A synopsis of the Characeae'

BULLETIN OF THE TORREY BOTANICAL CLUBVOL. 91, No. 1, pp. 35-46 JANUARY-FEBRUARY, 1964A synopsis of the Characeae'R. D. WoodDepartment of Botany, University of Rhode Island, KingstonWOOD, R. D. (Botany Dept., Univ. Rhode Island, Kingston). A synopsis of theCharaceae. Bull. Torrey Bot. Club 91: 35-46. 1964.—A synopsis to the species ofCharaceae as delimited in the manuscript of the forthcoming monograph on Characeaeof the world is presented. The bases for the author's revised classification issued inTaxon 11: 7. 1962, are revealed, and 81 species are recognized. The illegitimate nameNitella gelatinosa is replaced by Nitella tamanica, and the new subsp. gelatinifera isestablished.The most recent synopsis to the Characeae of the world was one byNordstedt which appeared in "Fragmente einer Monographie der Characeen" (Braun & Nordstedt, 1882: 8-25) as " Clavis synoptica characearum,"and which was later translated from Latin to English and augmented withspecial reference to North American taxa by T. F. Allen (1888: 38-64).The present treatment is taken with but slight modification from the unpublishedmanuscript for the "Monograph of the Characeae" which constitutesPart I of the "Revision of the Characeae" (Wood, R. D., and K.Imahori. m ed. Revision of the Characeae. Part I. Monograph of the Characeaeby R. D. Wood ; Part II. Iconograph of the Characeae by R. D. Woodand K. Imahori).The manuscript of the monograph was completed in June 1959, butpublication has continued to be delayed. In order to release the morecritical findings, certain parts are being reported in advance. The first, anoutline of the revised classification system and nomenclature, was issuedin 1962 (Wood, 1962). The present article is the second to be issued inadvance of the monograph, and it presents a synopsis indicating the majorreadily evident features by which the writer subdivides the taxa. It is1Supported in part by a grant from the National Science Foundation.Received for publication October 30, 1963.35

36 BULLETIN OF THE TORREY BOTANICAL CLUB[ Vol.. 91intended to reveal the system of classification, and is not designed as a toolfor identification. Both synopses of the present type and analytical keysappear in the forthcoming "Revision."The concept of the species has plagued phycologists for years. Oneanswer, that of continued finer subdivision into even smaller and smallerspecies, has been carried to extreme in various groups. It has caused extensiveproliferation of taxa in the Characeae. Another alternative, one employedon 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 thatof using broad species. Nordstedt (1882: 13) cited these as "sensu latissime"taxa. The present author concludes that this is the more realisticconcept, and has employed it throughout his revision. Thus, there are but81 species, rather than the earlier 314 (Wood & Imahori, 1959), includedin the synopsis. More narrowly delimited taxa (species of earlier workers)are assigned to lower ranks. Those which retain some consistency over certainareas are assigned to varieties or, in occasional complex cases whereseveral infraspecific taxa exist, to subspecies; whereas, those which aredistinguishable primarily by single traits or by recombinations of charactersare retained only as forms. The infraspecific taxa are treated in detailin the author's "Revision," but it is anticipated that the need for theformae 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 immediatelyprecede each taxon name, the first indicating positioh within the nextencompassing category, and the second (in parentheses) indicating sequentialposition for that taxon within the entire synopsis. Distribution is suggestedby 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 (SouthAmerica). Certain popular synonyms are included in parentheses after thenew names (Wood, 1962) of higher taxa.IA. Coronula of 5 cells in 1 tier ; coronula large; branchlets undivided, with1-celled laterals (bract-cells) ....................................................... Tribe 1(1) CHAREAEHA. Stipulodes developed (occasionally rudimentary) ; bract-cells normally4-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; coronularcells elongate, narrowing distally ......................... Subtribe I(1) CHARINAEIIIA. With or without cortication; stipulodes in 1-2 tiers, 1-2 (-4)per branehlet (if 1, then alternate—rarely opposite) ; monoeciousor dioecious; conjoined gametangia borne on antheridialstalk, thus with oogonium above antheridium (in dioeciousspecies antheridium replaced by bractlet which subtendsoogonium) ......................................................................... Genus I(1) CHARA

1964] WOOD: A SYNOPSIS OF THE CHARACEAE 37la. Stipulodes essentially in 2 tiers, one or both of which may bereduced .................................. 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) CHARA4a. Robust; axis 1000-2200 p. in diameter; bract-cellsverticillate, inflated; antheridia 640-1425 12in diameter; mature oospore 760-900 IAlong; generally dioeeious .............................................................. 1(1) Chara tomentosa [Eu, Af, As]4b. More slender; axes 200-1100 p, in diameter;bract-cells generally unilateral, slender (orovoid when small) ; antheridia to 600 (-800)p, in diameter; mature oospores 360-780p. long ............................. 2(2) Chara vulgaris [cosm][C. pseudohydropitys [As] has naked basalbranchlet cell—see 10b.]3b. Spine-cells predominantly fasciculate, rarely solitaryor geminate; monoecious ........ Subsect. 2(2) HARTMANIA.......................................... 1(3) Chara hispida [Eu, Af, As][3e. Four taxa which may be difficult to place include theecorticated 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 inwhich the cortex resembles a 3-corticated onein appearance, C. globularis f. Capensis [Af].]2b. Axial cortex essentially 1-corticate (occasionally secondariespartly developed) ............................. Sect. 2(2) DESVAUXIA..................................... 1(4) Chara eaneseens [Eu, Af, As, NA][Taxa with imperfect cortex belonging elsewhereinclude C. vulgaris vars. inconnexa, denudata, andimperfeeta (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 segmentnormal) ...........................................................................Subsect. 1(3) GROVESIA (Phloeopodes; Phloeobasalia)6a. Bract-cells small to fairly large, rarely as long asbranchlet segments, slender; axes to 1000 p,in diameter .................... 1(5) Chara globularis [cosra]6b. Bract-cells large, c. 1 1 / 2 times as long as branchletsegments, stout; axes c. 1500 1.1, indiameter .................................. 2(6) Chara globata [As]5b. Cortex of basal branchlet segment present, but distinctivelydiscolored or diaphanous....................................................... Subsect. 2(4) KuErziNotik....................................................... 1(7) Chara setosa [Af, As]5e. Gymnopodous (i.e., cortex absent on basal branchletsegment) .............................................................. Subsect.3(5) WILLDENOWIA (Gymnopodes; Gymnobasalia). . . 1(8) Chara zeylanica [Af, As, Aust, 0e,NA, SA] [C. pseudohydropitys [As] has 2-corticatedaxis, see 10b.]lb. Stipnlodes essentially in 1 tier .................................................................................................. Subgen. 2(2) CHAaorsis (TIaplostephanae)

38 BULLETIN OF THE TORREY BOTANICAL CLUB[ VOL. 917a. Totally ecorticate ....................... Sect. 1(4) CHAROPSIS (Ecorticatae)8a. Branchlets with terminal corona of reduced bract-cells;stipulodes and bract-cells well formed; gametangiaabsent 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 atbranchlet 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 ; gametangiaabsent from base of whorl; monoeordioecious ................... 3(11) Chara socotrensis [Af, As]i7b. Axes corticated ............................ Sect. 2 (5 ) AGARDHIA (Corticatae)9a. Branchlets corticate or ecorticate, without terminalcoronula of reduced bract-cells .... Subsect. 1(6) AGARDTHA10a. Stipulodes in 1 tier; branchlets with or withoutcortication.ha. Stipulodes (and bract-cells) tapering; stipulodes1-2 per branchlet, alternate.12a. Basal branchlet segment not inflated;axes 1-3 cortieate.13a. Gametangia restricted to branchlets............................ 1(12) Charafibrosa [ Af, As, Aust, 0c, NA, SA]13b. Gametangia also at base of whorl.............. 2(13) Chara leptopitys [ Aust]12b. Basal branchlet segment of upperbranchlets inflated with storedfood; axes 1-corticate ......................................... 3(14) Chara submollusca [ Aust]11b. Stipulodes and bract-cells large and cylindrical,blunt (i.e., allantoid) ; stipulodes1 per branchlet, opposite (i.e.,below branchlet) ....... 4(15) Chara ecklonii [Af]10b. Stipulodes in 2 tiers; branchlets with 3-4-corticatedsegments ..................................................................................... 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 reducedbract-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) WALLAIANIA14a. Medium to large (to 60 cm. high) ; stipulodes2 per branehlet; end segment 1-celled;bract-cells spreading ................................................................................. 1(18) Chara hornemannii [NA, SA]14b. Small to medium (to 15 cm.) ; stipulodes 1 perbranchlet ; 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 39IIIB. Without cortex; stipulodes in 1 tier (rarely irregular, sparse orabsent, see 15b), 1 per branchlet, opposite (i.e., below eachbranchlet), strongly decumbent ; monoecious; gametangia froma single peripheral cell, and when conjoined generally withoogonium below (position varies in 15b) antheridium ..................................................................................... Genus II(2) LAMPROTHAMNIUM15a. Stipulodes well developed, uniform; oogonia below antheridia.16a. Branehlet segments 4-5 (i.e., 3-4 nodes) ; gametangia atbranchlet nodes and at base of whorl .............................................. 1(20) Lamp. papulosum [Eu, Af, As, Aust, Oc]16b. Branchlet segments 3(-4) ; gametangia not at base ofwhorl ................................................... 2(21) Lamp. hansenii [Eu]15b. Stipulodes irregular, small, some vestigial or absent; oogoniumposition 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 separateperipheral nodal cell, generally with central oogonium flankedby 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 tobranchlet segment; branchlet segments 2-5; ecortieate; dioecious;bulbils developed, 1- or more-celled; coronular cells spheroid................................................................................. Subtribe II(2) NITELLOPSINAE.............................................................................................. Genus 1(4) NITELLOPSIS17a. Fertile branchlets with 2-3 segments (i.e., 1-2 nodes) ; branchletswithout terminal mucro; bulbils multicellular (stellate).................................................................... 1(24) Nit. obtusa [Eu, As, NA?]17b. Fertile branchlets with 3-5 segments; with or without terminalmucro; 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 terminalmucro; 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 orif monopodial then with lateral rays 2-more-celled ............ Tribe 1(2) NITELLEAEIVA. Branchlets furcate (occasionally with central ray) ; rays verticillate ;antheridia terminal on the end of branchlet rays (or at base ofwhorl) ; oospores laterally compressed; generally without evidentlime, or if present, usually in bands or at dactyl tips. [but see Sect.Migularia, 36h, which nearly matches IVB] ................ Genus I(5) NITELLA19a. 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 convolutionsswelling at maturity ............................................... Sect. 1(6) NITELLA21a. Dactyls 1-celled; branchlets 1 (-2) -furcate.22a. Without mucus; gametangia 1-2(-3) at branchletnodes, 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. 9120b. Dactyls extended into prolonged point, acuminate; oogoniaconvolutions not swelling.23a. Branchlets l(rarely 2)-furcate; oospore membrane notreticulate (except Y. bastinii).24a. Typical N. acuminata ; homoeoclemous; mucus uncommon............................................................ Sect. 2(7) RAJIA25a. 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 withtiny 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) PALIA29a. 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 axisnodes; dactyls acuminate or acute ................................................................................ 2(39) Nitella allenii [As]23c. Branchlets 1-2 (-3) -furcate ; dactyls 2-3 (-4)-celled; endcells 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 compactedinto tight heads [but see forms of N. dualis];with or without mucus; primary branchlet rays oftenenlarged.31a. Dactyls allantoid; dense mucus; primary branchlet raysoften enlarged (occasionally forms head) .......................................................... 1(41) Nitella dualis [As, Af, Aust][see also 35b].

1964 WOOD: A SYNOPSIS OF THE CHARACEAE 4131b. Dactyls irregular, allantoid to mucroniform; withoutmucus (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 intotight heads; with mucus; primary rays normal, notexceeding 1 / 2 of branchlet length.32a. Dactyls elongate; end cell elongate, often twice as long aspenultimate cell; branchlets very slender .......................................................... 3(43) Nitella myriotricha [Aust, As]32b. Dactyls not elongate; end cells variable but not exceedinglength of penultimate cell; branchlets of normaldiameter.33a. Fertile branchlets (2–)3(-4)-furcate; central secondarybranchlet rays present; monoecious ............................................................... 4(44) Nitella capillata [NA]33b. Fertile branchlets 1(-2)-furcate; branchlets withoutcentral secondary branchlet ray; monoecious ordioecious.34a. Heads spicate; monoecious ............................................................... 5(45) Nitella leptostachys [Aust, Oc]34b. Heads variable, generally spheroid, rarely somewhatspicate; dioecious.35a. Axial internodes shorter than branchlets;sterile branchlets 1-2 (-3 ) -furcate ; 00-spore membrane with 6-8 meshes acrossfossa 6(46) Nitella grovesii [AS]35b. Axial internodes 1-2 times as long as sterilebranchlets ; 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 sterilebranchlets; sterile branchlets 1-2-furcate;oospore membrane (1) ......................................... 8(48) Nitella polycephala [Aust][Two pluricellulate taxa might besought here, N. gracilis subsp. lechleriand subsp. zeyheri.]19c. Dactyls 1-3-, 2-, 2-3-, or 2-5-celled of which the end cell is differentfrom 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 thepenultimate cell; heads occasionally formed, but withoutmucus ............................. Sect. 1(13) TIEFFALLENIA (Mucronatae)[ A broad group easy to recognize but hard to define so asto clearly differentiate from Sect. Gioallenia, Sect. Tieffalleniaincludes forms resembling the European "N. niucro-2The 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 thecorrect name. N. tasmanica is here construed to include subsp. tasmanica and subsp.gelatinifera R.D.W., nom. et stat. nov. (basonym: Chara gelatinosa Braun, Linnaea17: 115. 1843; non Roth, Cat. Bot. 3: 125. 1806).

42 BULLETIN OF THE TORREY BOTANICAL CLUB [Von. 91nata," having the tendency for plants to be broad above,mucroniform dactyls, reticulate oospore membrane, nodifferentiation of fertile whorls, absence of mucus, andabbreviated dactyls. Gioallenia, on the other hand, includesforms resembling the European "N. graeilis" withtendencies for plants to be narrowed above, tapereddactyls and end cells, granulate oospore membranes, formationof heads, presence of mucus.] [Three taxa whichhave mucronate dactyls, and would seem to fit here, areassigned 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; antheridia140-540 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; antheridialarge (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 whorlscompact, spheroid and isolated, 0.3-0.5 cm. indiameter; 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 cellabout as wide as distal end of penultimate cell, the twocell outlines tapering smoothly ............................................................................................................................ Sect. 2(14) GIOALLENIA[A few forms have somewhat mucronate dactyls, but aresmaller plants with upper whorls smaller than the lower.]39a. Oospore membrane smooth, granulate, papillate or vermiform,but not reticulate.40a. Medium to large (to 60 cm.) ; some branchlets exceeding5 cm. in length.41a. Upper whorls lax and without mucus, or rarelysomewhat spicate with mucus.42a. Medium sized (20-30 cm.) ; with or withoutcentral 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.) ; withoutcentral secondary branchlet ray (exceptsubsp. 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 isolatedalong a shortened terminal (or axillary)axis resembling an inflorescence; withoutmucus ................ 5(57) Nitella rigida [As, Oct]40b. Small, 2-5(-13) cm. high; branchlets tiny (to 0.5cm. 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; withoutheads; 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. DECANDOLLEA45a. 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 ofnormal 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 andfused into 2-3-partite end cell ..................... Sect. 5(17) MUELLERIA................................................................... 1(65) Nitella partita [Aust]36f. Small; upper branchlets each compacted into 1-2 spheroidheads borne terminally on radiating stalklike branches................................................................................... Sect. 6(18) VOGANIA................................................................... 1(66) Nitella tumida [Aust]36g. Robust; brachydactylous; sterile branchlets often appearingto be undivided and terminated with a tiny corona of reduceddactyls; heads tiny and compact, with or withoutmucus .................................................................. Sect. 7(19) PERSOONIA47a. 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 isin diameter) ; dioecious ... 3(69) Nitella praeclara [Af]48c. Dactyls (1—)2(-3)-celled; axes normal; headsspicate, 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, withpercurrent central ray and verticillate to bilateral lateralrays at the nodes .......................................... Sect. 8(20) MIGULARIA

44 BULLETIN OF THE TORREY BOTANICAL CLUB [ VOL. 9149a. 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; dactyls2(-4)-celled; dioecious; with mucus ( / in N. penicillata).[Seem to have features both of sect. Tie ffailenia and sect.Gioallenia] ........................................................... Sect. 9(21) INCERTAE52a. 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, withlaterals differentiated from ab- or adaxial rays; antheridia lateralon adaxial side of branchlet nodes, or at base of whorl; oosporesterete, not compressed; uniformly lime incrusted.[similar to Nitella sect. Migularia, see 36h] ........ Genus II(6) TOLYPELLA53a. 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, sharppointed; coronula persistent; oogonial convolutions not swellingbelow coronula ........ Sect. 2(23) RommA (Acutifolia; Conoideae)...................................... 1(81) Tolypella intricata [Eu, Af, As, NA, SA?]IndexSynonyms are in italics.Acutifolia 53b canescens, C. 2bacuminata, N. 26a capillata, N. 33aAgardhia, sect. of C. 7b eernua, N. 24dsubsect. of C. 9a Ohara IIIAAllantoideae 53a subgen. laallenii, N. 29b sect. 2aAnarthrodactylae 19a subsect. 3aarechavaletae, N. 47e Chareae IAArt hrodactylae 19b, 19c Charinae IIAaxillocarpa, N. 36c Charopsis, subgen. of C. lbbarbatus, Lych. IIIC sect. of C. 7abastinii, N. 28b clavata, N. 24c,baueri, C. 9b congesta, N. 46bBraunia, subsect. of C. 9b eonifera, N. 48cbraunii, C. 8a Conoideae 53bBrownia, sect. of N. 24c corallina, C. 8bbuckellii, C. 14b Corticatae 7bbulbilifera, Nit. 18b eristata, N. 50a

19641 WOOD: A SYNOPSIS OF THE CHARACEAE 45Decandollea, sect. of N.denudata, var. of C. vulgarisDesvauxia, sect. of C.DiplostephanaeDiplostichaedoidgeae, N.dregeana, N.dualis, N.duthieae, N.Earthya, sect. of N.ecklonii, C.Ecorticataefibrosa, C.flexilis, N.furcata, N.gelatinifera, subsp. ofN. tasmanicagelatinosa, N.Gioallenia, sect. of N.globata, C.globularis, C.gracilis, N.Grovesia, sect. of C.subsect. of C.grovesii, N.GymnobasaliaGymnopodeshansenii, Lamp.HaplostephanaeHartmania, subsect. of C.heterodactyla, N.heteroteles, N.hispida, C.hookeri, N.hornemannii, C.huillensis, N.hyalina, N.Hyella, subgen. of N.imahorii, N.imperfecta, var. of C. vulgarisIncertae, sect. of N.also sect. of N.intricata, T.Knightia, sect. of N.knightiae, N.Kuetzingia, subsect. of C.Lamprothamniumlechleri, subsp. of N. gracilisleptopitys, C.leptostachys, N.lhotzkyi, N.Lyehnothamnusmacounii, N.Migularia, sect. of N.mirabilis, N.Muelleria, sect. of N.Mucronataemyriotricha, N.36d nidifiea, T.3c Nitella2bsubgen.lasect.2a Nitelleae, tribe of N.52c Nitellopsinae, subtribe48d Nitellopsis31a obtusa, Nit.38b Obtusifolia36c Palia, sect. of N.lib papulosum, Lamp.7a partita, N.13a penieillata, N.22a Persoonia, sect. of N.38a PhloeobasaliaPhloeopodes35h plumosa, N.35b Pluricellulatae36b polycephala, N.6b polygyra, N.(3c), 6a praeclara, N.42b pseudoflabellata, N.2c pseudohydropitys, C.5a Rajia, sect. of N.35a remota, N.5c Riddellia, sect. of N.5c rigida, N.16b robertsonii, N.lb Rothia, sect. of T.3b sarcularis, Nit.27a setosa, C .31b socotrensis, C.3b sphaeroeephala, N.23c stephensia, N.14a struthioptila, N.52b stuartii, N.45a submollusea, C.19b subtilissima, N.51b succinetum, Lamp.3c syncarpa, N.23c tasmanica, N.36i tenuissima, N.53b24bTieffallenia, subgen. of N.sect. of N.24b Tolypella5b sect. of T.IIIB tomentosa, C.42b translucens, N.13b tricellularis, N.34a Triplostichae46a tumida, N.1110 vertieillata, N.21b Vogania, sect. of N.36h vulgaris, C.22b Wallmania, subsect. of C.36e Willdenowia, subsect. of C.36a zeyheri, subsp. of N. gracilis32a zeylanica, C.53aIVA19a20aIBIIBIIB17a53a23b16a36e52a36g5a5a38c19b35e27b48b42a10b24a43a24d41b43b53b18a5b(3c), 8c44a37b51a29a12b40b15b28a35b44b19c36aIVB53a4a47a48a2c36f50b36f(3c), 4bfic5c42b5c

46 BULLETIN OF THE TORREY BOTANICAL CLUB [VOL. 91LITERATURE CITEDAllen, 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, seuFundamenta Florae Indiae Occidentalis 7: 30-44. Berolini.NORDSTEDT, C. F. 0. 1882. Clavis synoptica characearum. In A. Braun & C. F. 0. Nordstedt(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. TorreyBot. Club 86(3): 172-183.