soc. 67:133-161. Rosenvince, L. K. 1917. The marine algae of Denmark, Part II, Rhodophyceae Il (Cryptonemiales). D. Kgl. Danske Vidensk. Selsk. 7:155-283. West, J. A. 1968. Morphology and_ reproduction of the red alga Acrochaetium pectinatum in culture. J. Phycol. 4:58—99. Womenrs.ey, H. B. S., anp R. E. Norns. 1971. The morphology and taxonomy of Australian Kallymeniaceae (Rhodophyta). Aust. ]. Bot. Supp. Ser. 2: 1-62. A COMPARISON OF THE SPECIES OF CIRRULICARPUS (KALLYMENIACEAE, RHODOPHYTA ) GayYLE I. HANSEN ABSTRACT The three known species of Cirrulicarpus have complanate, cartilaginous, repeatedly branched thalli which reach similar maximum heights. The plants are narrowly cuneate at their holdfasts and have apices which are broadly rounded or dichotomously or irregularly lobed. All the species have filamentous medullary systems and carpogonial branch apparati consisting of elongated or lobed cells. They differ in the number of carpogonial branches and subsidiary cells located on the supporting cell, the appear- ance and arrangement of cystocarps, the presence or absence of “giant cells” in the medulla, and in minor features of blade morphology. INTRODUCTION The genus Cirrulicarpus was established by Tokida and Masaki in 1956. The name refers to the characteristic ringed arrangement of the cystocarps on the surface of the fronds in the type species. The genus now includes the following three species: C. gmelini (Grunow) Tokida & Masaki from Hokkaido and the Aleutian Islands: C. australis Womersley & Norris from Port Phillip Heads in Australia; and C. carolinensis Hansen from the coast of North Carolina. All three species of Cirrulicarpus have erect thalli that may reach 16 cm in height. The thalli are complanate, cartilaginous, and repeatedly branched in a subdichotomous manner, and they are attached to the substratum by a narrow holdfast. The blades have broadly rounded or irregularly lobed apices and margins that are entire or occasionally coarsely spinulose or erose. The genus resembles Kallymenia in having an essentially filamentous medullary system and a cortex of large inner cells grading into small outer cells. In species of Cirrulicarpus, the ‘arpogonial branch systems consist of large, elongate or lobed supporting cells bearing three-celled carpogonial branches and subsidiary cells. The number of carpogonial branches and subsidiary cells present on a sup- porting cell varies with the species. Cells of the carpogonial branch apparatus are elongated and lobed and decrease in size from the support- ing cell to the carpogonium. Cystocarps in all three species form in the upper portions of the fronds. The ring-like appearance of the cystocarps in C. gmelini is usually not evident in the other two species where cystocarps are generally solitary or in’ small irregular clusters. In section, cystocarps consist of numerous patches of carposporangia in chains separated by vegetative and gonimoblast filaments. 24 GAYLE I. HANSEN Cirrulicarpus gmelini (Grunow ) Tokida & Masaki When the genus Cirrulicarpus was first established, it included only C. gmelini. This species is based on Kallymenia gmelini, the name proposed by Grunow (1867) for a specimen collected in the Kurile Islands and deposited at the Botanisches Museum in Berlin-Dahlem. Grunow chose the specific epithet gmelini because the specimen resem- bled Gmelin’s drawings (1768, pls. 22 and 23) of Fucus palmetta. Grunow included the species in Kallymenia because it is similar to other species of this genus in vegetative and reproductive structure. J. Agardh (1899) transferred Grunow’s species to Rhodymenia without explanation. In 1915 Yendo transferred the species to Erythrophyllum because the vegetative structure of the thallus is similar to that of E. delesserioides, although thalli of K. gmelini lack the prominent midrib that occurs in E. delesserioides. Tokida and Masaki (1956) established the genus Cirrulicarpus for K. gmelini on the basis of reproductive character- istics. Cirrulicarpus differs from Erythrophyllum in having cystocarps located in rings (vs. in papillae) and intercalary tetrasporangia (vs. terminal). Norris, Tokida, and Masaki (1960) further studied the reproductive process in C. gmelini and confirmed Cirrulicarpus to be a distinct genus in the Kallymeniaceae. Many reports have established the distribution of Cirrulicarpus gmelini. Okamura (1921) reported that it was common along the coast of Hok- kaido. Nagai (1941) noted that it occurred only on the east coast of the northern and middle Kurile Islands. Setchell and Gardner (1903) found it off Agattu Island, Okamura (1933) off Atka Island, and recently Wynne (1970) off Amchitka Island, all islands in the Aleutians. Vegetative Structure. Cirrulicarpus gmelini displays a subdichotomous to polychotomous branching pattern (Fig. 1). The plant is perennial, and older specimens have a thickened basal portion which is the previous year’s growth. The older blades frequently become as narrow as 0.25 cm in width as a result of weathering and appear somewhat like an elongated portion of the stipe (Yendo, 1915). Dichotomies in older portions of the plant are usually acute and often appear torn while those in the younger portions are rounded. Apical bladelets formed during the most recent growing season are fan-shaped and irregularly dissected. The cross-sectional anatomy of Cirrulicarpus gmelini is somewhat different from the other two species (Fig. 2). The outer cortical cells, though usually subglobose in shape, are often elongated anticlinally. The cells of the subcortex are ellipsoidal in shape and elongated peri- clinally. The medullary filaments consist of cells which vary from ellip- soidal to filiform in shape. Intermixed with these filaments are very large, irregularly elongated medullary cells which extend as filaments down from the apex and then irregularly throughout the thallus. These “giant cells,” which are particularly prominent in female plants (Tokida & COMPARISON OF CIRRULICARPUS SPECIES 25 Masaki, 1956), contain a “yellow homogeneous substance which is similar to the hyaline content of the carpogones of various red algae” (Yendo, 1915). These cells stain as intense ‘ly with aniline blue as do reproductive cells (Fig. 3). The occurrence of similar ‘ ‘giant cells” has been reported to occur in Erythrophyllum delesserioides and Crosso- carpus lamuticus by Yendo. Reproductive Structure. The carpogonial branch system of Cirruli- carpus gmelini was described by Norris, Tokida, and Masaki (1960) consisting of one carpogonial branch of three cells, and two or three subsidiary cells on a lar ge supporting cell. The cells are characteristically elongated and lobed (Fig. 5). During the development of the carpogonial beiuch: all of the cells, including the carpogonium with its trichogyne, are present before clongation and lobing of any of the cells occurs (Fig. 6). Post-fertilization processes in the alga were described as nonprocarpic by Norris et al., (1960). After fertilization the carpogonium, supporting cell, and nearby subsidiary cells fuse. The large fusion cell is abundantly lobed and often appears connected to “giant cells” in the medulla (Fig. 7). According to Norris et al., fusion gets form nonseptate connecting fila- ments which grow outward through the thallus and contact auxiliary cells which then form gonimoblast filaments with carposporangia at their apices. They suggest that the cystocarpic rings in this species are produced by a ring of aude ary cells all of which are diploidized by connecting Alaments from a central, fertilized, car pogonial branch sys- tem. Young post-fertilization stages other than fusion cells were not observed by these authors. Cystocarps form in large patches on the upper portions of the fronds. Within the patches, groups of oval cystocarps, ranging from less than 0.1 to 1.0 mm in size, are arranged in elliptical and occasionally confluent rings (Fig. 4). Within cystocarpic areas, the thallus is considerably etene ‘d but the cystocarps themselves raise the surface of the frond only slightly. In section, the cystocarps appear as clusters of carpospor- angia intermixed with vegetative and gonimoblast filaments (Fig. 8). Tetrasporangial plants are similar to female plants in both habit and general morphology (Fig. 9). Tetrasporangia are abundant and scattered among the cortical cells as described by Yendo (1915). They are formed by direct conversion of cells one or two layers below the surface and thus are intercalary in origin (Tokida & Masaki, 1956). In Figs. 10 and 11 the upper or lower pit connections between tetrasporangia and nearby vegetative cells are clearly visible. The tetrasporangia are described by Tokida and Masaki as “obliquely cruciate in division.” Cirrulicarpus australis Womersley & Norris A second species of Cirrulicarpus, C. australis, was established by Womersley and Norris (1971) for specimens from Port Phillip Heads, 26 GAYLE I. HANSEN COMPARISON OF CIRRULICARPUS SPECIES 27 Australia. These algae previously had been referred to Cryptonemia inequalis, a nomen nudum of Wilson (1892), or to Kallymenia poly- coelioides |. Agardh, a different species recently redescribed by Womers- ley (1973). The authors put the species in Cirrulicarpus because the plants resembled C. gmelini in the cartilaginous and branched nature of the thallus and in the characteristic shape of the cells of the carpo- gonial branch system. Recently, Womersley (1973) has called C. australis and Meredithia nana synonyms; this is discussed on page 32. Vegetative Structure. Plants of Cirrulicarpus australis are branched in a subdichotomous or occasionally polychotomous pattern as in other members of the genus (Fig. 12). However, in the specimens I have studied, no evidence of growth from a previous year is evident. There are no thickened basal portions and the dichotomies are always rounded and untorn. Thus the plant appears to be an annual even though individual plants are comparable in size and degree of branching to the other two species. Young branches of C. ein are usually narrower than those of the other species and often are about 0.5 cm in width. Due to these narrow upper branches, C. australis is usually less foliose in appearance than the other two species, which have broad apical bladelets. The apices of C. australis are uniformly rounded or variously lobed. The cross sectional anatomy of Cirrulicarpus australis is essentially identical with that of C. carolinensis (Fig. 13). The outer cortex consists of subglobose cells which rarely appear elongated anticlinally. The sub- cortical cells are large and subglobose. The medulla consists of stellate cells that extend peridermally in a longitudinal direction in the thallus and filiform cells that extend in all directions. Reproductive Structure. The carpogonial branch system in Cirruli- carpus australis differs from the other species in having up to five carpogonial branches. Five trichogynes are evident in Fig. 15. Womersley and Norris (1971) report that two to five carpogonial branches and up to six subsidiary cells may occur. The cells of the carpogonial branch system are similar in shape to those in the other two species though they frequently appear less broadly lobed. Early post-fertilization stages have not been observed in this alga. Womersley and Norris presumed "that the female reproductive system was non-procarpic, as observed for C. gmelini, Fic. 1-11. Cirrulicarpus gmelini.—1, female plant collected by Masaki on March 27, 1945, at Shireto-Misaki, Kushiro, Hokkaido; scale, 5 cm.—2, cross section of the blade; scale, 30 pm.—3, a large, deeply staining “‘giant cell’; scale, 30 um.—4, fertile area with cystocarps grouped in rings; scale, 1 cm.—5, mature carpogonial branch system showing the elongated and lobed nature of the cells (tr = trichogyne); scale, 30 pm.—6, young carpogonial branch system in which the trichogyne has formed before the cells of the carpogonial branch have become elongated and lobed (cb 1, cb 2, cp = cells of the carpogonial branch, sc = supporting cell, sub c = subsidiary cell); scale, 30 pm.—7, fusion cell derived from cells of the carpogonial branch system with a pit connection (arrow) to a “‘giant cell’? in the medulla; scale, 30 um.—8, cross section through a mature cystocarp; scale, 30 pm.—9, tetrasporic plant collected by Masaki on November 30, 1955, at Shireto-Misaki; scale, 5 cm.—10, 11, sections showing the upper and lower pit connections (arrows) of the tetrasporangia (tsp); scale, 10 pm. 28 GAYLE I. HANSEN though they noted no fusion cells or connecting filaments. Short-celled filaments have been observed penetrating both reproductive and vege- tative cells in this species as they do in C. carolinensis. In one specimen studied, a disintegrating carpogonial branch system had outward radiat- ing filaments that appeared to be maturing into carposporangia (Fig. 16). Cystocarps of Cirrulicarpus australis are scattered in small groups on the mid to upper portions of the fronds (Fig. 14). They are hemispherical and about 1 to 2 mm in diameter. When mature, the cystocarps push up the thallus surface more on one side than on the other, and they often become wart-like in appearance. The grouping of the cystocarps is irregular, and only occasionally are rings formed. When carpospores have been released from the central portion of a large cystocarp, a white central region or hole is formed giving the illusion of a ring of several confluent cystocarps; however, these structures actually develop from a single cystocarp. The same situation occurs in C. carolinensis (Fig. 54 of the previous paper). In section, cystocarps are composed of many clusters of carposporangia (Fig. 17) as seen in the other two species. Though both male and tetrasporangial plants were reported as un- known by Womersley and Norris (1971), I found a tetrasporic specimen preserved in the Farlow Herbarium (Fig. 18). The specimen, identified as Cryptonemia inequalis J. Agardh, was collected by J. B. Wilson in 1895 from Port Phillip Heads. The plant is identical in habit and cross sectional anatomy to the female plants of Cirrulicarpus australis. Mature tetrasporangia are scattered among the cortical cells on both sides of the thallus. However, the tetrasporangia are not intercalary in origin: instead, they are formed Von surface cortical cells which, during conversion to tetrasporangia, expand and sink below the surface leaving a vacant area in the cortex above them (Fig. 19). At times the wall of the sporangium is visible on the surface above the retracted, maturing tetrasporangium (Fig. 20). Tetraspores are cleaved from the parent sporangium in an imperfectly cruciate pattern. Cirrulicarpus carolinensis Hansen This species is described in detail in the previous paper, and thus it is discussed here only for comparative purposes. Vegetative Structure. Cirrulicarpus carolinensis is widely foliose in habit, and branches in the typical subdichotomous pattern of species Fic. 12-20. Cirrulicarpus australis.—12, holotype collected by J. B. Wilson on January 15, 1895, at Port Phillip Heads; scale, 3 cm.—13, cross section of the blade; scale, 100 pm. —14, fertile area showing the grouping and wart-like nature of the cystocarps; scale, 3 cm. —15, carpogonial branch system with five trichogynes (arrows); scale, 30 um.—16, carpogonial branch systems with filaments radiating outward and forming what appear to be young carpospor: ingia (arrow); scale, 100 pm. 17, cross section through a mature cystocarp; scale, 100 ,ym.—18, tetrasporic plant collected by “a B, Wilson on January 30, 1888, at Port Phillip Heads; scale, 3 cm.—19, section showing a sunken tetrasporangium cleaved in an irregularly cruciate fashion; scale, 10 pm.—20, section with tetra- sporangium retracted below the surface leaving behind the empty wall; scale, 10 pm. COMPARISON OF CIRRULICARPUS SPECIES 30 GAYLE I. HANSEN in this genus. Because the plant is perennial like C. gmelini, the older dichotomies are acute and show evidence of tearing, while the younger ones are rounded. Older portions of the thalli become thickened and worn from weathering, though they never become as narrow as the stipe- like older blades of C. gmelini. Young apical bladelets in C. carolinensis are obovate-lobed and easily distinguish the plant from C. gmelini which has fan-shaped apical bladelets. The cross sectional anatomy of Cirrulicarpus carolinensis consists of a medulla of stellate and filiform cells and a cortex of large subglobose cells that grade into smaller nearly globose outer cells very much like those of C. australis. “Giant cells” never occur in the medulla of this species. Reproductive Structure. The carpogonial branch system of Cirruli- ». . * et ; 1 r Fw e Fic. 21-23. Meredithia nana.—21, lectotype collected by J. B. Wilson on January 27, 1883 at Port Phillip Heads; scale, 1 cm.—22, cross section of the blade; scale, 50 mm. 23, section showing gonimoblast development from a cell (arrow) which appears to be vegetative in nature; scale, 50 pm. COMPARISON OF CIRRULICARPUS SPECIES mY | carpus carolinensis consists of one or two carpogonial branches of three cells, and occasionally one or two subsidiary cells on the lobed support- ing cell. The branch system resembles that of C. gmelini which consis- tently has only one carpogonial branch. During development, the cells of the carpogonial branch apparatus clongate and lobe before the formation of the carpogonium—unlike the development of the branch apparatus in C. gmelini. Development of the carposporophyte in Cirrulicarpus carolinensis is unique in the genus. Gonimoblast filaments are produced from vegeta- tive cells and frequently can be seen penetrating and digesting both vegetative and reproductive cells before forming carposporangia. Cysto- carps occasionally form in rings. However, the rings are caused by the outward growth of secondary gonimoblast filaments from the central cystocarp and not by the outward growth of primary gonimoblast or connecting filaments to auxiliary cells as is reported to occur in C. gmelini (Norris, et al., 1960). No tetrasporophyte generation is present in Cirrulicarpus carolinensis. The alga is able to bypass this stage in its life history and forms erect gametophytic blades directly from carpospore germlings. The occurrence or lack of a tetrasporophytic generation among the species of one genus is not a serious taxonomic problem. Martin (1969) has pointed out that variations of this kind occur in the life histories of numerous red algae and may, at times, even be caused by environmental factors. DISCUSSION The genus Cirrulicarpus comprises a group of species within the Kallymeniaceae that are cartilaginous and complanate, have repeatedly branched thalli, filamentous medullary systems, and carpogonial branch systems consisting of elongated or lobed cells. Womersley and Norris (1971) considered the divided nature of the blades an important charac- teristic for distinguishing Cirrulicarpus from Kallymenia. Womersley (1973) also pointed out that the cartilaginous texture of Cirrulicarpus distinguishes it from the softer Kallymenia. Species of Cirrulicarpus differ from one another in the characters outlined in Table 1. The most im- portant characters differentiating the species appear to be the number of branches in the carpogonial branch systems, the presence or lack of “giant cells” in the medulla, and the appearance and arrangement of the cystocarps. Norris, et al., (1960) suggested that Cirrulicarpus gmelini was a primitive member of the Kallymeniaceae because it had a filamentous medullary system and because it bore carpogonial branches on its auxiliary cell sys- tem. Auxiliary cell systems have not been seen in all species of Cirruli- carpus, but all do have filamentous medullary systems. Norris (1957) also established that within the Kallymeniaceae, species with fewer carpo- 32 GAYLE I. HANSEN TABLE 1. A COMPARISON OF THE SPECIES OF CIRRULICARPUS Apical bladelets or branches Branching in mature blades Growth Medullary cells Carpogonial branch systems Cystocarp arrangement Male gametophytes Tetrasporophytes Distribution gonial branches C. gmelini C. australis C. carolinensis Fan-shaped, irregularly dissected Subdichotomous- acute Perennial Filiform to ellipsoidal cells; «e . >» giant cells” present 1 carpogonial branch, 2-3 subsidiary cells Clusters, usually in small elliptical rings Unknown Present, tetra- sporangia intercalary Hokkaido, Kurile and Aleutian Islands Narrow, dichotomously or polychotomously lobed Subdichotomous- rounded Annual Filiform and stellate cells; “giant cells” absent 2-5 carpogonial branches, 0-6 sub- sidiary cells Solitary, in clusters, rarely in rings Unknown Present, tetraspor- angia terminal Port Phillip Heads, Victoria, Australia Obovate, broadly rounded or dichotomously lobed Subdichotomous- acute Perennial Filiform and stellate cells; “giant cells” absent 1-2 carpogonial branches, 0—2 sub- sidiary cells Solitary, in clusters, rarely in rings Present Absent North Carolina on their supporting cells are more advanced. Thus C. australis with two to five carpogonial branches can be considered the most primitive, C. carolinensis with one or two carpogonial branches inter- mediate, and C. gmelini with only one branch, the most advanced. The disjunct distribution of the species of Cirrulicarpus presents a problem. It is difficult to believe that species so widely separated geo- graphically could share a common ancestor. However, as the marine algal flora of the intermediate localities is more thoroughly investigated, additional species of Cirrulicarpus may be discovered that better link the three established species. EXCLUDED SPECIES Meredithia nana |. Agardh In 1973 Womersley transferred Meredithia nana J. Agardh (1892) to Cirrulicarpus making the new combination Cirrulicarpus nana. He con- sidered the type specimen of M. nana to be a young frond of C. australis, and therefore placed C. australis in synonymy under C. nana. Earlier Womersley and Norris (1971) had expressed doubt as to whether M. nana was a member of the Kallymeniaceae due to the different nature of the carpogonial branch system and suggested that it might be a COMPARISON OF CIRRULICARPUS SPECIES 33 member of the Gigartinales. In proposing the transfer, Womersley stated that the identical vegetative structure of the plants in cross section and the similarity of their type localities were sufficient evidence of the synonymy of the two species. I have studied several specimens of Meredithia nana J. Agardh, includ- ing the lectotype (Fig. 21), which is cystocarpic. The cross sectional anatomy of the plant (Fig. 22) is similar to both Cirrulicarpus australis and C. carolinensis. Although the carpogonial branch systems in M. nana have not been observed in a young cnough state to describe precisely, they do appear to differ from those of C. australis. Post-fertilization stages have been observed, and gonimoblast filaments appear to develop from an auxiliary cell which is vegetative in nature (Fig. 23). This indicates that M. nana is most likely a member of the Gigartinales, as originally suggested by Womersley and Norris, and that it is not a member of the genus Cirrulicarpus. ACKNOWLEDGE MENTS This paper is based on a portion of a dissertation submitted to the faculty of the University of North Carolina in partial fulfillment of the requirements for a Ph.D. degree in botany. I am very grateful to Dr. Max Hommersand for invaluable sugges- tions during the entire course of this investigation. I most appreciate the very helpful comments on the manuscript by Drs. Norton Miller, Donald Pfister, and Elizabeth Shaw. Financial support for the preparation of the manuscript was provided by the Farlow Herbarium of Harvard University. Herbarium specimens of Cirrulicarpus australis, C. carolinensis, C. gmelini, and Meredithia nana were generously loaned by the following institutions and individuals: Duke University (puKE), Farlow Herbarium of Harvard University (FH), Botanical Museum at Lund (1p), National Herbarium of Victoria (MEL), University of North Carolina (Ncu), University of California at Berkeley (uc), U.S. National Museum (us), Dr. G. Kraft, University of Melbourne, Australia, Dr. T. Masaki, Hokkaido University, Japan, and Dr. C. Schneider, Trinity College, Connecticut. LITERATURE CITED Acarou, J. G, 1892. Analecta algologica. Acta Univ. Lund, 28:1-182, pls, 1-3. ——-————— . 1899, Analecta algologica. Cont. V. Acta Univ. Lund. Avd. 2, 10:1—160. GMELIN, S. G. 1768. Historia fucorum, Petropoli. 239 p., 35 pls. Grunow, A. 1867. Reise seiner majestiit Fregatte Novara um die Erde. Bot. Theil. 1. Vienna. 104 p., pls. I-11. Martin, M. J. 1969. A review of life-histories in the Nemalionales and some allied genera. Br, Phycol. J. 4:145-158. Nacat, M. 1941. Marine algae of the Kurile Islands. Il. Rhodophyceae. J. Fac. Agric. Hokkaido Univ. 46:139-319, pls. 1-3. Norris, R, E. 1957. Morphological studies on the Kallymeniaceae. Univ. Calif, Publ. Bot. 28:251-334. a , J]. Tokina, ann T. Masaki, 1960. Further studies on Cirrulicarpus gmelini (Grunow) Tokida ect Masaki. Bull. Fac. Fish. Hokkaido Univ. 11:29-36. Oxamura, K. 1921. Icones of Japanese algae. 4:85-107; pls. 171-175. ——————— - 1933. On the algae from Alaska collected by Y. Kobayashi. Records of Oceanographic Works in Japan, 5:85-97. SETCHELL, W. A., AND N. L. Garpner. 1903. Algae of northwestern America. Univ. Calif. Publ. Bot. 1:165-418, 34 GAYLE I. HANSEN Tokina, J., AND T. Masaxt. 1956. Studies on the reproductive organs of red algae. I. On Erythrophyllum gmelini (Grun.) Yendo. Bull. Fac. Fish. Hokkaido Univ. 7:63-71. Wison, J. B. 1892. Catalogue of algae collected at or near Port Phillip Heads and Western Port. Proc. R. Soc. Vict. 4:157-190. WomersLey, H. B. S. 1973. Further studies on Australian Kallymeniaceae ( Rhodo- phyta). Trans. R. Soc. S. Aust. 97:253-256. ———_——— , AND R. E. Norns. 1971. The morphology and taxonomy of Australian Kallymeniaceae (Rhodophyta). Aust. J]. Bot. Supp. Ser. 2:1-62. Wynne, M. 1970. Partial checklist of marine algae identified from 1962 to present, Amchitka Island, Alaska. (Mimeograph from Symposium on Amchitka Island Bioenvironmental Studies, Aug. 27, 1970), 2 p. Yenvo, K. 1915. Erythrophyllum gmelini (Grun.) nov, nom, Bot. Mag,, Tokyo 29 :230-237. STUDIES IN CRYPHAEACEAE III. SPHAEKROTHECIELLA FLEISCH, NEW TO THE AMERICAS! MontrE G. MANUEL? ABSTRACT The genus Sphaerotheciclla Fleisch., previously known from = eastern Asia, is reported new to the neotropics (Mexico and Venezuela). The combination S. pinnata (B.S.G.) Manuel is made. Cryphaca corrugata Card. and Pilotrichum decurrens C. Mill. are conspecific with S. pinnata. All names are lectotypified. The presence of endosporic germination and intrathecial protonemata in Sphaerotheciella is discussed. INTRODUCTION Bryophytes (and vascular plants) having a disjunct distribution be- tween eastern Asia and northern Latin America are rather well-known (Sharp, 1953 & 1972; Sharp & Iwatsuki, 1965; Wood, 1972). It is there- fore not phytogeographically surprising to find the genus Sphaerotheciella Fleisch. in the bryoflora of northern Latin America. It is known otherwise from a single species (S. sphaerocarpa (Hook.) Fleisch.) found in Nepal, Sikkim, Bhutan, and southern China (Szechwan and Yunnan ) (Noguchi, 1966 & 1971). A second species is herein added to the genus by the transfer of Cryphaea pinnata B.S.G. This taxon has been recognized since 1850. Sphaerotheciella pinnata (B.S.G.) Manuel occurs in Mexico and Venezuela and is very likely to be found in Central America (Fig. 1). MORPHOLOGICAL NOTES Sphaerotheciella Flcisch. is distinguished from the other six genera of the Cryphaeaceae in having erect gametophores, short lateral perichaetial branches, globose thecae, double peristomes, and endosporic germination associated with precocious intrathecial protonemal development. Al- though endosporic germination is known to occur in several other genera of “true mosses” e.g., Dicnemon Schwaegr., Encamptodon Mont., Mesotus Mitt. in Hook., and Cleistostoma Brid. (Ruhland, 1925), it is considered rare. However, such protonemata are rather common in Hepatophytina particularly in the orders Jungermanniales and Metzgeriales. Nehira (1966 & 1974) recognized three basic groups for 18 different sporeling patterns in the Jungermanniales. “Group III” consists of pro- tonemata that form inside enlarged exospores. From my_ preliminary observations the pattern of sporeling development in S. pinnata seems to coincide with Radula type of Group III, viz. “spore germination occurs inside the enlarged exospore and a usually unistratose, multicellular, ‘Part II of this series was published in the Bryologist 76:521—527 (1973). *Present address: Farlow Herbarium, Harvard University, 20 Divinity Avenue, Cambridge, Massa- chusetts 02138, U.S.A. 35 36 MONTE G. MANUEL disk-like protonema is formed through intercalary divisions” (Nehira, 1974, p. 157). Further studies are underway to clarify the pattern of sporeling development in Sphaerotheciella. SYSTEMATIC TREATMENT KEY TO THE SPECIES Leaf apices acuminate to somewhat abruptly contracted into wide, short, irregularly denticulate acumina; costae 0.75—-0.88 the length of leaves ............-.2.00055 je Basate uw dp be bund etna een Gee ercuitaere q*aneuse axecalers eats S. pinnata (B.S.G.) Manuel. Leaf apices acuminate to abruptly contracted into narrow acumina, acumina short to long, entire, rarely denticulate; costae 0.5-0.6(—0.7) the length of leaves ......... dpla es Pack ated Oa eg ted deere ea ene a Se ease ae S. sphaerocarpa (Hook.) Fleisch. Sphaerotheciella pinnata (B.S.G.) Manuel, comb. nov. Fics, 2-10 Basionym, Cryphaeca pinnata B.S.G., Bryol. Eur, 5:35. 1850 (fase. 44-45, mon. 5). Vide nomen, remark 1, Pilotrichum decurrens C. Mill., Syn. Muse. 2:172. 1851 (syn. nov. ).—Cryphaea decurrens (C. Miill.) Mitt., Jour. Linn. Soc., Bot. 12:414, 1869.—Cryphaea patens var. decurrens (C. Miill.) Schimp. ex Par., Index Bryol. 290. 1894.—Cryphaca patens var. decurrens (C. Miill.) Schimp. & Par. ex Par. ex Wijk et al., Index Muse. 1:517. 1959 (err. cit.). Type: Mexico. S. loc., Ehrenberg, s.n. (lectotype: BM!). Vide nomen, remark 2. Cryphaea corrugata Card., Rev. Bryol. 38:39. 1911 (syn. nov.). TYPE: Mexico. Etat de Puebla. Esperanza, Purpus 4294 (lectotype: s—pal; isotype: pc?). Vide nomen, remark 3. Gametophores green to dark green. Stoloniform shoots creeping, gen- erally inconspicuous; stems erect, up to 7 cm tall, pinnately to irregularly pinnately branched; branches 0.5-2.0 cm_ long. Flagelliform branches not observed. Stem leaves imbricate to loosely imbricate, occasionally Fic. 1. Distribution map of Sphaerotheciella pinnata. STUDIES IN CRYPHAEACEAE or Figs. 2,3; 8-lIO H—40.5mm Figs.4-6 k#—11Oum Fig. 7+——425ym Fic. 2-10. Sphaerotheciella pinnata:—2. leaves.—3a. outer perichaetial leaves.—3b. inner peri- chaetial leaves.—4. apical margin of leaf.—5. apical cells of leaf.—6. medial cells of leaf.—7. protonemata.—8. operculum.—9. calyptra.—10. theca. 38 MONTE G. MANUEL wrinkled when dry, spreading when wet, ovate-lanceolate, concave, (0.4—)0.6-0.9(-1.0) x (1.0—)1.2-1.8(-2.4) mm; apices acuminate to somewhat abruptly contracted into wide, short acumina; bases decurrent; margins revolute proximally to 0.50-0.75 the length of leaves, plane and irregularly denticulate at apices, entire below; unicostate, costae 0.75- 0.88 the length of leaves, 30.8-61.6 um wide at bases; cells thick- walled, smooth; basal marginal cells quardrate to transversely oval; inner basal cells fusiform; medial cells narrowly elliptic to widely elliptic, 4.4-11.0 (-13.0) x (5.8—)11.0-17.6 pm; apical cells narrowly elliptic to fusiform, 5.5-13.2 x 13.2-24.2(-26.4) pm. Branch leaves similar to stem leaves except smaller and less denticulate. Autoicious. Perigonal bracts ovate to ovate-lanceolate, 0.2-0.3. x 0.45-0.6(-0.8) mm; apices acuminate; generally ecostate; cells smooth. Outer perichaetial bracts ovate to lanceolate, 0.4-0.7(-1.0) x 2.0-3.0 mm; apices acuminate to abruptly acuminate into long, smooth awns; margins plane to revolute, entire; unicostate, costae percurrent to excurrent; cells smooth. Inner perichaetial bracts obovate, 0.4-0.7(-0.9) x 2.3-3.5 mm; apices abruptly contracted into long, smooth awns; shoulders sometimes revolute; margins entire throughout or occasionally slightly serrulate at shoulders; unicostate, costae excurrent into long awns. Spore germination endosporic, protonemata intrathecial, thalloid, 30-66 x 44.0-96.8 um. Thecae globose to widely obovate, (0.6—)0.7-1.0(-L.1) x 0.9-1.3(-1.5) mm; stomata not observed; exostome teeth narrowly lanceolate, smooth, proximally, papillose distally, 0.4-0.58 mm long; endostome segments rudimentary, smooth, ca 0.25 mm long ; cilia none; annuli compound; opercula conic, 0.3-0.5 x 0.4—0.5 mm. Calyptrae mitrate, smooth, 0.2—0.4 x 0.5-0.6 mm. Type. Mexico. S. loc., Liebmann, s.n. (lectotype: BM!). Habitat. Corticolous or ramicolous on trees and shrubs between 1700 and 3100 meters. Distribution. Mexico and Venezuela. Nomenclatural Remarks. 1. Only a single specimen could be found which matched the scant information contained in the protologue of Cryphaea pinnata B.S.G. This specimen, herein designated lectotype, is in the British Museum. It was formerly in the herbarium of T. Gumbel, one of the three publishing authors. 2. Carl Miiller’s types were destroyed during World War II. Thus, the name Pilotrichum decurrens needs lectotypification. From among the several specimens examined which were collected in Mexico by Ehren- berg, I have designated a specimen in the British Museum to serve as lectotype. 3. Cardot (1911) failed to mention a holotype in the protologue of Cryphaea corrugata. Accordingly I have designated the isotype at S-PA STUDIES IN CRYPHAEACEAE 39 a lectotype. Presumably an isotype also exists in Cardot’s herbarium at PC, Taxonomic Remarks. 1. Cryphaea corrugata Card., C. pinnata B.S.G,. and Pilotrichum decurrens C. Miill. are based upon differences in leaf size, shape of leaf apices, costal lengths, and the degree to which the leaves are wrinkled. After careful study, I have concluded that C. Miiller and J. Cardot described minor variations of C. pinnata, i.e., Sphaero- theciella pinnata (B.S.G.) Manuel. 2. The decision to reduce Cryphaea corrugata was based on a con- sideration of the gametophore. The lectotype lacks sporophytes and protonemata. 3. There seems to be a weak correlation between leaf areolation and the shape of the leaf apex. Leaves having acuminate apices have narrowly elliptic medial and apical cells, whereas leaves with apices somewhat abruptly contracted into wide, short acumina have elliptic to widely elliptic medial cells and narrowly elliptic to fusiform apical cells. A few exceptions to this correlation, i.e., intermediates, are known. 4, Bartram’s (1949) report of Cryphaea pinnata from Guatemala was based on misdeterminations. SPECIMENS EXAMINED. Mexico. Hidalgo: near Pellillos along Hwy 85, ca 24 mi § of Jacala, Norris 17054 (MANUEL). Near Honey Station, Pringle 10490 (rH,TENN), 15047 (FH), Above Real del Monte, 9400 ft, Sharp 519 (ny). Below Las Ventanas near Mineral el Chico, 3100 m, Sharp ct al. 1687 (stANuEL). Jacala, Sharp, 17 Dec. 1962 (TENN). Oaxaca: Llano de Las Flores, N of Oaxaca, 2000-2500 m, Iwatsuki & Sharp 5264-b (TENN). Cerro do San Felipe on Hwy 175 through Sierra Juarez, 2500 m, Sharp et al. 2358 (taNueL). Puebla: Chinantla, Liebmann 49 (BM), Liebmann 8110 (co), Liebmann s.n. (4—Br). San Luis Potosi: along Hwy 70, 23 mi E of San Luis Potosi, near k-226, 7600 ft, Magill 2582 (TENN). Alvarez, Orcutt 6523 (FH). Veracruz: El] Puerto, 7600 ft, Crum 581b (TENN). Santa Rita, Sharp et al. 3016a (TENN). Atzalan, 5700 ft, Sharp 5581 (MANUEL). Without locality. Liebmann sn. (BM). Venezuela. Merida: Mucurubé, 2600 m, Gehriger 280 (¥rH—Bart.). ACKNOWLEDGEMENTS I thank Dr. Norton G. Miller for reading the manuscript, the curators of the herbaria cited for the loan of specimens, and the Farlow Reference Library and Herbarium of Cryptogamic Botany for a post-doctoral research fellowship. LITERATURE CITED BarrraM, E. B. 1949. Mosses of Guatemala. Fieldiana, Bot. 25:1—442. Carport, J. 1911. Diagnoses préliminaires de Mousses mexicaines. Rev. Bryol. 38 :33-43. Nenira, K. 1966. Sporelings in the Jungermanniales, Jour. Sci. Hiroshima Univ., ser. B. Bot. 11: 1-49. ~- a . 1974. Phylogenetic significance of the sporeling pattern in Junger- manniales. Jour. Hattori Bot. Lab. 38:151—160. Nocucut, A. 1966. Musci. In H. Hara, “Flora of eastern Himalaya,” 1:537-591. a . 1971. Musci. In H. Hara, “Flora of eastern Himalaya,” 2:241-258, RunLanp, W. 1924. Musci, Allgemeiner Teil. In A. Engler & K. Prantl, “Die natiir- lichen Pflanzenfamilien” ed. 2, 10:1—100. 40 MONTE G. MANUEL Snanrp, A. J. 1953. Generic correlations in the flora of Mexico and eastern Asia. Jour. Tennessee Acad. Sci. 28:188. ——————— . 1972. Phytogeographical correlations between the bryophytes of eastern Asia and North America. Jour, Hattori Bot. Lab. 35:263-268. —-—— AND Z. Iwarsuxi. 1965. A preliminary statement concerning mosses common to Japan and Mexico, Ann. Missouri Bot. Gard. 52:452—456. Woop, C. E., JR. 1972. Morphology and phytogeography: the classical approach to the study of disjunction, Ann. Missouri Bot. Gard. 59: 107-124. No. 11. No. 12. Robert K. Edgar: An Annotated Bibliography of the American Microscopist and Diatomist Jacob Whitman Bailey (1811-1857) Donald H. Pfister: A Note on Phaeofabraea and its Placement in the Leotiaceae Subfamily Encoelioideae (Discomycetes) (Feb- ruary 1977). Gayle I. Hansen: Cirrulicarpus Carolinensis, A New Species in the Kallymeniaceae (Rhodophyta). A Comparison of the Species of Cirrulicarpus (Kallymeniaceae, Rhodophyta). Monte G. Manuel: Studies in Cryphaeaceae III. Sphaerotheciella Fleisch. New to the Americas (April 1977). 7 Seat ree me eee CE yaaa N ena sats ‘Li ee a 5, ia d 7 a ; A — . ate A ta. nie ves aries De) (Nee DY Hepes F aan, : Tae on ae aes