A Synopsis of the Characeae' R

BULLETIN OF THE TORREY BOTANICAL CLUB VOL. 91, No. 1, pp. 35-46 JANUARY-FEBRUARY, 1964

A synopsis of the Characeae' R. D. Wood Department of Botany, University of Rhode Island, Kingston WOOD, R. D. (Botany Dept., Univ. Rhode Island, Kingston). A synopsis of the Characeae. Bull. Torrey Bot. Club 91: 35-46. 1964.—A synopsis to the species of Characeae as delimited in the manuscript of the forthcoming monograph on Characeae of the world is presented. The bases for the author's revised classification issued in Taxon 11: 7. 1962, are revealed, and 81 species are recognized. The illegitimate name Nitella gelatinosa is replaced by Nitella tamanica, and the new subsp. gelatinifera is established. The most recent synopsis to the Characeae of the world was one by Nordstedt which appeared in "Fragmente einer Monographie der Chara- ceen " (Braun & Nordstedt, 1882: 8-25) as " Clavis synoptica characearum," and which was later translated from Latin to English and augmented with special reference to North American taxa by T. F. Allen (1888: 38-64). The present treatment is taken with but slight modification from the un- published manuscript for the "Monograph of the Characeae" which con- stitutes Part I of the "Revision of the Characeae" (Wood, R. D., and K. Imahori. m ed. Revision of the Characeae. Part I. Monograph of the Chara- ceae by R. D. Wood ; Part II. Iconograph of the Characeae by R. D. Wood and K. Imahori). The manuscript of the monograph was completed in June 1959, but publication has continued to be delayed. In order to release the more critical findings, certain parts are being reported in advance. The first, an outline of the revised classification system and nomenclature, was issued in 1962 (Wood, 1962). The present article is the second to be issued in advance of the monograph, and it presents a synopsis indicating the major readily evident features by which the writer subdivides the taxa. It is

1 Supported in part by a grant from the National Science Foundation. Received for publication October 30, 1963. 35 91 36 BULLETIN OF THE TORREY BOTANICAL CLUB [ Vol.. intended to reveal the system of classification, and is not designed as a tool for identification. Both synopses of the present type and analytical keys appear in the forthcoming "Revision." The concept of the species has plagued phycologists for years. One answer, that of continued finer subdivision into even smaller and smaller species, has been carried to extreme in various groups. It has caused exten- sive proliferation of taxa in the Characeae. Another alternative, one employed on a limited scale by both Alexander Braun (1834: 355; 1867: 907; e.g., C. foetida) and Groves and Groves (1911, e.g., C. zeylanica), is that of using broad species. Nordstedt (1882: 13) cited these as "sensu latis- sime" taxa. The present author concludes that this is the more realistic concept, and has employed it throughout his revision. Thus, there are but 81 species, rather than the earlier 314 (Wood & Imahori, 1959), included in the synopsis. More narrowly delimited taxa (species of earlier workers) are assigned to lower ranks. Those which retain some consistency over certain areas are assigned to varieties or, in occasional complex cases where several infraspecific taxa exist, to subspecies; whereas, those which are distinguishable primarily by single traits or by recombinations of char- acters are retained only as forms. The infraspecific taxa are treated in detail in the author's "Revision," but it is anticipated that the need for the formae will not support their retention in the future. The arrangement of the synopsis may be self-evident, but note that (1) Roman numerals head entries for taxa of or above the rank of genus (e.g., "IA. Tribe I, Chareae") and Arabic numbers head infrageneric taxa (e.g., "la. subgen. Chara"; or "6a. Chara globularis"). Two numbers immedi- ately precede each taxon name, the first indicating positioh within the next encompassing category, and the second (in parentheses) indicating sequen- tial position for that taxon within the entire synopsis. Distribution is sug- gested by the abbreviations in brackets in order and as follows : cosm (cosmopolitan), Eu (Europe), Af (Africa), As (Asia), Aust (Australia), Oc (Pacific Oceania), NA (North and Central America), and SA (South America). Certain popular synonyms are included in parentheses after the new names (Wood, 1962) of higher taxa. IA. Coronula of 5 cells in 1 tier ; coronula large; branchlets undivided, with 1-celled laterals (bract-cells) ...... Tribe 1(1) CHAREAE HA. Stipulodes developed (occasionally rudimentary) ; bract-cells normally 4-more at a node and including both anteriors and posteriors; branchlet segments 3-more; plant with or without cortication; bulbils unusual, 1-celled, only known in corticated species; coronular cells elongate, narrowing distally ...... Subtribe I(1) CHARINAE IIIA. With or without cortication; stipulodes in 1-2 tiers, 1-2 (-4) per branehlet (if 1, then alternate—rarely opposite) ; monoecious or dioecious; conjoined gametangia borne on antheridial stalk, thus with oogonium above antheridium (in dioecious species antheridium replaced by bractlet which subtends oogonium) ...... Genus I(1) CHARA 1964] WOOD: A SYNOPSIS OF THE CHARACEAE 37

la. Stipulodes essentially in 2 tiers, one or both of which may be reduced ...... Subgen. 1(1) CHARA (Diplostephanae) 2a. Axial cortex essentially 2-corticate ...... Sect. 1(1) CHARA (DiplOStiChae) 3a. Spine-cells solitary or rarely geminate ... Subsect. 1(1) CHARA 4a. Robust; axis 1000-2200 p. in diameter; bract-cells verticillate, inflated; antheridia 640-1425 12 in diameter; mature oospore 760-900 IA long; generally dioeeious ...... 1(1) Chara tomentosa [Eu, Af, As] 4b. More slender; axes 200-1100 p, in diameter; bract-cells generally unilateral, slender (or ovoid when small) ; antheridia to 600 (-800) p, in diameter; mature oospores 360-780 p. long ...... 2(2) Chara vulgaris [cosm] [C. pseudohydropitys [As] has naked basal branchlet cell—see 10b.] 3b. Spine-cells predominantly fasciculate, rarely solitary or geminate; monoecious ...... Subsect. 2(2) HARTMANIA ...... 1(3) Chara hispida [Eu, Af, As] [3e. Four taxa which may be difficult to place include the ecorticated form of C. socotrensis f. nuda [As], two with imperfect cortication C. vulgaris var. denudata [Eu, Af, As] and var. invert ecta [Eu, Af, As], and one 2-cortieated taxon in which the cortex resembles a 3-corticated one in appearance, C. globularis f. Capensis [Af].] 2b. Axial cortex essentially 1-corticate (occasionally second- aries partly developed) ...... Sect. 2(2) DESVAUXIA ...... 1(4) Chara eaneseens [Eu, Af, As, NA] [Taxa with imperfect cortex belonging elsewhere include C. vulgaris vars. inconnexa, denudata, and imperfeeta (4b), and C. tomentosa var. disjuneta (4a).] 2e. Axial cortex essentially 3-corticate ...... Sect. 3(3) GROVESIA ( Triplostichae) 5a. Phloeopodous (i.e., cortex of basal branchlet segment normal) ...... Subsect. 1(3) GROVESIA (Phloeopodes; Phloeobasalia) 6a. Bract-cells small to fairly large, rarely as long as branchlet segments, slender; axes to 1000 p, in diameter ...... 1(5) Chara globularis [cosra] 1 6b. Bract-cells large, c. 1 /2 times as long as branchlet segments, stout; axes c. 1500 1.1, in diameter ...... 2(6) Chara globata [As] 5b. Cortex of basal branchlet segment present, but dis- tinctively discolored or diaphanous ...... Subsect. 2(4) KuErziNotik...... 1(7) Chara setosa [Af, As] 5e. Gymnopodous (i.e., cortex absent on basal branchlet segment) ...... Sub- sect. 3(5) WILLDENOWIA (Gymnopodes; Gymnobasalia) . . . 1(8) Chara zeylanica [Af, As, Aust, 0e, NA, SA] [C. pseudohydropitys [As] has 2-corticated axis, see 10b.] lb. Stipnlodes essentially in 1 tier ...... Subgen. 2(2) CHAaorsis (TIaplostephanae) 38 BULLETIN OF THE TORREY BOTANICAL CLUB [ VOL. 91

7a. Totally ecorticate ...... Sect. 1(4) CHAROPSIS (Ecorticatae) 8a. Branchlets with terminal corona of reduced bract-cells; stipulodes and bract-cells well formed; gametangia absent from base of whirl; monoecious ...... 1(9) Chara braunii [nearly cosm] 8b. Branchlets with either a terminal mucro or corona ; stipulodes and bract-cells generally rudimentary; gametangia at either base of whorl and/or at branchlet nodes; monoecious or dioecious ...... 2(10) Chara corallina [ Af, As, Aust, Oc] Sc. Branchlets with terminal mucro, but not a corona ; stipulodes and bract-cells rudimentary ; gametangia absent from base of whorl; monoe- or dioecious ...... 3(11) Chara socotrensis [Af, As]i 7b. Axes corticated ...... Sect. 2 (5 ) AGARDHIA (Corticatae) 9a. Branchlets corticate or ecorticate, without terminal coronula of reduced bract-cells .... Subsect. 1(6) AGARDTHA 10a. Stipulodes in 1 tier; branchlets with or without cortication. h a. Stipulodes (and bract-cells) tapering; stipulodes 1-2 per branchlet, alternate. 12a. Basal branchlet segment not inflated; axes 1-3 cortieate. 13a. Gametangia restricted to branchlets ...... 1(12) Chara fibrosa [ Af, As, Aust, 0c, NA, SA] 13b. Gametangia also at base of whorl ...... 2(13) Chara leptopitys [ Aust] 12b. Basal branchlet segment of upper branchlets inflated with stored food; axes 1-corticate ...... 3(14) Chara submollusca [ Aust] 11b. Stipulodes and bract-cells large and cylindrical, blunt (i.e., allantoid) ; stipulodes 1 per branchlet, opposite (i.e., below branchlet) ...... 4(15) Chara ecklonii [Af] 10b. Stipulodes in 2 tiers; branchlets with 3-4-corticated segments ...... 15 (116) Chara pseudohydropitys [As] [Placed here despite the 2-tiered stipulodes. Entries also occur at 4b and 5c.] 91). Branchlets ecorticate, with terminal corona of reduced bract-cells; medium to small in size (to 30 cm. high) ...... Subsect. 2(7) BR k UNIA ..... 1(17 ) Chara baueri [Eu, Aust ; to be expected in As] 9c. Branchlets ecortieate, without a corona ; plants robust, often 50(-60) cm. high ...... Subsea. 3(8) WALLAIANIA 14a. Medium to large (to 60 cm. high) ; stipulodes 2 per branehlet; end segment 1-celled; bract-cells spreading ...... 1(18) Chara hornemannii [ NA, SA] 14b. Small to medium (to 15 cm.) ; stipulodes 1 per branchlet ; end segment commonly 2-celled; lower bract-cells often decumbent ...... 2(19) Chara buckellii [ NA]

1 for var. bulbilifera (in Wood, 1962: 13), see Nitellopsis belbilifera, 18b. 19641 WOOD: A SYNOPSIS OF THE CHARACEAE 39

IIIB. Without cortex; stipulodes in 1 tier (rarely irregular, sparse or absent, see 15b), 1 per branchlet, opposite (i.e., below each branchlet), strongly decumbent ; monoecious; gametangia from a single peripheral cell, and when conjoined generally with oogonium below (position varies in 15b) antheridium ...... Genus II(2) LAMPROTHAMNIUM 15a. Stipulodes well developed, uniform; oogonia below antheridia. 16a. Branehlet segments 4-5 (i.e., 3-4 nodes) ; gametangia at branchlet nodes and at base of whorl ...... 1(20) Lamp. papulosum [Eu, Af, As, Aust, Oc] 16b. Branchlet segments 3(-4) ; gametangia not at base of whorl ...... 2(21) Lamp. hansenii [Eu] 15b. Stipulodes irregular, small, some vestigial or absent; oogonium position variable, below, beside or above antheridium ...... 3(22) Lamp. succinctum [ Af, As, Aust, Oc] IIIC. With or without cortex; stipulodes in 1 tier, 2 per branchlet, spreading; monoecious; gametangia each from a separate peripheral nodal cell, generally with central oogonium flanked by 2 lateral antheridia ...... Genus III(3) LYCHNOTHAMNUS ...... 1(23) Lych. barbatus [Eu, Af, As, Aust] IIB. Stipulodes absent or rudimentary; bract-cells normally 2 at a node (occasionally 1 or 3), anterior only, similar in size and shape to branchlet segment; branchlet segments 2-5; ecortieate; dioecious; bulbils developed, 1- or more-celled; coronular cells spheroid ...... Subtribe II(2) NITELLOPSINAE ...... Genus 1(4) NITELLOPSIS 17a. Fertile branchlets with 2-3 segments (i.e., 1-2 nodes) ; branchlets without terminal mucro; bulbils multicellular (stellate) ...... 1(24) Nit. obtusa [Eu, As, NA?] 17b. Fertile branchlets with 3-5 segments; with or without terminal mucro; bulbils 1-celled (I). 18a. Sterile branehlets with 3-4 segments, without terminal mucro; 1 bract-cell at a branchlet node; bulbils (I) ...... 2(25) Nit. sarcularis [As] 18b. Sterile branchlets with 4-5 segments, with short terminal mucro; 1-2 bract-cells at a node; bulbils 1-celled ...... 3(26) Nit. bulbilifera [SA] IB. Coronula of 10 cells in 2 tiers; coronula small; branchlets divided, furcate or if monopodial then with lateral rays 2-more-celled ...... Tribe 1(2) NITELLEAE IVA. Branchlets furcate (occasionally with central ray) ; rays verticillate ; antheridia terminal on the end of branchlet rays (or at base of whorl) ; oospores laterally compressed; generally without evident lime, or if present, usually in bands or at dactyl tips. [but see Sect. Migularia, 36h, which nearly matches IVB] ...... Genus I(5) NITELLA 19a. Dactyls 1-celled (except N. Inacounii [NA], 21b; N. hookeri [ Aust, Oc], 23c) ...... Subgen. 1(3) NITELLA (Anarthrodactylae) 20a. Dactyls obtuse or acute, not acuminate; oogonial convolutions swelling at maturity ...... Sect. 1(6) NITELLA 21a. Dactyls 1-celled; branchlets 1 (-2) -furcate. 22a. Without mucus; gametangia 1-2(-3) at branchlet nodes, sessile ...... 1(27) Nitella flexilis [Eu, Af, As, Aust?, NA, SA] 22b. With mucus; gametangia 2-3 at branchlet nodes, also stipitate at base of whorl ...... 2(28) Nitella mirabilis [As] 21b. Dactyls 1 (-2-3) -celled, branchlets (1—) 2 (-3) -furcate ...... 3(29) Nitella macounii [NA] 40 BULLETIN OF THE TORREY BOTANICAL CLUB [VoL. 91

20b. Dactyls extended into prolonged point, acuminate; oogonia convolutions not swelling. 23a. Branchlets l(rarely 2)-furcate; oospore membrane not reticulate (except Y. bastinii). 24a. Typical N. acuminata ; homoeoclemous; mucus un- common ...... Sect. 2(7) RAJIA 25a. Without mucus. 26a. Branchlets 1-furcate ...... 1(30) Nitella acuminata [Af, As, 0c, NA, SA] 26b. Branchlets 1(-2-3)-furcate. 27a. Dactyla apices thick-walled at tip; branchlets 1-2-furcate ...... 2(31) Nitella heterodactyla [Af] 27b. Dactyla apices with normal wall; branchlets 1(-2-3)-fureate ...... 3(32)Nitelia polygyra [Aust] 25b. With mucus. 28a. Oospore membrane not reticulate ...... 4(33) Nitella syncarpa [Eu] 28b. Oospore membrane reticulate ...... 5(34) Nitella bastinii [NA] 24b. Wall of end cell of branchlet extremely thickened; heads tiny; dactyls tiny ...... Sect. 3(8) KNIGHTIA ...... 1(35) Nitella knightiae [Af] 24c. Cells stout; often swollen; branchlets heteroclemous, of two kinds; plant without heads or mucus ...... Sect. 4(9) BROWNIA ...... 1(36) Nitella clavata [NA, SA] 24d. Cells stout, cylindrical; homoeoclemous; plant with tiny heads with mucus ...... Sect. 5(10) RIDDELLIA ...... 1(37) Nitella cernua [NA, SA] 23b. Branchlets 1- and 2-furcate; oospore membrane reticulate ...... Sect. 6(11) PALIA 29a. Oospore membranes with 8-13 meshes across fossa; accessory branehlets 2-6; dactyls obtuse or acute ...... 1(38) Nitella stuartii [ As, Aust] 29b. Oospore membrane with 8-9 meshes across fossa; with occasional accessory branchlets at axis nodes; dactyls acuminate or acute ...... 2(39) Nitella allenii [As] 23c. Branchlets 1-2 (-3) -furcate ; dactyls 2-3 (-4)-celled; end cells allantoid to mucronate; oospore membrane ( ?) ...... Sect. 7(12) INcEaTez ...... 1(40) Nitella hookeri [Aust, Oc] 19b. Dactyls (1—)2-5-celled, of which the end cells are cylindrical, similar to other dactyl cells (generally allantoid) ...... Subgen. 2(4) HYELLA (Arthrodactylae, in part, cf. 19c; Pluricellulatae, in part). 30a. Upper whorls normal or reduced, but not modified or compacted into tight heads [but see forms of N. dualis]; with or without mucus; primary branchlet rays often enlarged. 31a. Dactyls allantoid; dense mucus; primary branchlet rays often enlarged (occasionally forms head) ...... 1(41) Nitella dualis [As, Af, Aust] [see also 35b]. 1964 WOOD: A SYNOPSIS OF THE CHARACEAE 41

31b. Dactyls irregular, allantoid to mucroniform; without mucus (1) ...... 2(42) Nitella heteroteles [ Af] (several taxa may be sought here: N. macounii [NA] —see 21b; N. hookeri [ Aust, Oc]—see 23c). 30b. Fertile whorls of modified, reduced branchlets compacted into tight heads; with mucus; primary rays normal, not 1 exceeding /2 of branchlet length. 32a. Dactyls elongate; end cell elongate, often twice as long as penultimate cell; branchlets very slender ...... 3(43) Nitella myriotricha [Aust, As] 32b. Dactyls not elongate; end cells variable but not exceeding length of penultimate cell; branchlets of normal diameter. 33a. Fertile branchlets (2–)3(-4)-furcate; central secondary branchlet rays present; monoecious ...... 4(44) Nitella capillata [NA] 33b. Fertile branchlets 1(-2)-furcate; branchlets without central secondary branchlet ray; monoecious or dioecious. 34a. Heads spicate; monoecious ...... 5(45) Nitella leptostachys [Aust, Oc] 34b. Heads variable, generally spheroid, rarely somewhat spicate; dioecious. 35a. Axial internodes shorter than branchlets; sterile branchlets 1-2 (-3 ) -furcate ; 00- spore membrane with 6-8 meshes across fossa 6(46) Nitella grovesii [AS] 35b. Axial internodes 1-2 times as long as sterile branchlets ; sterile branchlets (1–) 3 (-4)-f urcate; 2-4 meshes across fossa ...... 7(47) Nitella tasmanica2 [Aust, Oc] [See N. dualis [As, Af, Aust?], 31a, which has 6-more meshes across fossa.] 35c. Axial internodes 1-2 times as long as sterile branchlets; sterile branchlets 1-2-furcate; oospore membrane (1) ...... 8(48) Nitella polycephala [Aust] [Two pluricellulate taxa might be sought here, N. gracilis subsp. lechleri and subsp. zeyheri.] 19c. Dactyls 1-3-, 2-, 2-3-, or 2-5-celled of which the end cell is differ- ent from other cells and is not allantoid; end cell a small, tapering, pointed cell ...... Subgen. 3(5) TIEFFALLENIA (Arthrodactylae, in part, cf. 19b) 36a. Medium to large plants; some or all dactyl end cells mucronate, narrower at the base than the rounded end of the penultimate cell; heads occasionally formed, but without mucus ...... Sect. 1(13) TIEFFALLENIA (Mucronatae) [ A broad group easy to recognize but hard to define so as to clearly differentiate from Sect. Gioallenia, Sect. Tieffal- lenia includes forms resembling the European "N. niucro-

2 The taxon Nitella gelatinosa A. Br. (nom. illeg.) is here expanded to include N. tasmanica Miill. ex A. Br. Since the former name is illegitimate, the latter becomes the correct name. N. tasmanica is here construed to include subsp. tasmanica and subsp. gelatinifera R.D.W., nom. et stat. nov. (basonym: Chara gelatinosa Braun, Linnaea 17: 115. 1843; non Roth, Cat. Bot. 3: 125. 1806). 42 BULLETIN OF THE TORREY BOTANICAL CLUB [Von. 91

nata," having the tendency for plants to be broad above, mucroniform dactyls, reticulate oospore membrane, no differentiation of fertile whorls, absence of mucus, and abbreviated dactyls. Gioallenia, on the other hand, includes forms resembling the European "N. graeilis" with tendencies for plants to be narrowed above, tapered dactyls and end cells, granulate oospore membranes, for- mation of heads, presence of mucus.] [Three taxa which have mucronate dactyls, and would seem to fit here, are assigned elsewhere: N. tasmanica [Aust, Oc], N. huillensis [Af], and N. doidgeae [Af].] 37a. Branchlets uniform in a whorl; homoeoclemous. 38a. Gametangia never at base of whorl; antheridia 140-540 p, in diameter; heads rare; monoeeious (except subsp. fiagelliformis) ...... 1(49) Nitella furcata [cosm] [includes Macrodactylae and Brachydactylae]. 38b. Gametangia occasionally at base of whorl; antheridia large (to 600 1.1, in diameter) ; without heads; dioecious ...... 2(50) Nitella duthieae [Af] 38c. Gametangia never at base of whorl; antheridia 320- 500 .11 in diameter; without heads, but whorls compact, spheroid and isolated, 0.3-0.5 cm. in diameter; without mucus (4) ; dioecious ...... 3(51) Nitella plumosa [Af] 37b. Branchlets in a whorl of two kinds, heteroclemous; dioecious ...... 4(52) Nitella stephensia [Af] 36b. Medium to small; dactyl end cells not mucronate (except N. pseudollabellata var. penicillata), but base of end cell about as wide as distal end of penultimate cell, the two cell outlines tapering smoothly ...... Sect. 2(14) GIOALLENIA [A few forms have somewhat mucronate dactyls, but are smaller plants with upper whorls smaller than the lower.] 39a. Oospore membrane smooth, granulate, papillate or vermi- form, but not reticulate. 40a. Medium to large (to 60 cm.) ; some branchlets exceeding 5 cm. in length. 41a. Upper whorls lax and without mucus, or rarely somewhat spicate with mucus. 42a. Medium sized (20-30 cm.) ; with or without central secondary branchlet ray ; branchlet furcations variously 2-5-furcate; rarely dioecious ...... 1(53) Nitella pseudoflabellata [Af, As, Aust, Oc] 42b. Medium to small (5-15 cm.) ; without central secondary branchlet ray (except subsp. gracillima [As]) ; 1-3-furcate; monoecious or dioecious ...... 2(54) Nitella gracilis [cosm] [includes two pluricellulate taxa, subsp. lechleri [SA] and subsp. zeyheri [Af]]. 42e. Robust (to 60 cm.) ; axes 1000 la in diameter; dioecious. 43a. Dactyls 2-3-celled ...... 3(55) Nitella remota [Aust] 43b. Dactyls (1—)2-celled ...... 19641 WOOD: A SYNOPSIS OF THE CHARACEAE 43

...... 4(56) Nitella robertsonii [Aust] 41b. Upper whorls reduced, forming small whorls isolated along a shortened terminal (or axil- lary) axis resembling an inflorescence; without mucus ...... 5(57) Nitella rigida [As, Oct] 40b. Small, 2-5(-13) cm. high; branchlets tiny (to 0.5 cm. long) ; dense mucus; plant generally moniliform ...... 6(58) Nitella subtilissima [Aust] 39b. Oospore membrane clearly reticulate. 44a. Branchlets without central secondary ray; heads compact, spheroid, with dense mucus ...... 7(59) Niteila sphaerocephala [Af] 44b. Branchlets with a central secondary ray; without heads; no mucus ...... 8(60) Nitella tenuissima [Eu, Af, As, OA NA] 36e. Robust; similar to N. translucens (47a), but heteroclernous ...... 3(15) Sect. EARTHYA ...... 1(61) Nitella axillocarpa [Af] 36d. Medium in size; similar to N. gracilis (42b), but heteroclemous ...... 4(16) Sect. DECANDOLLEA 45a. Accessory branchlets in 2 tiers; monoecious ...... 1(62) Nitella hyalina [cosm] 45b. Accessory branchlets in 1 tier; dioecious. 46a. Accessory branchlets few, 1-6(-20) ; whorls of normal habit ...... 2(63) Nitella lhotzkyi [Aust] 46b. Accessory branchlets many, c. 40; whorls spheroid, dense, isolated along main axis, moniliform ...... 3(64) Nitella congesta [Aust, Oct] 36e. Small; similar to N. gracilis (42b), but with dactyls small and fused into 2-3-partite end cell ...... Sect. 5(17) MUELLERIA ...... 1(65) Nitella partita [Aust] 36f. Small; upper branchlets each compacted into 1-2 spheroid heads borne terminally on radiating stalklike branches ...... Sect. 6(18) VOGANIA ...... 1(66) Nitella tumida [Aust] 36g. Robust; brachydactylous; sterile branchlets often appearing to be undivided and terminated with a tiny corona of reduced dactyls; heads tiny and compact, with or without mucus ...... Sect. 7(19) PERSOONIA 47a. Dactyls 2-celled; without mucus; sterile branchlets 1-2- furcate ...... 1(67) Nitella translucens [Eu, Af, As, NA, SA] 47b. Dactyls 1-3-celled. 48a. Dactyls (2—)3-celled; axes normal; monoecious ...... 2(68) Nitella tricellularis [Aust, Oc] 48b. Dactyls 1-3-celled; axes greatly inflated (to 2500 is in diameter) ; dioecious ... 3(69) Nitella praeclara [Af] 48c. Dactyls (1—)2(-3)-celled; axes normal; heads spicate, conelike, 1-2 cm. long; dioecious ...... 4(70) Nitella conifera [Af] 48d. Dactyls 1(-2)-celled, generally reflexed; dioecious ...... 5(71) Nitella dregeana [Af] 47c. Dactyls 3-4-celled; monoecious ...... 6(72) Nitella arechavaletae [SA] 36h. Small to large; branchlets predominately monopodial, with percurrent central ray and verticillate to bilateral lateral rays at the nodes ...... Sect. 8(20) MIGULARIA 44 BULLETIN OF THE TORREY BOTANICAL CLUB [ VOL. 91

49a. Branchlets with 1-3 central axial (along central ray) nodes or furcations. 50a. Dactyls 3(-4)-celled ...... 1(73) Nitella cristata [Aust] 50b. Dactyls 1-2-celled ...... 2(74) Nitella verticillata [Aust] 49b. Branchlets with only 1 node or furcation. 51a. Sterile branehlets numerous, 5-12, undivided; dioecious ...... 3(75) Nitella struthioptila [Af] 51b. Sterile branchlets few, 5-6, generally undivided; monoecious ...... 4(76) Nitella imahorii [As] 36i. Small to large; end cells more or less mucronate; dactyls 2(-4)-celled; dioecious; with mucus ( / in N. penicillata). [Seem to have features both of sect. Tie ffailenia and sect. Gioallenia] ...... Sect. 9(21) INCERTAE 52a. Dactyls 2 (-3 ) -celled ; not clearly mucronate ; mucus I; oogonia 1 at a node; oospore membrane papillate ...... 1(77) Nitella penicillata [Aust] 52b. Dactyls 2-4-celled, clearly mueronate; with thick mucus; oogonia 1 at a node; oospore ( ...... 2(78) Nitella huillensis [Af] 52c. Dactyls 2 (-3) -celled, clearly mucronate ; with mucus ; oogonia 1-2 at a node; oospore membrane reticulate ...... 3(79) Nitella doidgeae [ Af] IVB. Branchlets monopodial, not furcately divided; rays bilateral, with laterals differentiated from ab- or adaxial rays; antheridia lateral on adaxial side of branchlet nodes, or at base of whorl; oospores terete, not compressed; uniformly lime incrusted. [similar to Nitella sect. Migularia, see 36h] ...... Genus II(6) TOLYPELLA 53a. End cells or rays allantoid and rounded at tip; coronula deciduous; oogonial convolutions swelling below coronula ...... Sect. 1(22) TOYLPELLA ( Obtusifolia; Allantoideae) ...... 1(80) Tolypella nidifica [Eu, Af, As, Aust, NA, SA] 53b. End cells of rays not allantoid; end cell small, conical, sharp pointed; coronula persistent; oogonial convolutions not swelling below coronula ...... Sect. 2(23) RommA (Acutifolia; Conoideae) ...... 1(81) Tolypella intricata [Eu, Af, As, NA, SA?]

Index Synonyms are in italics. Acutifolia 53b canescens, C. 2b acuminata, N. 26a capillata, N. 33a Agardhia, sect. of C. 7b eernua, N. 24d subsect. of C. 9a Ohara IIIA Allantoideae 53a subgen. la allenii, N. 29b sect. 2a Anarthrodactylae 19a subsect. 3a arechavaletae, N. 47e Chareae IA Art hrodactylae 19b, 19c Charinae IIA axillocarpa, N. 36c Charopsis, subgen. of C. lb barbatus, Lych. IIIC sect. of C. 7a bastinii, N. 28b clavata, N. 24c, baueri, C. 9b congesta, N. 46b Braunia, subsect. of C. 9b eonifera, N. 48c braunii, C. 8a Conoideae 53b Brownia, sect. of N. 24c corallina, C. 8b buckellii, C. 14b Corticatae 7b bulbilifera, Nit. 18b eristata, N. 50a 19641 WOOD: A SYNOPSIS OF THE CHARACEAE 45

Decandollea, sect. of N. 36d nidifiea, T. 53a denudata, var. of C. vulgaris 3c Nitella IVA Desvauxia, sect. of C. 2b subgen. 19a Diplostephanae la sect. 20a Diplostichae 2a Nitelleae, tribe of N. IB doidgeae, N. 52c Nitellopsinae, subtribe IIB dregeana, N. 48d Nitellopsis IIB dualis, N. 31a obtusa, Nit. 17a duthieae, N. 38b Obtusifolia 53a Earthya, sect. of N. 36c Palia, sect. of N. 23b ecklonii, C. lib papulosum, Lamp. 16a Ecorticatae 7a partita, N. 36e fibrosa, C. 13a penieillata, N. 52a flexilis, N. 22a Persoonia, sect. of N. 36g furcata, N. 38a Phloeobasalia 5a gelatinifera, subsp. of Phloeopodes 5a N. tasmanica 35h plumosa, N. 38c gelatinosa, N. 35b Pluricellulatae 19b Gioallenia, sect. of N. 36b polycephala, N. 35e globata, C. 6b polygyra, N. 27b globularis, C. (3c), 6a praeclara, N. 48b gracilis, N. 42b pseudoflabellata, N. 42a Grovesia, sect. of C. 2c pseudohydropitys, C. 10b subsect. of C. 5a Rajia, sect. of N. 24a grovesii, N. 35a remota, N. 43a Gymnobasalia 5c Riddellia, sect. of N. 24d Gymnopodes 5c rigida, N. 41b hansenii, Lamp. 16b robertsonii, N. 43b Haplostephanae lb Rothia, sect. of T. 53b Hartmania, subsect. of C. 3b sarcularis, Nit. 18a heterodactyla, N. 27a setosa, C . 5b heteroteles, N. 31b socotrensis, C. (3c), 8c hispida, C. 3b sphaeroeephala, N. 44a hookeri, N. 23c stephensia, N. 37b hornemannii, C. 14a struthioptila, N. 51a huillensis, N. 52b stuartii, N. 29a hyalina, N. 45a submollusea, C. 12b Hyella, subgen. of N. 19b subtilissima, N. 40b imahorii, N. 51b succinetum, Lamp. 15b imperfecta, var. of C. vulgaris 3c syncarpa, N. 28a Incertae, sect. of N. 23c tasmanica, N. 35b also sect. of N. 36i tenuissima, N. 44b intricata, T. 53b Tieffallenia, subgen. of N. 19c Knightia, sect. of N. 24b sect. of N. 36a knightiae, N. 24b Tolypella IVB Kuetzingia, subsect. of C. 5b sect. of T. 53a Lamprothamnium IIIB tomentosa, C. 4a lechleri, subsp. of N. gracilis 42b translucens, N. 47a leptopitys, C. 13b tricellularis, N. 48a leptostachys, N. 34a Triplostichae 2c lhotzkyi, N. 46a tumida, N. 36f Lyehnothamnus 1110 vertieillata, N. 50b macounii, N. 21b Vogania, sect. of N. 36f Migularia, sect. of N. 36h vulgaris, C. (3c), 4b mirabilis, N. 22b Wallmania, subsect. of C. fic Muelleria, sect. of N. 36e Willdenowia, subsect. of C. 5c Mucronatae 36a zeyheri, subsp. of N. gracilis 42b myriotricha, N. 32a zeylanica, C. 5c

46 BULLETIN OF THE TORREY BOTANICAL CLUB [VOL. 91

LITERATURE CITED Allen, T. F. 1888. The Characeae of America. Part 1. Publ. by author, New York. Braun, A. 1834. Esquisse monographique du genre Chara. Ann. Sci. Nat. II. Bot. 1: 349- 357. 1867. Die Characeen Afrika's. Monatsber. K. Akad. Wiss. Berlin (1867) : 782-800, 873-944 [reprinted and repaged 782-872]. Braun, A., and C. F. 0. NORDSTEDT. 1882. Fragmente einer Monographie der Cbaraceen. Abh. K. Akad. Wiss. Berlin 1882: 1-211, pl. 1-7. 1883. Groves, H., and J. Groves 1911. Characeae. In Ignaz Urban, Symbolae Antillanae, seu Fundamenta Florae Indiae Occidentalis 7: 30-44. Berolini. NORDSTEDT, C. F. 0. 1882. Clavis synoptica characearum. In A. Braun & C. F. 0. Nord- stedt (1882: 8-25). Wood, R. D. 1962. New combinations and taxa in the revision of Characeae. Taxon 11: 7-25 [issued 30 January]. Wood, R D., and K. IMAHORI. 1959. Geographical distribution of Characeae. Bull. Torrey Bot. Club 86(3): 172-183.