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    Taxonomy and Ecological Distribution of Chaetophoraceae

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    64 visualizações33 páginas

    Taxonomy and Ecological Distribution of Chaetophoraceae

    Enviado por

    Geraldo Ramos
    Taxonomy and Ecological Distribution of Chaetophoraceae

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    © All Rights Reserved
    Baixe no formato PDF ou leia online no Scribd
    Sinalizar o conteúdo como inadequado
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    Teaarseaisoe [AS] Saat Ano Taxonomy and ecological distribution of Chaetophoraceae (Chaetophorales, Chlorophyta) in lotic ecosystems from Sao Paulo State, southeastern Brazil By Ciro Cesar Zanini BRaNco UNICENTROICEDETEG, Depto, Biologia, Guarapuava, PR, Brazil ‘Optanbo Neccut JONtoR and Luts Henrique Zantst Branco LUNESP/BILCE, Depto. Zoologia e Botanica, Sio José do Rio Preto, SP, Brazil With 35 figures and 3 tables in the text Abstract: 172 stream segments were Sampled forthe presence of Chactophoraceae repre- ‘hhuatincs among macroalgal communities from May to October in 1992-1993 and 1996— 1907 in five natural repions (parts of biomes or selected geological areas) and one dis- turbed area of Sio Paulo State, soueaster Brazil (19°45 -21°25'S, 49°45'-S1°30°W), “This study is based on the examination of 97 simples, from which ten species of Choeto~ phoraveae were identified: Chactophora attenuata, C. elegans, C. psiformis, Draparnal- tia mutabils, Siigeoclonium amoenum, S. farctum, S. fascicular, 8. helveticum, Stu ‘iricum and § subsecundum. Chactophora attenuata and S. subsecundum were reported for the fist time for Brazil For each species the following information was provided: de- ‘Cription, diagnostic fetuteilustration, ist of populations examined, known distribution tn Brasil and associated macroalgal species. Speties of Chaetophore and Stigeoctonium ‘Were evaluated according to @ new set of taxonomic eriteria combining characters from prostate and erect systems. The numberof species found in streams from So Paulo State ‘Nas relatively high when compared to other world regions; however, species abundance (be cover) showed lower values. No correlations between number and abundance of the family and environmental variables were found but the following combination of environ mcnal parametcts was the most closely related to the occurrence of Chaetophoraceae: fisher temperatures and current velocity intermediate specific conductance, oxygen sat- lation and turbidity and pH circumnedtra, All species reported for Sao Paulo State can bbe considered as having wide geographical distribution, occurring in tropical and temper- Ie regions of both hemispheres. members of Chaetophoraceae were found in all biomes! ‘gions, The most widespread species was C. elegans, occurring in four ofthe six regions [tcatigated: on the ot hand, 30% of the total spocies were recorded ina single biome fezion. A reappraisal of specific identification in Chaetophoraceae is clearly needed. ‘Ghoul include larger number of populations and habitats, and should embrace classical, ‘morphological characters, as well 8s ultrastructural and molecular data (0342-1123 02/0184.085 5825 © 2002 Schweizer sche Velagshucbhndhag.D-70176 Stata “Agog Stuties 106 Arch, Hyranol.SupL 144 “4 C.C.Z, Branco eta Key words: Chaeiophora, Chactophoraccare, Draparnaldia distribution, macroa Stigeoclonium, steam, w@xonom’. Introduction ‘The classification of the green algae included under Chaetophoraceae is a major source of discussion with divergent opinions and litle agreement about taxo- nomic concepts (JOHN 1984, Sar 1986) and, thus, of doubtful validity. How- ever, that author comments that genera and species delimitation are closely associated with vegetative instead of reproductive characteristics, because the re= productive process is not very specialized and exhibits a narrow variation among different taxa. These considerations are readily evident when the taxonomy of some of the main genera of the family is investigated ‘The taxonomy of Chaerophora Scikavk is traditionally based on character istics ofthe gelatinous thallus, including properties of erect filaments and mui laginous matrix, . g. ype and orientation of branching, morphometric attributes, of vegetative cells, hair presence or absence and texture and shape of mucilage (Hazen 1902, Prescorr 1962, Privrz 1964, Starwancn 1972, Sansa. 1986), However, these taxonomic characteristics are recognized as variable under env ronmental changes which cause high levels of plasticity and polymorphism, The genus Draparnaldia Bory also has a very complicated and confused tax- ‘onomy, with sharply divergent opinions concerning the characteristics to be used to separate different species. Prescort (1962), Pristz (1964), Starwaxcu (1972) aand Saxwia (1986) adopted characteristics related with fascicles for species de- limitations, such as shape and width of lateral branches, presence or absence of ‘rachis, cell dimensions and type of chloroplast. On the other hand, many studies have indicated a broad variation of these attributes duc to environmental condi tions (Forrst 1956, 1957, Sarma 1964, Jouvstove 1978, Loxuiorst 1984, van Bren & Srwons 1988), Van Been & Sivions (1988) studied the growth and mor- Phology of D. muabils in culture and concluded thatthe boundaries among dif ferent species are frequently indistinct and only four or five species should be recognized. ‘The genus Stigeoclonium Korzisc is certainly the most intensively investi- gated of the Chaetophoraceae. The occurrence of variation in the relative devel. pment ofthe heterotrichous thallus in the different species mae its taxonomy very difficult and controversial, involving a heated discussion of the eritera tha: should be adopted for species determination (Is. am 1963, Cox & Bo.o 1966, Sanais 1986, Franckt & Sinons 1984, Stioss et al. 1986). Thus, two divergem, {taxonomic approaches emerged, one essentially based on characteristics ofthe erect system and the other relied on the prostrate system. Is. aut (1963), in his rex vision of Stigeoclonium, chose the erect portion of thallus a the most mmpor tax. ‘nomic character. Several authors have followed IsLaM's position (Haze 1902, Heexivo 1914, Coutins 1928, Prvtz 1964, Sranwanct 1972, Sansa 1986). Dif. ‘Chactophoraceae in lotic ecosystems from So Paulo State, Brasil 45 ferently, Cox & Botn (1966) proposed a new organization in the classification system of these algae, mostly based on morphological and metric characteristics Of the prostate portion of the plant. Therefore, it is evident that the dominant tax- ‘onomic schemes for Stigeoclonium represent extreme views with one part ofthe hheterotrichous thallus prevailing over the other, ‘Members of Chaetophoraceae are commonly reported in lotic ecosystems ‘around the world, specially among macroalgal communities, particularly species of the genera Chaetophora, Draparnaldia and Stigeoclonium (Houwies & Wut TON 1981, JonANssow 1982, SHEATH et al. 1986, 1988, 1989, Bioas & Price 1987, Enrwiste 1989, Siteari & Cove 1992, Branco & Neccit 1996a). However, there has been no recent taxonomic account of the Chactophoraceae which attempts t0 incorporate characteristics of both erect and prostrate systems on the basis of a large set of field populations. Most taxonomic information concerning Brazilian species is found in old general floras (Dickie 1881, Montvs 1892, Eowatt 1896, Scinapce 1901, Borat 1918, 1925, Kueteexorer 1955, Prescorr 1957). Some papers have been published more recently (Is.aw 1963, Dias 1989, 1997, Fas. ccescust 1992, Fraycescuint & Court, 1992). ‘Considering that the Chaetophoraceae has a wide distribution in macroalgal ‘communities and the lack of more specific taxonomic treatments about this green algal group in streams, the present study was carried out to identify and describe genera and infra-generic taxa occurting in lotic habitats of Sao Paulo State, southeastem Brazil. In addition, we analyzed the populational variability for ‘morphological characters of taxonomic importance. Simultaneously, species o¢- currence was investigated in accordance with stream environmental data in order {o evaluate ecological tendencies and geographic distribution patterns for lotic Chactophoraceae. This study is part of broader project dealing with macroalgal ‘communities in lotic ecosystems of the region. Material and methods The occurrence of macroscopic populations of Chaetophoraceae was investigat- din five natural regions (parts of biomes or geological areas) and one disturbed area of S20 Paulo State, southeastern Brazil, 19°45°-21°25'S, 49°45" -S1°30'W (Fig. 1). The total area of the State is 247,898 km2, The sampling program was conducted to include the main biomes (ALoxso 1977) and geological regions rep- resented in Sdo Paulo State: tropical rainforest (TF), subtropical rainforest (SF), Atlantic rainforest (AF), cerrado (CE, Brazilian savanna), hard water regions (HW, draining calcareous rocks) and northwest region (NW, a highly disturbed region with a mixture of cerrado and tropical rainforest, which was sampled for comparison). Emphasis was given to the best preserved areas representing each biome (according to Kronka et al. 1993). Two areas draining caleareous rocks were sampled (Fig. |, E-F), according to the map elaborated by the Technolog- ical Research Institute (IPT), Sao Paulo State. We have previously published 46 .Z, Branco etal Fig. 1. Distribution of natural vegetation cover in Sio Paulo State showing the sampling ‘areas with specification of biomes (adapted from Kxoska etal. 1993). A ~ northwest re- gion (NW); B-C — Tropical rainforest (TF); D ~ Cerrado (CE); E-F ~ Han water regions (HW), G-lT~ Adlantic rainforest (AF); I~ Subtropical rainforest (SF). Biomes represent ‘the dominant vegetation type in each area, parts ofthis general approach for northwest region (Neccut etal. 1995, 1997) and castem Atlantic rainforest (Branco & Necexi 1996a, 19966). Here we summa- Fize the entie study, including the data ofthese previous investigations 172 stream segments were sampled from late fll through early spring (May to October) in 1992-1993 and 1996-1997, typically the period with the highest diversity and abu dance of macroalgae in the region (Neca & Pascoatoto 1993) Sites were visted once and were 10-40 km apart Each sampling site consisted of 10-m reaches in streams of vat: ible sizes (Ist to Sth order), which were thoroughly examined for the presence of Chae ‘ophoraceae representatives among the macroalgal communities. In addition, abundance ‘of each species was estimated in terms of percentage cover. Algal samples were preserved immediately after collection in 4 96 buffered formal dehyde oniaxsson 1982). A representative set of algal samples was incorporated in the Herbarium SJRP (Hounciex & Hovwckex 1993), Microscopical examination, measure ‘ments and photomicrographs were made, respectively, whith a BIS OLYMPUS micto- Scope, a micrometer eyepiece and a PM-10 AD photomicrographic system coupled to the ‘microscope. 20 measurements were randomly taken for momphometric structures consid {ered to be of taxonomic importance in previous studies. This number was et ‘the equation (Sournwooo 1978): n= (s/x)2, where s~ standard deviation x~ mean, E= predetermined standard-eror (here 0.05). For each species the following information is Provided! a) deseripion; b) diagnostic features; illustrations: d) ist of populations ex: amined ¢) known distribution in Brazil (in States); e) associated macroaleal species (For cach algal division), In addition, remarks, inluding taxonomic characters and difficulties in identification, are presented. Values for morphometric characters appear in parentheses in descriptions only if they represent exceptions, i. ., do not fit within the typical 95 ‘confidence interval or te typical values along the frequency distribution curve ‘Chaetophoraceae in lotic ecosystems from Sao Paulo State, Brasil 47 Four type specimens were analyzed, as follows (herbaria acronyms according to Howwcnts etal. 1990): 1) Stigeoctonium amoenum Korzixc (1845; 198). L $1292 (holotype). 2) Stigeoctonium lubricum (Dicwyn) Kerase (1845: 198). Herd. Kerzne, L 51283 (holotype). 3) Stigeoclonium tenue (Acaxpi) Kovzin« (1843: 253). C (lectotype) 4) Stigeoctonium subsecundum (Korat) Koraise (1843: 253) Herb. Kovzine, L 51290 (lectotype here designated), ‘The following stream variables were measured: temperature, specific conduct- ance, turbidity, current velocity, pH and oxygen saturation, as previously de- seribed (Necow et al. 1995, Branco & Neccn 1996b). In addition, nutrients (total nitrogen, nitrate, ammonium, total phosphorus, calcium and potassium) ‘were also analyzed. They were measured with Orion ion selective electrodes and «Horiba F-24 ion meter, except total N and P, which were analyzed according to the technique of VaLpereaMa (1981). Current velocity was measured at a depth ofS em in three equidistant points within the stream reach with a 2030R General Oceanies mechanical flowmeter, In addition, substratum type (accord- ing to particle size classes given by Gobo et al. 1992) and shading, on the basis ‘ofa four-class scale (open, partly shaded, shaded and heavily shaded, modified from DiNicota et al, 1992), were also annotated treatment Key to species of Chaetophoraceae in lotic ecosystems of So Paulo State. 4a Branched filaments immersed within a firm gelatinous thallus .....2 (Chaetophora) 2a Basal system composed of prostrate filament; erect filaments fasciculated at the some mit elegans 2b Basal system without prostate filaments; erect filaments not fasciculated at the coms ‘mit 3 32 Basal system composed of palmetoid ceils; erect filaments sparsely branched, stent. ated atthe summit C.aiteruata 3b Basal system composed of thizoids;crect filaments densely branched, not attenuated atthe summit C pisiformis 1b Branched filaments not immersed ina frm gelatinous thallus 4 ‘4a Main axis of erect filaments composed of cells strongly differentiated fom eels of iat. eral branches Draparnalia mutabilis ‘4b Main axis of erect filaments composed of ces similar ia shape to cells of lateral branches. 5 (Siigeacionium) Sa Basal system composed exclusively ofrhizoids 6 {6a Main axis of erect system differentiated in two kinds of celis, short and long ‘Svamoenum { Main axis of erect system composed of celis similar in sie and shape 7 7a Lateral branches fascicuated atthe summit, S fasciculare To Lateral branches not fasciculated at the summit S helvericum 48 C.C.Z Branco etal Si Basal sytem composed of prostrate filaments or 3 psewloparenchymatous disks ‘8 Basal system composed of pseudoparenchymatousdise-likestrcture; erect system lit- tle developed S farctum ‘8b Basal system composed of prostrate filaments; erect system well developed 9 9a Mam axis feet te diferentated in tw Kids of ells, sort and long S lubricum ‘9p Main axis oF erect system composed of cells similar in size and shape 'S.subsecundum Chactophora attenuata Wxres (Figs 2-4) Mem. Torey Bot Club, 11 (2): 1902 Plants with a firm, variously shaped, light green gelatinous matrix 0,2-1.0 cm in diameter; erect filaments di oF trichotomously branched; branches loose and spreading, not fasciculated atthe suramit; terminal branches with strongly atten~ uated apices, often setiferous, 12-19 (-25) per main axis, basalapical diameter 2.0-2.6; cells of main axis cylindrical, 3.0-67.5 ym long, 5.0-8.0 um diam Jength diameter 5.0-7.0; chloroplast consisting of« median band in cells of main axis and laminate in cells of peripheral regions of the thallus; basal system com posed of « mass of palmelloid cells, producing erect filaments and thizoids, thi zoids also developing from the branched filaments. Diagnostic features: Plants with basal system formed by a mass of palmelloid cells producing rhizoids; erect filaments with apices strongly attenu- ated (basal/apial diameter 2.0-2.6) Distribution in Brazil: First record of the species. Associated species: Chlorophyta: Chaetophora elegans (Rov) C. A. ACARD, Mic rospora stagnorum (Korz0sG) Lasceaint, Nitella subgiomerata 8. Beaty, Spirgyra Spp.; Cyanophyta: Phormidium rerzii Gowoxt ; Rhodophyta: Batrachospermum mac rosporum Moxtacst, “Chantransia” stage. Populations examined: TF — Ocaugu, |.Skm from town, Corego Taruma, 22°25'19'S, 49°55°10'W; Teodoro Sampaio, $0 Sebastido Farm, 20 km from town, be ‘sveen Teodoro Sampaio and Planalto, Ribeirde Culaba, 22°23 08°S, $2°15°28W, 14.vi, 19: NW ~ Guapiagu, Corrego do Ipé, 20°46'45°S, 49°55'10-W, 21.1997. Remarks: In protologue of C. atenuata, Haze (1902) pointed out the features ‘of muscilaginous matrix and the branching pattern of erect filaments and com- pared this species with C elegans (Rom) C.A. Acton and C. pisiformis (Rom) C.A. Acaaoii, He ascribed the presence of thinner and attenuated filaments as an essential charaeteristic to distinguish C- attenuaa from the remaining species in this genus, as well as also the abundant occurrence of rhizoids arising ftom a ‘mass of palmelloid cells. The material collected in Sdo Paulo State fully agrees with the original circumscription. The only incongruence isthe presence of fila- ‘ments with large diameter observed inthis study. However, C.atfemuata was the species with smallest diameter values (X= 7.0 am; C. elegans X = 7.3 ym and C pisiformis x = 7.4 ur) occurring in Sio Paulo State. Saka (1986) proposed a new variety, C. attenuata var. claytoni, based on differences of eel size and shape in erect system, However, these characteristics were not observed in the populations studied. ‘Chaetophoraceae in lotic ecosystems from Sio Paulo State, Brasil 49 ee ZAI 19 Figs 2-10. Morphological features of Chaetophora species. Figs 2-4: Chaetophora attenuata. 2 ~ Basal system showing thizoids and palmelioid cell (arrow), 3 — Not fasciculate branching rect filaments. 4~ Attenuated apices of ene fila Figs 5-7: Chaetophora elegans. 8 Basal system showing prostate filament (arrow).6 Prostrate filament (arrow) with rhizoids and erect branches. 7 Apical region sowing a fascicle Figs 8-10: Chactophora pisiformis. 8~ Basal system showing hizoids (arrow) 9 Plant showing thizoids (arrow) and erect branches. 10 ~ Not fasciculate branching erect fil {Scale bars: $0 um 0 Figs 3-10; 10 um to Fig. 2] 50 C.C.Z, Branco etal Chaetophora elegans (Rovi) CA. AGaxoH (Figs 5-7) Dispositio Algarum Sueciae 4:42, 1812 Plants with a sof, variously shaped, ight green gelatinous matrix 0.5-2.0 em in diameter; erect filaments di- or trichotomously branched; branches loose and spreading, fasciculated near to periphery; terminal branches with rounded or pointed apices, rarely ending ina multicellular hair, 9-20 per main axis; cells of ‘main axis cylindrical, rarely inflated, 42.-95.0 um long, 5.0-10.0 um diam, lengti/diameter 5.7-8.2 (-11.9); chloroplast consisting of « median band in cells ‘of the main axis and laminate in cells of peripheral regions of the thallus; basal system composed of prostrate branched filaments with cylindrical cells, produc- ing erect filaments and rhizoids Diagnostic features: Plants with well-developed basal system, formed by pros- trate branching filaments, with thizoids; erect filaments strongly developed with lax branches and fasciculated at summit. Distribution in Brazil: Gotés: Prrscorr (1957); Rio de Janeiro: Diss (1997); Sio Paulo: Branco & Neccnt (1996, 19966, 1997), Neccat & Mottita (1995), Neceat & Pascoaloto (1993), Necem etal. (1991, 1992, 1998, 1995, 1997). Associated ‘species: Chlorophyta: Chaetophora atenuauta Hazen, Chara mar- ana Ws.ustan, Microspora floccosa (Vavcuen) THURET, Mlcrospora stagnorum (KOT? 's0) Lacennmst, Ntella furcata (Roxsuncn ex BRU7ELIS) C.A. ACARD emend RD. ‘Woo subsp. mucronata (A. Brain) Woon, Oedogonium spp, Spirogyra spp, Stigeve “onium helveticum Viscier, 5. amoenum Krzivs, Chrysophyta: Vaucheria spp ; Cyano phyta: Calosrir fisca Bows et FLaniav, Oscllatoria princeps Vavcuee ex Gown? Phormidium aerugineo-caeruleum (Gosioxt) ANacwostiDs et KOMAREK, P. resi Gowosr, P: willl (Gaxonte) AnacxosTIDNs et KoMAnEK, Rhodophyta: Batrachospermun ‘ambiguaam Mow act, B. delicatulum (Sku) Neccwi et ENTwistt, B. macrosporum Mon. ‘TaNE, “Chantransi’ stage, Compsopogon coeruleus (Bats ex C. Avast) Mont aon Populations examined: TF ~ Ocaugu, 1-5 km from town, Comego Tarum: 22°25'19°S, 49°55'10°W, I7-vi, 1997; AF - Redengio da Serra, road, SP-I21, 6 kn fr town, 23°15°00°S, 45°20°00'W, 18.vii.1993; HW ~ Apiai, road SP-245, 16 km frr Apiai,24°23'50°S, 48°52°35'W 08.x. 1996; Ribeirio Branco, road SP-249, 35 km frr Apiai, 24°17°33°S, 48°49°06 W, 07.x.1996; NW ~ Cardoso, near to the town, Corre ‘20 Tomazinho, 17.v.1993;,Potirendaba, road to Nova Alianga, Ribeirio do Bot. 21°01 00"S, 49°27°00-W, ix.1992; v.1993, Riolindia, road 322, I km from town, 49°55'00'S, 20°04'00°W, 17.v.1993; Sio José do Rio Preto, Cérrego da. Lagoa, 20°49 06°, 49°20'00°W, 16 sii. 1995; O1.ix. 1995; 18.1995. Remarks: This species was considered by Haven (1902) and Sansa (1986) as extemely variable. The populations collected in lotic systems of Sdo Paulo State were highly polymorphic. The gelatinous matrix, for instance, generally ‘exhibited a soft consistency, but, in many populations plants with marked differ= «ences in relation to ths typical condition were found, thus specimens with firm ‘or much softer gelatinous matrix were encountered. Similar variations were observed as to the shape of the gelatinous matrix: the globulous shape can be considered typical of C. elegans, but tuberculous and confluent (amorphous) gelatinous thalli were found in some plants. Saxsta (1986) described this species as having slender apical branches or ending in a multicellular hair. In Sio Paulo State, plants with slender apical branches where of limited occurrence Chactophoraceae in lotic ecosystems from So Paulo State, Brasil $1 (20 % of all populations) and those with multicellular hair were rarely observed (© 10%). The present results give conflicting morphometric data for C. elegans when. ‘compared with those of Paescorr (1962) and Sanwa (1986). Cell length variation in the main axis was considerably lower in those works (15.0-30.0 um and 13.0~ 31.0 um, respectively). However, cell diameter variation was very consistent in the three studies (7.0-12.0 um and 8.0-9.0 jm, respectively). Considering that C. elegans showed high morphological plasticity (Hazen 1902, Sara 1986, Dias 1997), and that cell length is a widely variable morphological character, one ‘can admit that these differences consist of an expression of the polymorphism of this alga, Hazen (1902) and Sazaa (1986) reported morphometric data more closely related with those reported for our populations, which reinforces the con- sideration above. ‘The recognized polymorphism of C: elegans indicates that it is, probably, a complex of species without clearly delimited circumscription. In this study, the presence of prostrate filaments in basal system was considered as an important taxonomic criterion and a differential and consistent feature of this species. Chaetophora pisiformis (Rov) C.A. AGaRDI (Figs 8-10) Dispositio Algarum Sueciae 4:43, 1812 Plants with a sof, light green gelatinous matrix, 0.3-4.0 em in diameter: erect fi aments di- or trichotomously branched; branches lax and densely, not fasciculat- ‘ed near to periphery; terminal branches with rounded apices, rarely ending in a ‘multicellular hair, 8-15 per main axis; cells of main axis cylindrical, rarely in- flated, 27.5-72.0 um long, 5.0-12.5 ym diam., length/diameter 5.4-9.2: chloro- plast a median band in cells of main axis and laminate in cells of peripheral regions of the thallus; basal system composed exclusively of rhizoids raising from initial cells of erect filaments, occasionally from cell of median region of filaments. Diagnostic features: Plants with basal system formed exclusively by thizoids raising from initial cells of erect filaments or occasionally from cells of ‘median region; erect filaments system strongly developed, characterized by pres- ‘ence of densely branched filaments not fasciculated in regions near to periphery. Distribution in Brazil: Golds: Prescort (1957), Associated species: Chlorophyta: Chaetophora elegans (Rom) CA. ACARD, Mougeotia spp., Nitella furcata (Roxmurcni ex Bruits) CA, Acakon emend RD. ‘Woon subsp. mucronata (A. Braun) Wooo var. mucronata Woon et IMaiont, Oedoge- hum spp.. Spirogyra spp.. Zygnema spp.; Chrysophyta: Vancheria spp; Cyanophyta: C= lindrospermum cf minutissimum Couns, "Geitlerinema splendidum (GoMOSt) Axacxostips, Hapalosiphon fhacuosus Boe, Microcoleus subtorulasus Gosost, Oscil- latoria princeps Vavcute ex Gonos, Rivularia ef. beccariana Boaset et Faint; Rhodophyta: Barrachospermum delicatulum (Skus) Necew et Extwis., “Chantransia™ stage, Compsopogon leptocladus MONTAG Populations examined: AF ~ Eldorado Paulista, road to Sete Barras, 2 km from town, 24°33/00"S, 48°05 00" W, 09.ix. 1993; CE ~ Espirito Santo do Turvo, 1 km from road SP-225, Cérego Rangel, 24°42°14°S, 49°25'42 W, 14.ix.1997; Lengois Paulista, 52 C.C.Z, Branco et al. Rio turvnho, road SP-261, 22°44°42'S, 49°06'5S"W. 164.1997; NW — Amério de Campos, 3km from town, Ribeiro das Aguas Paradas, 20°18'00S, 49°46 00-W. 181-1983: Cedral near to the road to Poirendaka, Rio Preto, 20°85'00"S, 49°19°00°W, 4x. 1992: xii.1992: 1.1993: i. 1998, Guzolindia, oad SP-S9S, Corrego das Lagos, 21°52'00'S, S0°400'W, 15.v.1998; Sao José do Rio Preto, Cortego Talhadino, 20°43 00'S, 49°1500°W, 28.4. 997; 12.v,1997: 28.1997, 12.1. 1997, 28.9197, 8.i.199%; 23.1997; 1245.1997, 278.1997, 155.1993, Vila Alves, 3m frm town, Cérrego Rods, 20°11°00°S, 49°88 00-W, 189.1993. Remarks: Hazex (1902) commented that C. pisiformis occasionally shows some similarities with C. elegans, but, as a rule, these two species can be easily distinguished, The same author pointed out that C. pisiformis included plants with firm and more resistant gelatinous matrix and thinner filaments, more densely aggregated and less numerous inthe apices than C. elegans. These char- acteristics were observed by us indistinetly in populations of both species inves- tigated. Therefore, they are not appropriate to identify these species. The relative similarity between erect filaments of C. pisiformis and C. elegans and the general tendency to identify those Chaetophoraceae with a gelatinous thallus as a mem- ber of the later (Dixs 1997), imposed the need of application or use of basal sys- tem characteristics to separate these species. Thus, C. elegans has a basal system ‘mainly composed of prostrate filaments, whereas, C. pisiformis has a basal sys- tem formed exclusively of rhizoids, Draparnatdia mutabitis (orn) Bory (Figs 11-13) Ann, Mus. National Hist, Natu. 12: 405, 1808, Thallus consisting of branched filaments, involved by soft, almost fluid muci- lage; main axis distinet with inflated cylindrical cells, (42.0-) $0.0-152.0 um long, 44.0-110.0 um diam., length’/diameter 0.9-1.5; lateral fascicle nearly al- ways without distinct rachis, ovate branching, di or trichotomic, alternate or ‘opposite; cells of lateral branches cylindrical, 26.0-S0.0 um long, 8.0-14.0 um diam, length/diameter 2.6-4.6; chloroplast fimbriated, typically forming a ‘median band in main axis cells and laminated in lateral fascicle cells, covering ‘most of the cell wall; terminal cells of lateral fascicles acuminate or rounded. ‘multicellular hairs absent; basal system formed by profuse mass of branched rhizoids, Diagnostic features: Plants with basal system formed exclusively of hiz~ ‘ids raising from inital cells of erect filaments or occasionally from cells of me- dian region; erect filaments system strongly developed, characterized by the presence of densely branched filaments not fasciculated in regions near to pe- riphery. Distribution in Brazil: Golds: Prescorr (1987, as D. glomeratay: Rio de Janeiro: Dias (1985, as D. glomeraia), Dus (1997); So Paulo: Braxce & Neccr (1996a, as D.glomeraia) Associated species: Chlorophyta: Spirogyra spp.: Rhodophyta: Batrachosper- ‘mum ambiguum MoxtAcrt, B. vogesiacum Skits Chactophoraceae in lotic ecosystems from So Paulo State, Brasil 53 1 : Figs 11-19. Morphological features of Draparnaldia and Stigeoclonium species. Figs 11-13: Draparnaldia mutabils. 11 ~ Basal system showing rhizoids (arrow). 2 Plant showing main axis and lateral branches. 13 ~ Detal of main axis with lateral branch, Figs 14-16: Stigeoelonium amoenum. 14 ~ Basal system showing thizoids (arrow). 18, Main axis showing two consecutive short cells bearing lateral branches. 16 ~ Detail of a main axis short cell bearing lateral branches. Figs 17-19: Stigeoclonium farctum. 17 ~ Basal system showing pseudoparenchimatous, sise-like structure, 18 ~ Defail of a pseudoparenchimatous, dse-lke structure. 19 "Erect system showing branching cell similar to other from main axis, {Scale bars: 1.5 mm to Figs 17-18; 100 um to Fig. 12; 50 ym to Figs 11, 13-14; 10 um to Figs 15-16, 19.) sa C.C.Z. Branco etal, Populations examined: AP —Iguape, road SP-222, between km 28-29, Bairro Trains, 24°30°00"S, 47°33°00°W, 08.ix.1993; Juquid, 1 km from BR-116, Ribeirio das ‘Ongas, 24°22'00°S, 47°40'00"W, 08 ix. 1993; HW ~ Apiai, road SP-165, 4 km from Api- ai, 24°32 25°S, 48°49 407 W, 09.1996, Remarks: Several authors, such as Haze (1902), Pkescorr (1962) Sraawanct (1972) and Sarma (1986), used the presence of a distinct rachis and lateral fasci- cle shape as diagnostic characteristics for delimitation of Draparnaldia species. Forest (1956) found great variability in diagnostic characteristics indicated by Hazen (1902) and, not rarely, a single plant could have more than one kind of lateral fascicles and represent more than one species. Thus, Forest (1956) con- ‘cluded that most secimens analyzed represented a widely variable taxonomic unit, named D. mutabilis (Rovi) Bory, ‘Vaw Bren & Snions (1988) described the growth and morphology of some lineage of D. mutabilis in different culture conditions and from nature. The re- sults of their observations showed the same variability previously recorded by Forest (1956), indicating a large overlapping of the limits for several species. Dias (1997) analyzed some tropical populations of Draparnaldia, found no evi- dent limits in fascicle characteristics, and identified all specimens as D. mutabi- lis. In this study a high morphological variability was also observed and ‘corroborated the data in Forest (1956), Vaw Been & Sivons (1988) and Dias (1997), Stigeoctonium amoenum Kevane (Figs 14-16) Phycologia Generals: 1918, 1845 Plants light-ereen, 0.9-2.0 (-3.0) cm long; basal system composed exclusively ‘of profuse mass of rhizoids; erect system well developed, main axis cells differ. entiated in two distinct kinds of cells, long and short; branching regions formed by 2-3 (-4) short cells in a row; long cells cylindrical, 45.0-127.5 um long, 17.5-20.0 um diam, lengtidiameter 2.3-6.1; short cells eylindrical, 17.5 27.5 um long, 10.0-16.0 jm diam.,length/diameter 1-1.6, commonly producing primary lateral branches; lateral branches alternate or opposite, often two branches arising from a single short cell; cells of lateral branches cylindrical, 12.5-27.5 jm long, 5.0-12.5 um diam, lengthdiameter 1.2-1.9; apices of main axis acuminate rarely with multicellular hairs; chloroplast consisting of a medi- an band in main axis cells and laminated in peripherical regions, covering most ‘of the cell wall; oospore formation evident by the presence of multseriate fila- ments Diagnostic features: Plants with basal system formed exclusively of pro fuse mass of rhizoids, raising from the base or rarely from median region of the erect filaments; erect filaments differentiate in two distinct kinds of cells, short and long, the former commonly bearing lateral branches; branching regions often composed ofa row of short cells. ‘Chaetophoraceae in lotic ecosystems from Sao Paulo State, Brasil 5S Table 1. Morphological characteristics of types specimens of four Stigeoclonium species analyzed, Gharacertes —Somoenm ——_Sihdvicum —_‘Siahaccudhon 5 me PROSTRATE STEM, as sytem — hizo prose recat proses Fame anes lament ‘ea indica oindral ———eyindieal_ ined tes bein iS satuiae yGlobular avert present absent stoent ‘eal ERECT SYSTEM. Branching ype oppose, erate, opposite, aerate, a Aching repos: arching revs, kere. Sone fichiomes Gime emer —— Sichomeus i) Branching cells distinc, 2 kinds: simior sitar stint, 2 Kinds imcompursion long tnd son Sing and sb Mainrite preset escat shat presen 1) Malet — present rset ahent preset ©) Shape ofrmsintatshaped cynic slit linat Size ofmain | 20-28 82-120 LHX 268412 6xAO-OK 12-16% 26-58 also ‘vi Shape ofthe eying erin slid linia it rach i) Sie often $1420.40 LOKI 104123850 OHH IB 2 pan cc Sis repeat ameter and gi apne Distribution in Brazil: Rio de Janeiro: Dias (1985). Associated species: Chlorophyta: Chactophora elegans (Roti) C.A. AGARDH, -Microspora floceosa (Vavcuex) Taker, M, tumidula Hazts, Mougeota spp. Spirogora spp.; Chnysophyta: Fragillaria javanica Husteor, Vaucheria bursaia (MOLLER) CA Aoanon; Cyanophyta: Phormidiwm aerugineo-coeruleum (GoMON!) ANAGNOSTIIS. et Kowsnek, P. iriguum (Govonr) ANacnostoss et KowAtex: Rhodophyta: Batrachosper- ‘mum ambigaaum Moxracst, B. macrosporum Moxtacnt, B. puggarianum GH NOW, “Chantransia” stage, Populations examined: AP ~ Paraibuna, secondary road to Redengio da Serra- Paraibuna, 8 km from Paraibuna, 23°20°00"S, 45°37 O(°'W, 18 vii 1993; Ubatuba, Pein ‘zuaba, road Rio-Sanios (BR-IO1), Rio da Fazenda, 23°06°00"S, 44°51'00°W, 17 ii 1993; HW ~ Apiai oad SP-249, 16 km from Apiai,24°23'S0"S, 48°52 38°W, 08 x 1096: Apiai, road SP-280, km 289-290, 32 km fom Apiai, 24°22°30'S, 48°38 11'W, 08 1996, NW — Mirassol, near 1o the road 10 Jaci, Rib Sto José dos Dourados, 20°4°00°S, 49°33'00'W, 1.192, xi. 1992; i, 1992; Sdo José do Rio Preto, road near tothe bridge of the road SP-510, 20°52°00'S, 49°1800°W, ix. 1990; ix. 1990; x. 1990; 1 1991 56 C.C.Z. Banco etal Remarks: [stax (1963) regarded S. amoenum as a well-established species of ‘Stigeoclonium. On the other hand, he commented on a possible difficulty in di ferentiating this species from S. lagelliferum Korzin6, since both produce main axis with long and short cells. Hazew (1902) admitted that S. flagelliferum could be a form of S. amoenum differing only by the lower cell diameter and presence of attenuated or setferous apices in terminal branches. Cox & Bo.o (1966) did not accept S. amoenum as a good species from studies carried out with culture ‘material. They argued against the presence of two kinds of cells in the main axis, «a characteristic considered as diagnostic in the circumscription of the species by Is.aw (1963). Authors from several distinet regions followed Is.w’s concept in identifying S. amoenum, e.g., Stawsanct (1972), SaRMA (1986) and Pristz (1964). In the present study, the populations fully agree with the typical features of S. amoo- ‘num, including information taken from type specimens (Tab. 1). However, Is Law's varieties were not recognized because the set of characteristics used to citcumscribe them were considered to be excessively subjective litle inflated, very inflated, not or little inflated) and broadly variable due to environmental conditions. Stigeoclonium farctum BrxrwoD (Figs 17-19) Nova Acta Kgl. Leopold. Carol. Dt, Akad. Naturforscher 40: 201, 1878 Plants light-green, 0.3-0.5 em long; basal system well-developed, forming a pseudoparenchymatous dise-like structure composed of globular oF cylindrical cells, 10-20 um long, 6.0-8.0 um diam., length/diameter 1.22.7; erect system less developed in comparison to prostrate thallus; erect filaments little branched, ‘main axis cells similar in shape and size, cylindrical, 28,0-40,0 um long, 6.0- 10.0 jum diam., length/diameter 3.0-5.7; lateral branches alternate, opposite ot dichotomous, apices of main axis acuminate, rarely with multicellular hairs chloroplast consisting of a median band in main axis cells and laminated in pe- ripheral regions, covering most of the cell wall; zoospore formation process ev- ident by the presence of multiseriate filaments, Diagnostic features. Plants with well-developed basal system forming typically an evident pseudoparenchymatous, dise-like structure; erect system less developed than the prostrate, main axis cells similar in shape and size Distribution in Brazil: Mato Grosso: Duss (1989); Rio de Janeiro: Diss (1997), Associated. species: Chrysophyta: Pleurosira laevis (Enmexoera) Cowrtne, Vaucheria spp.: Rhodophyta: “Chantransia” stage, Compsopogon coeruleus (Bais & CA. Acanpt) Moxtaci Populations examined: HW ~ Perc 49°57 OW, [8in.1997. near (0 railway station, 23°01'38°S, Remarks: The relation between basal and erect systems in S. faretum are so ex- treme that the prostrate portion of the thallus is the taxonomic character consid ceed as most informative by authors such as Istawt (1963), Prixt2 (1964) and Sanat (1986). ‘Chaetophoraceae in lotic evosystems from Sao Paulo State, Brasil $7 Isawt (1963) attributed more importance tothe presence of a well-developed prostrate system composed of pseudoparenchymatous, dise-like structure, inde~ pendently of little variations in erect system, Cox & Bop (1966) admitted that the circumscription of S.farctum is incompletely understood and suggested stud- ies under controlled culture conditions to establish the variation levels inthis spe- cies. Afler Cox & Bou (1966), several works were conducted with culture ‘material (Franck 1982, Francke & Sioxs 1984, Sivoxs et al. 1986), confirm- ing the presence of predominant basal portion, despite some variations. In Sao Paulo State only a single population of S. farctum was found. Speci- ‘mens showed basal system composed of a pseudoparenchymatous, dise-like structure, in accordance with protologue and description and illustrations of sev- eral authors (Printz 1964, Cox & Bo. 1966, Saxsta 1986, Sivons et al. 1986) ‘Stigeoclonium fasciculare KCraNe (Figs 20-22) Boe. Ztg. 8: 177, 1887 Plants light-green, 0.8-3.5 em long; basal system composed of rhizoids raising from a mass of spherical cells; erect system well-developed and branched; branches opposite or alternate, often forming fascicles near apical regions; main, axis cells cylindrical, similar in shape and size throughout the plant, 25.0— 42.0 um long, 11.0-17.5 wm diam., length/diameter 2.5-6.2; branch cells eylin- drical or slightly inflated, 7.5-20.0 um long, 6.5-15.0 ym diam., length/diameter 11-17; branch apices acuminate; chloroplast consisting of a median band in ‘main axis cells and laminated in peripheral regions, covering most of the cell, ‘wall; zoospore formation evident by the presence of multiseriate filaments. Diagnostic features: Plants with basal system composed of hizoids aris ing from a mass of spherical cells; erect system well-developed and branche forming fascicles near apical regions; main axis cells similar in shape and size ‘throughout the plant Distribution in Brazil: Rio Grande S.glomeratum ‘Associated species: Chrysophyta: Melosira varians C.A. Acanon; Rhodophyta: Barrachospermum ambigiaum Most Ae, B. puiggarianum GRNOW. Populations examined: AF Sio Luis do Paraiinga, toad to Catugaba, Rio do ‘Chapeu, 23°15 00'S, 45°12 00 W, 18.1993: Tapirai, toad SP-79, km 174-175, Rio Cora, 24°05 00'S, 47°37 00°W O8.ix.1993; NW. Sio José do Rio Preto- Masso, ‘Correo do Macado, seeondary road to Mirasol, 20°48 00'S, 49°27°00'W, 23.vi.1995, s.as (1963), KLsenexoPeR (1955, as Remarks: The circumscription of this species has caused deep divergence among specialists. Hazen (1902), based on differences in morphometric charac~ teristics of main axis cells in comparison to. fasciculare, proposed a new and closely related species, 5. glomeratum, with main axis cells much longer than ob- served in S.fasciculare. After analysis ofthe S.fasciculare type specimen, ISLAM (1963) enlarged the morphometric limits of the main axis cells merging S. glo- ‘meratum into S. fasciculare and created a new variety named S, fasciculare var, -glomeratum, characterized by the constant presence of longer cells in main axis, 38 C.C.Z, Branco etal. Figs 20-25. Morphological features of Stigeoclonium species. Figs 20-22: Stigeoclonium fasciculare. 20~ Basal system showing thizois (arow). 21 — Feasciculated erect filaments. 22 ~ Detail ofa fascicle Figs 23-25; Srigeuclonium helveticum. 23 Plant showing well-developed erect system 24~ Basal system showing thizoids arising from the base ofa erect filament.25 Erect, System showing branching cells (arrow) similar to other from main axis, {Scale bars: 100 jum to Fig. 21; 50 pm to Fig. 22; 30 um to Fig. 23; 20 um to Figs 24-25; 10 um 0 Fig. 201) Specimens from Sio Paulo State showed wide variation for this character, and no pronounced differences in main axis cells length were found. Thus, we ‘considered this character alone insufficient to determine varieties and treat our material only at specific level. Saraaa (1986) proposed two new varieties of S. faseiculare: var. henriquesii and var. amrutii, The former differs from other varieties by the presence of fasciculated branches in main axis instead of lateral branches; however, the populations analyzed by us exhibited both types of fasci- cles, thus demonstrating the weakness of this character and, therefore, the incon sistency of the variety, Stigeoclonium fasciculare var. amrutit was defined by differences in main axis cell dimensions and the presence of unilateral branches ‘Chactophoraceae in lotic ecosystems from Sio Paulo State, Brasil 59 Is.an (1963), Prvtz (1964) and Sraveanci (1972) reported wide ranges for cell dimensions inS.fascculare, rising serious doubts about the validity of this va- riety. Thus, the tendency to produce unilateral branches must be analyzed with ‘mote detail to determine if this feature is really sufficient to keep the var. amruti. ‘Stigeoctonium hetveticum Viscnrn (Figs 23-25) Beih, Bot. Zbl. $1: 36, 1933, Plants light-green, 0.9-7.0 em long; basal system composed of thizoids raising from the base of erect filaments, or less frequently from median regions of the plant; erect system well-developed and branched; main axis cells cylindrical, similar in shape and size throughout the plant, 20.5-67.5 um long, 8.2-15.9 (19.0) um diam. length/diameter 1.5-4.9; branches alternate, dichotomous or, less frequently, with two branches arising from a single cel, branch cells cylin- drical, 12.5-27.5 (-30.0) um long, 5.0-10.0 (-11.0) jm diam,, lengthidiameter 1.6-3.7; apices of primary branches acuminate, frequently forming hairs; ehlo- roplast consisting of a median band in main axis cells and laminated in peripheral regions, covering most of the cell wall; zoospore formation evident by the pres cence of multiseriate filaments. Diagnostic features: Plants with basa system composed of thizoidsrais- ing from the base of erect filaments, o less frequently, from cells of median re- gion of the plant; erect system well developed and branched, main axis cells similar in shape and size throughout the plant. Distribution in Brazil: Sdo Paulo: Branco & Nucci (1996a, 1996b, 1997), Necent & Montna (1998), Nicci eta. (1994, 1997), Associated species: Chlorophyta: Chaetophora elegans (Rom) C.A. AGARDM, Microspora floccasa (Vaveusn) Twxtr, Nitella cermua A. Bkatn, Oedogonii@n sp. Palmellopsis gelainosa Koxsinov, Rhizoclonium hieroglsphicum (C.A. Agatti) Kev 'No, Schizomeris leiblein Kcvains; Chrysophyta: Vaucheria spp. Cyanophyta: Colin- Arospermum ct. minutissimum Co.is, Geitlerinemaspiendidum —(Goniont) “Axasnosnins, Lyng allorgel Fess, Microcoleus vaginarus GoNoXt, Phormidium er- ‘ouani Gowons, Pf, jadinianum Gost, P-retit GoMont, P tenue (C.A. AGARDM eX ‘GoMot) ANAGNOSTIDIS et KOMANEK, P.willet (GARDNER) ANaGNostins et KOMARE Rho- dophyta: Barrachospermum ambiguum Mostacne, 8. dlicanulum (Sk.34) Neccut et ES wit, “Chantransia”" stage, Compsopogon coeruleum (Batnis ex CA. ACARDA) Mowtacnt Populations examined: TF Marla, road to Amadew Amaral, 6 km from road 'SP.333, Corrego da Cobra Aguada, 2°20 42°S, 50°02'48 W, 18.vi.1997; Ocaugu, 5 him from Nova Colombia, secondary road to Mariia, Cérrego Santo Antonio, 22°22 13'S, 49°54 "49°W, 17 vii, 1997; Oscar Bressane, I km from the town, Cérrego da Banancira 22°19°M'S,'50°16'45°W, 16.ii.1997; Teodoro Sampaio, road SP-563, km 19-20, 22°21'45'S, $2°04°37°W, 15.1997; Teodoro Sampaio, Copacabana rego da Cachoeira, 2°27 09°S, $2°20'52"W, I4.vi, 1997, Teodoro Sampaio. road SPV-31, km 4, Cérrego do Lajeadinho, 22°29'27°S, 52°11 37°W, |5.¥4,1997: Ch Piraju, Cérnégo Agua do Padre, 3km from road SP-270, 23°07°19°S, 49°25'03°W, 15.in.1997; NW — Américo de Campos, 3 km from the town, Ribeiro das Aguas Paradas, 20°18'S, 49°46'W, 18.v.1993, Dice Reis, 4km da road SP-463, Sidnei Farm, 20°29° 04'S, 50°34'00°W, 16,¥1,1993: Femandépolis, km from the town, Cémego Gatio, 20°16'00"S, 50°12°00'W, 18.1993; Neves Paulista, Barra Dourada, road to Neves Paulista, Rio Sao José dos Dourados, 20°48°00"S, 49°37°00"W, x 1092: x1 1902, 60 C.C.Z. Baanco et al xi1992s iy. 1993; xi. 1993: Orindiuva, SP-322, 500 m from the wn, Cérrego Barrer 20°10°00'S, 49°21 00-W, 17.v.1993: Potirendaba, road to Bady Bassit,Ribeiio do Bor, 20%S8'00"S, 49°25'00°W, in-1992: x 1992; xi.1992: xi1992; Potrendabs, road to Nova Alianga, Ribeitdo do Bord, 21°01 00'S, 49°47 00°W, ix.1992;x.1992; 192: xi 1992; 1.1993, 1.1993 iv-1993; v 1993; vi, 1993; vi 1993; vi 1993; i, 1995. Remarks: Viscurx (1933) proposed 5. helveticum with two varieties, differen- tiated by cell size: var. majus and var. minus. Is-aMt (1963) observed a broader variation in cel size than that described by Viscue (1933) and for this reason did not recognize the two varieties. Cox & Bo. (1966) showed a new perspec- tive for characterization of S. helveticum, transferring the emphasis of taxonomic information from the erect to the basal system, In this scheme, the simple pres- ence of short rhizoids arising from basal cells or from other cells of the filaments, was considered sufficient to characterize this species. Following Ist aw (1963), Cox & Bot (1966) did not accept the varieties originally proposed by Viscures (1933), France & Srwons (1984) included the patter of zoospore germination san important taxonomic attribute in S. helvericum, Thus, they recognized this species as having short rhizoids and erect germination of zoospores, SiMONs etal (1986) used the same information presented by Fraxcke & Sivons (1984) and enlarged the description incorporating features of the erect system (“drapamal- dioid characteristics”), i... differentiation between main axes and lateral branches: chloroplasts of main axis cells more or less fimbriate; lateral branches, frequently originate from short isodiametric cells, often in opposite pairs and branches ending in short pointed hairs, In Sio Paulo State, S.helveticum was the most frequent species found in the sampling sites. In all populations examined the basal system was formed of a 'more or less profuse mass of hizoids. In the same way, the erect system showed cconstant characteristics in most populations. However, the presence of drapar- naldioid features (Siwoxs et al. 1986) was not common, Multicellular hairs oc curred inconsistently in the populations analyzed, showing the variability ofthis attribute. The features that best define this species, in our view, ate the presence of a basal system formed only by rhizoids, erect system very well-developed and cells of main axis similar in size and shape throughout the plant. Stigeoctonium lubricum (Duswyn) Ke Phycologia Generali: 198, 1845 Plans light-green, 0.5-2.0.m long; basal system composed of unbranched prostate filaments and rhizoids, basal cells globulous, 8.0-14,0 pm long, 8.0 10.0um diam,, lengthidiameter 1.0-1.5; erect system well developed and branched; main axis cells differentiated in two distinct kinds, long and short: long cells cylindrical, 30.0-67.5 (-87.5) um long, 8.0-14.0 um diam, length/di- ameter 3.0-6.5 (-9.7}; short cells eylindrica, 12.0-20.0 um tong. 80-10.0 um diam. length/diameter 13-2.5, commonly producing primary lateral branches: branches altemat, dichotomous or with two branches arising from a singe cell rarely forming a row of 2-3 consecutive cells bearing lateral branches succes” Ni (Figs 26-28) Chactophoraceae in lotic ecosystems from Sao Paulo State, Brasil 61 sively: branch cells cylindrical, 15.0-25.0 jm long, 5.0-7.5 ym diam,, length’di- ameter 2.5-40; apices of primary branches slightly accuminate, rarely forming ‘multicellular hairs; chloroplast consisting of a median band in main axis cells and. laminated in peripheral regions, covering most of the cell wall; zoospore forma- tion evident by the presence of multiseriate filaments. Diagnostic features: Plants with basal system composed of unbranched prostrate filaments and rhizoids; erect system well developed with main axis cells differentiated in long and shor, the latter commonly producing prinvary lat- eral branches. Distribution in Brazil: Cearé: [staw (1963); Minas Gerais: Isa (1963); Sto Pau- fo: Istave (1963). Associated species: Chlorophyta: Micrasporaflocensa (Vaucsen) Tier, Mtu- Imidula Hazes, Mougeotia spp, Niellafurcata (Bnuzvan's) C-A. Acaabi emend RD. ‘Wooo subsp. mucronata (A. Bean) Wooo var. mucronata. Oedogonium spp., Schizom= cris leibleimii KovzNe, Spirogyra spp., Tetraspora lubrica (Roti) C-A. Aanoni; Cyan Dhyta: Glocotrichia natans Bosner et Fialiauit, Lyngby majuscula Gowen, Phormidium aerugineo-caeruleum (Gowost) Axsovosnors &t KOMAREK, Schizothrix are- nara (Braet) Gostowr; Rhodophyta: Batrachospermum delicarulum (Sxuis) Necen et Exruistr, °“Chantransia” stage, Compsopogon coeruleus (C.A. AGARDH) MOTAGNe Populations examined: TF ~ Teodoro Sampaio, Copacabana Farm. Comrego da Cachoeira, 22°27 09'S, 52°30's2"W, 14.vi.1997; CE ~ Cerqueira César, Cotrepo da Jacuba, 1 kam from the town, 23°03'11°S, 49°09°18"W, 16,x.1997;Ipaugu road Raposo ‘Tavares (SP-270), km 339-340, 23°05'07°S, 49°32 05°W, 15.ix.1997; NW_ Sto Jost do Rio Preto, Vila Toninko, Rio Preto, 20°50' 00'S, 49°20'00°W, ix 1992: x.1992: 11998; Wv 1993; Sio José do Rio Preto, Engenhciro Schimidt, Rio Preto, 20°S1"00"S, 49°19°00'W, ¥.1993; vi.1993; Turiiba, road. from Buritama w Turitba, 4km fom the town, 20°59°00'S, 50°06'00°W, 14.01.1993. Remarks: Is.as (1963) showed his erticism with the previous studies ofthis species, considering them as insufficient to circumscribe what is actually known as 5. ubricum, In such case, he proposed the adoption of Berri0t0"s (1878) ‘work and his illustration as the starting point of this species. However, the adop- tion ofa late study instead of the original publication diverge fromally from the basic ules of botanical nomenclature. From this point of view, Is.aw’s dicussion referred to another taxonomic entity but nat o S /ubricum ‘careful comparison ofthe type specimen of S lubricum (Tab. 1) with the populations collected in Sao Paulo State was conducted and characteristics from basal and erect systems were perfectly consonant ‘Stigeoclonium subsecundum (KevaNc) Korzine (Figs 29-31) Phycotogia Generals 253; 1849 Plants light-green, 0.8-2.5 cm long; basal system composed of branched pros- trate filaments and rhizoids, basal cells cylindrical, 20,0-30.0 um long, 8.0— 10.0 um diam., length’diameter 2.0-3.0; erect system composed of filaments sparsely branched; main axis cells similar in shape and size, cylindrical, 45.0— 75.0 um long, 8.0-12.5 ym diam., length/diameter 4.5-7.0; branches alternate, dichotomous, rarely with two branches arising from a single cell; branch cells 62 C.C.Z. Branco etal Figs 26-31. Morphological features of Stigeocionium species. Figs 26-28: Stigeoc/onium lubricum. 26 ~ Basal system showing unbranched prostrate filaments. 27 ~ Detail of @ prosirae filament with rhizois (arrow). 28 ~ Erect system showing a short branching cel (arrow), Figs 29-31: Sigeocloniuo subsecundum. 29 — Basal sytem showing branched prostrate filaments (arow)-30~ Detail ofa branched prostate filament. 31 ~ Erect system showing branching cells arrow) similar to other from main axis. [Seale bars: 20 um to Figs 29-30; 10 um to Figs 26-28; 31.) cylindrical, 22.0-37.5 um diam, length/diameter 3.7-6.5; apices of primary branches acuminate, multicellular hairs present; chloroplast consisting of a me- dian band in main axis cells and laminated in peripheral regions, covering most of the cell wall; zoospore formation evident by the presence of multiseriatefila- ments, Diagnostic features: Plants with basal system composed of branched prostrate filaments and rhizoids; erect system sparsely branched with main axis cells similar in shape and size throughout the plant. Distribution in Brazil: First record ofthe species. Associcated species: Chlorophyta: Bulhochacte spp. Schizochlamys gelatinosa A. Brawn, Tetraspora lubrica (Rom) C.A. AcaRDM, Zina spp.: Cyanophyta: Phor= (Chactophoraceae in lotic ecosystems from Sto Paulo State, Brasil 63 ‘midium ef. ret (C.A. Acanea) Gowort; Rhodophyta: Batrachospermum atrum (Huo. Sox) Hanvev, B. delicarulum (Skuia) Necem et EX, “Chaniransa” sage. Populations examined: TF ~ Teodoro Sampaio, Copacabana Farm, Cérrego da Cachocira, 22°27 09°S, $2°20'S2"W, 14.vi 1997; SP —Pindamonhangaba, 9 km from the Horto Floresta, road tothe “Mirante” ~22°43°25°S, 48°27 12 W, 21.x.1996, Remarks: Stigeoclonium subsecundum was considered by Istast (1963) as a ‘well known and widely distributed species, commonly growing in cold and stag- nant waters. stam (1963) and Sawa (1986) described a particular type of in- complete and cylindrical chloroplast in this species. This characteristic was not ‘observed in Sio Paulo State material, Cox & Bot (1966) described S. subsecun- ‘dum with a basal system formed by small branched prostrate filaments of limited ‘growth and rhizoids, differing from Istast (1963) who indicated only the pres- ‘ence of long rhizoids in the basal system. Our observations agrees with the de- scription of the basal system by Cox & Bot (1966). Franekt & Sivons (1984) did not consider S. subsecundum to be a valid spe- cies, because of the close similarity to S. aestivale. They suggested that the pres- ence or absence of rhizoids reflect just the age of the culture and, in that ease, ‘younger isolates do not present rhizoids or they are very sparse typical feature Of S.aestivale sensu (Cox & Bown 1966), and older isolates show rhizoids arising profusely from basal cells (S. subsecundum sensu Cox & Bop 1966). ‘The type specimen was analyzed (Tab. 1) and the morphological characteris- ties of prostrate and erect systems were in accordance with material collected in lotic systems of Sio Paulo State. Cell dimensions were slightly higher in type specimens, but no disjunction was found to invalidate the identification as S sub- secundum, Geographical and environmental distribution The sampling program conducted in streams of So Paulo State resulted in the identification of ten species of Chaetophoraceae, distributed among the follow- ing genera: Chaetophora, Draparnaldia and Stigeoclonium (Tab. 2). Stigeo- 0.05). Macroalgal communities as a whole occurred un- der the following environmental conditions (Neccut et al. 2000): ture 19.7 + 2.9°C; current velocity 48 31 ems: spect S.cm-; turbidity 8 8 NTU; oxygen saturation 66 + 13 % and pil 6.9 + 0.5. 66 CZ. Brawcocet al Table 3. Abundance (means + standard deviation) and frequency of Chactophoraceae spe cies found in streams fom So Paulo State Twa ‘Abundance Frequency Chacipora aenuata Toroo 2 elegans 06203 5 © pforms 09206 5 ‘Total for Chactophora oxsos “ Draparnalate maabiis sper 3 Stigecclontum amram ass40 ‘ S farctun so 1 5 sascewlare hast 3 S heveticam Lse20 “ bri 07403 6 S subsecundum bo 2 ‘Total of Sidgoctonium assess 32 ‘Toad of Chactophoraceas Lwe27 3 Frequency distribution pattems of environmental variables for members of the family are presented in Figs 34-35. Chi-square est revealed similar frequen cy distribution pattems for Chaetophora and Siigeoctonium when compared to the entire family for all environmental variables. No significant corelation among abundance of Chaetophoraceae and stream variables were found, Similar results were obtained considering abundance within each genus. However, some correlations were significant for individual species. C: elegans abundance showed postive correlation with temperature (7 = 0.983, p < 0.001), and C. pi- siformis was negatively correlated with specific conductance (r = -0.868, p < 0.01). Oxygen saturation was positively correlated with abundance of S: amoe- ‘num (x = 0.708, p < 0.05) and negatively with . helvevcum (r = -0.789, p < 0.001), Chaetophoraceae in lotic ecosystems from Sao Paulo State, Brasil 67 Frequency (%) UW Low. fel Hp islam enlace : ] 7 qo fee LL a: [in Seema iie een bution of stream variables in sampling sites with Chactophora= pecies) in Sao Paulo State 70 €.€.Z. Branco etal Discussion ‘Taxonomic treatment ‘The taxonomic study of Chactophoraceae from lotic ecoxystem of Si0 Paulo ‘State resulted in the identification of 10 species from belonging to three genera: Chaetophora, Draparnaldia and Stigeoclontum. ‘The criteria for species delimitation in Chaetophora are traditionally based ‘on features ofthe gelatinous matrix and of the erect filaments branching, and sev- eral taxonomic investigations (Hazen 1902, Prescorr 1962, Privtz 1964, Sanwa 1986, Starstact 1972) revealed this tendency. However, such characteristics are widely variable and show high plasticity in response to different environmental ‘conditions (c. g. current velocity, irradiance and nutrients) and their taxonomic Usefulness is limited (Diss 1997). In the present study, the characteristics of the erect filament system were incorporated to the features of the prostrate portion, providing a set containing the most relevant information of each system and al- lowing a more satisfactory delimitation of the species from So Paulo State ‘Therefore, the three recognized species of Chaetophora (C. attenuata, C. ele- _gans and C. pisiformis) found in study area were identified on the basis of a com= bination of these plants characteristics, as described for each species in the diagnostic features item (see results), Conceming to Draparnaldia, just a single species (D. mutabilis) was found in Sio Paulo State streams, Taxonomic criteria applied to Draparnaldia were based on Forest (1956, 1957) and vaw Been & Simons (1988), which pointed out the fragility of the taxonomic criteria traditionally used for species definition and proposed a drastic reduction in the number of species, Specimens studies were highly variable regarding to theit morphology and could not be splitted in differ- cent taxa, since they showed a continuum for the morphological characteristics. ‘These data corroborate the findings of Forest (1956, 1957) and van BEEN & Si ‘Mons (1988) and suggest that the populations belong to a single species, D. mu- abil. ‘The heterotricheous thallus of Stigeoclonium and the high variation in devel- ‘opment of the prostrate system in relation to the erect one promoted extensive taxonomic divergences that resulted in two oposite taxonomies schemes, Some authors used characteristics of the erect system to delimit species (e. g. Haz 1902, Heexinc 1914, Coutins 1928, Is.aw 1963, Pruvtz. 1964, Srarwcact 1972, Sawin 1986), whereas others only emphasize the features ofthe prostrate system (6. 8. Cox & Bot 1966, Francie & Simons 1984, Sinons etal. 1986). However, Siwoxs etal. (1986) proposed to characterize S, helveticum on the basis of mixed information from the erect and prostrate systems, We found this scheme applica ble not only to 5: helvericum, but also to all species found in So Paulo State streams. This approach resulted in an analysis of the whole plant, which better represented the morphological variability of the species surveyed (diagnostic features adopted for cach species are showed in results). ‘Chactophoraceae in lotic ecosystems from So Paulo State, Brasil 7 ‘Therefore, in this study we extended the possibility of a new classification system, started by Simons et al (1986), to all species of Chaetophora and Stige- ‘oclonium from Sao Paulo State streams, mixing the information from erect and basal systems, which showed very satisfactory results. However, the analysis of a more wide range of specimens from other regions of the world and habitats is necessary for confirmation or expansion of this new approach to the infrageneric taxonomy of the Chactophoraceae. Additional techniques (e. g. molecular and cytogenetic) would be also strongly recommendable in such future studies. ‘Geographical and environmental distribution, Fow studies have focused on the geographical and environmental distribution patterns of Chaetophoracean algae in lotic ecosystems. Most of the available ‘ecological information used for comparisons is included in overall works con- ‘cerning macroalgal communities. ‘The number of Chaetophoracean species (10) found in this study is relatively high when compared to the 17 reported by Swear & Cour (1992) for 1,000 sam- pling sites from several North American biomes. The comparison of species number between the most similar North American region (the tropical rainforest) Sweatt & Cote 1992), with the results of the present study evidenced a higher richness in Sio Paulo State streams (4 vs. 10, respectively). In addition, the sim- ilaity in algal composition was low with only one common species, C. elegans. Species number per sampling site varied from one to two in all biomes/regions, ‘comparable to those from southcentral Alaska (Seats etal. 1986). Chaetopho- racean abundance was also low in the streams sampled, corresponding to the ‘same pattern observed in a previous study (SweaTH etal. 1986), We found no significant correlation of abundance and frequency of Chacto- phoraceae with stream environmental variables. However, on the basis of the en- tire data set, the combination of high temperature and current velocity, intermediate values of specific conductance, oxygen saturation and turbidity and circumneutral pH seemed to be the most closely elated to higher abundance and frequency of Chactophoraceae. Gissow & Wuirrox (1987) described effective growth of Chaetophoraceae species kept grown in culture stimulated by organic or inorganic phosphorous available in the medium. In the present study, no cor- relation of abundance of Chaetophoraceae with nutrients was observed. ‘The evaluation of the genus Siigeaclonium, considering all species together, with environmental variables resulted in no significant correlation, thus sugaest- ing that an overall evaluation for the whole genus is probably inconsistent Results of correlation analysis atthe species level (e.g. . amoenum and S. hel- veticum) pointed out specific strategies of the organisms to deal with the en ronment and reinforce the impossibility to make ecological generalizations on the basis of higher taxonomic entities. Rosewonn & Brawiry (1996) reported that Srigeoclonium (represented by ‘8. tenue) had lower photosynthetic capacity than other algae in high environmen n C.C.Z. Branco et al tal irradiance and low nutrient levels, indicating low competitive capacity, inthis specific circumstance. In Sto Paulo State, 76% of Stigeoclonium records were in open or partly shaded sites (high iradiance) and in low nutrients concentration ‘water (Necci etal. 2000), showing. different trend than observed by Rose MoD & Brantey (1996), Microhabitat data of two populations of S. helvericum (Brasco & Necext 1998) reinforce this tendency and reveal preferential occur rence of tis species under high values of irradiance (1,010-1,235 mol.m-2.s"!). ‘The absence of significant correlation with environmental variables was also ‘observed to the genus Chaetophora and suggests again that environmental ap- proaches must not be considered at higher taxonomic levels. However, some trends were found for individual species. Chaetophora elegans abundance was positively correlated to temperature, whereas for C. pisiformis it was negatively correlated with specific conductance. Different pattems were reported for C. ele- _gans (low temperature, with values commonly under 15 °C) and C. pisiformis (telatively high conductance, > 100 yS.cm-*) (Sieata & Cote 1992). CChaetophora elegans was the most widespread species in Sao Paulo State re- gions, which isin accordance to Star & Cote (1992) who reported tis species ‘curring in seven of eight biomes in North America. However, other well-i tributed species in So Paulo State, including the most often encountered species (S. helveticum), were not found in North America. All Chaetophoraceae species found in Sao Paulo State streams have been re- ported in previous studies for several areas of the world, from temperate to trop- ical regions of norther and southern hemispheres (Hazen 1902, Prescorr 1962, Istast 1963, Painrz 1964, STARMARCH 1972, SaRMA 1986, Exrwisté 1989, SEAT & Cote 1992), The occurrence of Chaetophoraceae in all biomes/regions studied suggests a broad distribution ofthe family members also in a more restricted spa~ tial scale Acknowledgements This research was supported by a FAPESP grant-in-aid (95/2758-8) to ONJ, a CNPq fellowship (14172095-6) to CCZB and CNPq research grants to ONJ (300379/96-2) and LHZB (300128/97-5). We are grateful to Jaws R. COLEMAN, UNESP, for reviewing the English and Mania Hetexa Carapot are forthe help in laboratory analyses, References ‘Atoxso, M.T.A. (1977): Vegetagio.— In: Fundagio IBGE, Geografia do Brasil, 3: Regiio Sudeste.p, 1-118, Serceat, Rio de Janeiro. Brarsoc, G. 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