The Taxonomy of Polysiphonia in Hawaii!
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The Taxonomy of Polysiphonia in Hawaii! ERNANI G. MENEZ2 ABSTRACT: An investigation of Polysiphonia collections from Oahu , Hawaii, Molokai, and Maui of the Hawaiian Islands has revealed the presence of seven species: Polysiphonia mollis Hook. & Harv., Polysiphonia pulvinata ( Roth) J. Ag., Polysiphonia subtilissima Mont., Polysiphonia ferulacea Suhr., Polysiphonia yona kuniensis Segi, Polysiphonia flabellttlata Harv., and Polysiph onia rhizoidea sp. nov. These seven species of Polysiphonia were recognized primarily by their morpho- . logical features. Some characteristics of Polysiphonia which have not been previously used by monographers but which appear to be important criteria for delimiting specific entities are discussed. One of these is the presence of more than one secondary pit connection between adjacent peri central cells, a condition present in P. rhizoidea and P. yonakuniensis but not in the other 'species mentioned above. The other is the presence of multicellular rhizoids, a condition which was observed only in P. rhizoidea. Previously, authors have accepted the rhizoids of Polysipho nia as being unicellular. ALTHOUGH there are a number of published evaluate the validity of these taxa and to de articles on the marine algae of Hawaii, none of termine if additional species of Polysiphonia are them deals specifically or intensively with the represented in Hawaii. The species included in taxonomy of Polysiphonia. Up to the present, this study were recognized primarily by morph o only six species of Polysiph onia have been re logical features. corded in the literature as occurring in Hawaii: Polysiphonia aquamara Abbott, Polyiiphonia ACKNOWLEDGMENTS calothrix Harv., Polysiphonia ferulacea Suhr., The author is indebted to Dr. Maxwell S. Polysiphonia mollis Hook. & Harv., Polysiphonia polyphysa Kuerz., and Polysiphonia tongatensis Dory, of the University of Hawaii, Dr. George Hollenberg, of the University of Redlands, Cali Harv. fornia, and Dr. Toshio Segi, of the Prefectural With respect to the published records of these six species, reference may be made to the works University of Mie, Japan, for their invaluable of Chamberlain (1860), Falkenberg (1901), comments; to Dr. Benjamin Stone, of Guam College, for preparing the Latin description of Tilden (1901), Lemmermann (1905 ), Mac Polysiphonia rhizoidea sp, nov.; to Mr. E. H. Caughey (1918 ), Yendo (1918), Weber van Bosse (1923), Neal (1930), and Abbott (1947). Bryan, Jr., and Miss M. Neal for permission to examine the exsicatti materials of Polysiph onia The objectives of the present study were to in the Bernice P. Bishop Museum; to Drs. 1 Part of a thesis submitted to the Graduate School . Charles Lamoureux and Sidney Townsley, of the of the University of Hawaii in partial fulfillm ent of University of Hawaii, for their critical reading the requirements for the degr ee of Master of Science in of this paper; and to the Botany Department of Botany, March, 196 2. Manuscript received March 8, 1963. the University of Hawaii, for providing field 2 Biology and Botany Department , Uni versity of and laboratory facilities and supplies for this Brit ish Columbia, Vancouver 8, Canada. research. 207 208 PACIFIC SCIENCE, Vol. XVIII, April 1964 MATERIALS AND METHODS THE GENUS Polysiphonia The collections of Polysiphonia in Dr. M. S. Th e genus Polysiphonia, established by Dory's herbarium (DUH) ; Polysiphonia col Greville in 1824, belongs to the family Rho lections deposited at the Bernice P. Bishop domelaceae in the Rhodophyta. According to Museum (BISH) by 1. Abbott, E. Bailey, J. Falkenberg (190 1), Polysipho nia should include Rock, D. Rogers, and J. Tilden; and my own all those radial1y symmetrical members of the collections around Oahu (M) were the materials . Rhodomelaceae in which ( 1) at least the ulti used in this study. mate branches are evidently polysiphonous, ( 2) most of the branches arise exogenously by a A. Collection of the Biological Materials more or less diagonal division of subapical cells before these have cut off pericentr al cells, (3) As much area as possible was covered at all branches are essentially similar and inde every station. The materials collected were first terminate, and (4 ) only one tetrasporangium is placed in plastic bags and later transferr ed to borne normally in each segment. wide-mouthed bottles filled with 10 per cent formalin. IMPORTANT TAXONOMIC FEA TURES B. Investigation of the Biological Materials in the Laboratory , A . T richoblasts Little difficulty was met in the preparation Rosenvinge ( 1903) referred to the structure of the materials for the detailed studies. There that other investigators have called "leaves" as were three types of preserved materials: old "rrichoblasrs." Commonly, one trichoblast occurs mounted dry specimens, materi als preserved in on each segment. Hollenberg ( 1942) com formalin for several years, and fresh collections mented on the rrichoblasrs as being usually ar in 10 per cent formalin. ranged spirally in a counter-clockwise direction A drop of water was added to each of the old toward the tip of the branch, looking at the branch from the apex. In the present paper this mounted specimens to make them soft enough would be referred to as a right-hand spiral. to avoid damaging any structures when prepar Hollenberg also stated that the divergence of ing samples for microscopic study. It was usually the trichoblasts in the spiral is relatively con necessary to boil such dry specimens for a few stant and usually bears some relation to the minutes to restore the normal cell shape. On number of pericentral cells. Th e trich oblasts the other hand, the old specimens preserved in may besimple or forked, consisting of one to formalin did not need to be boiled but had to several segments of cells. Th ey are usually pres be stained to distinguish the pericenrral cells, ent at or near the apices. Generally, they are using 1 per cent aniline blue or 1 per cent deciduous, leaving conspicuous -scar cells after safranin O. The newly collected materials were they fall off. soften ed in sea water-formalin solution for 24 48 hours before examination. B. Branches A dozen or more slides of each collection In the Rhodomelaceae, Falkenberg ( 1901) were prepared. A few thallus fragments were distinguished two types of branches: (l ) de taken from each collection and placed on one terminate branches, which do not ordinarily or more slides with a drop of 10 per cent give rise to further branches, and ( 2) inde glycerine solution. termin ate branches, those of potentially un For counting the number of pericentral cells limited growth. The branches are eith er of the materials were eith er crushed on the slide endogenous or exogenous origin . Branches aris or cross-sections were cut. Important charac ing after the peri central cells have been formed teristics of the different species were illustrated are designated as "endogenous," whereas those with a camera lucida. Measurements were made formed before the formation of the pericentral with an ocular micrometer. cells are referred to as "exogenous." The branches Polysiphonia in Hawaii-MENEZ 209 of all the Polysiphonia species described in this mal end or middle of the pericentral cells. They paper are exogenous. are commonly unicellular, but sometimes multi Hollenberg (1942) described two types of cellular. Each mayor may not be cut off by a exogenous branches : (1) normally exogenous cross-wall at its proximal end from the support branches, arising directly from the branch pri ing pericentral cell. The tip of the rhizoid may mordia, and (2) cicatrigenous branches , devel be simple, digitate, or discoid. oping from the scar cells after the trichoblast has fallen off. F. Pericentral Cells and Cortical Cells In the tetrasiphonous species of Polysiphonia C. Antheridia (Segi, 1951), the number of pericentral cells The antherid ia usually occur as primary is usually constant throughout the entire thallus, branches of the trichoblasts or may occur on the while in species with more than four siphons trichoblasts, covering the lower part of the latter. the number of peri central cells may vary in the At the apex of each antheridium a sterile tip of different parts of the thallus. one or more cells may be found . Each antherid Cortical cells may be observed in some species. ium is usually supported at the base by a two They are usually formed near the base of the celled stalk. thallus and arise from the pericentral cells. In these corticate species the rhizoids are produced Cystocarps D. from the cortical cells. All the Polysiphonia Polysiphonia cystocarps are very variable in species described in this paper are uncorticated. shape. They are usually urceolate, ovate, or oblong and seem to be little used for systematic OBSERVATIONS AND RESULTS or taxonomic purposes. Polysiphonia ferulacea Suhringar in J. Agardh, E. Rhizoids Spec. Alg. II:980, 1863 The rhizoids occur at the base of the thallus. Figure 1, A-E While they may also be found on the axis above the thallus base they are never found near the TYPE: From the Atlantic coast of Mexico, thallus apex. The rhizoids may occur singly or North America , and presumed to be in the several may be produced from a segment of the Agardhian herbarium, University of Lund. thallus. They arise as outgrowths from the proxi- Thalli up to 5 em in height, usually shorter. KEY TO THE HAWAIIAN SPECIES OF Polysiphonia 1. Pericentral cells 4 .__ _ __ __ __ __ _ __ _ 2 1. Pericentral cells more than 4 .______ __ ____________ __ ______ __ ___ ___ ____ ____ __ __ ___ __ ___ __ __ __ __ __ ____ ___ __ 5 2. Thalli chiefly erect; segments from as long as broad to 4 times as long as broad 3 2. Thalli chiefly prostrate; segments half as long as broad , Polysiphonia ferulacea 3. Rhizoids cut off by a cross-wall from the proximal ends of the supporting peri central cells .