Rodriguésia 62(4): 867-885. 2011 http://rodriguesia.jbrj.gov.

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Podostemaceae in Southern Brazil
Podostemaceae na Região Sul do Brasil

Anderson Santos de Mello1,3, Aldaléa Sprada Tavares2 & Rafael Trevisan2
Abstract
This study provides a taxonomic treatment of the Podostemaceae family in southern Brazil. Podostemaceae is the largest family of strictly aquatic angiosperms. The center of family richness is the equatorial region of South America. Taxonomic studies are still scarce in Brazil. For southern Brazil there are six genera and 10 species recognized. Dichotomous key and illustrations are presented for species identification. Key words: taxonomy, Malpighiales, aquatic macrophytes, river-weeds.

Resumo
O presente estudo apresenta um tratamento taxonômico para as Podostemaceae da Região Sul do Brasil. Podostemaceae é a maior família de angiospermas estritamente aquáticas com centro de riqueza na região equatorial da América do Sul. Os estudos taxonômicos ainda são escassos para o Brasil. Para a Região Sul do Brasil são reconhecidos seis gêneros e 10 espécies. São apresentadas chaves dicotômicas e ilustrações para a identificação das espécies. Palavras-chave: taxonomia, Malpighiales, macrófitas aquáticas, plantas de rio.
Introduction
Podostemaceae is distributed in rivers in tropical regions of the world and some temperate zones of North America and Asia (Tavares 1997), being the largest family of exclusively aquatic Angiosperms, with 50 genera and approximately 270 species (Philbrick & Novelo 2004). The plants of this family are aquatic herbs, that vary in size, and have unusual morphology for flowering plants. They have simple anatomical features, and generally are very distinct when compared to other groups of aquatic Angiosperms (Ameka et al. 2002). The representatives of this family grow on rocky substrates, in rivers with strong currents. Exceptionally they may occur in lentic ecosystems (Irgang et al. 2003). The life cycle is very singular when compared to other families of Angiosperms, especially because of the need to flower and fruit in synchrony with water-level fluctuations (Tavares et al. 2006). In periods when the rivers have low water levels, some of the plants or even the entire population begins the reproductive process, up to fruit maturity and seed dispersal (Imaichi et al. 2004; Tavares et al. 2006).
1

Due to their form, sometimes like lichens or bryophytes, the position of Podostemaceae has always been controversial (Tavares 1997) and the phylogenetic relationship discussed (Kita & Kato 2001). Different classification systems always had difficulty in relating the family to other taxonomic groups. When recognized as a family, it was initially placed among the monocots (Richard 1816; Martius & Zuccarini 1822). Lindley (1830) was the first to recognize Podostemaceae as a dicotyledonous family. Warming (1888) considered Podostemaceae related to Saxifragaceae. Cronquist (1981) included the family in the monotypical order Podostemales, subclass Rosidae. Cusset & Cusset (1988), studying the African Podostemaceae, established for the family the exclusive class Podostemopsida. Recently, the classification system APG III (2009) included the family in the order Malpighiales, having as a sister group Hypericaceae and affinities with Clusiaceae and Callophyllaceae. The species with neotropical distribution were recently investigated in a phylogenetical study (Tippery et al. 2011) and the clade clusioid, from the order Malpighiales in which Podostemaceae is included, presented for the first time well resolved phylogeny (Ruhfel et al. 2011).

Universidade Federal de Santa Catarina, Programa de Pós-Graduação em Biologia Vegetal, Campus Reitor João David Ferreira Lima, R. Dep. Antonio Edu Vieira, 88040-970, Florianópolis, SC, Brazil. 2 Universidade Federal de Santa Catarina, Centro de Ciências Biológicas, Depto. Botânica, Campus Reitor João David Ferreira Lima, 88040-970, Florianópolis, SC, Brazil. 3 Autor para correspondência: japabio@yahoo.com.br

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Mello, A.S., Tavares, A.S. & Trevisan, R.
Brazil and CTES and LP, from Argentina (siglas according to Thiers 2010). Part of the type material was not analyzed, because it is deposited in European herbaria; however, images of type material available at the sites of the main herbaria were consulted. The morphological characterization of the vegetative and reproductive structures is according to the terminology used for the family. We accepted the synonymization of Van Royen (1951, 1953, 1954), Tur (1997, 1999), Tavares (1997) and Philbrick & Novelo 2004.

Despite the wide distribution and richness of the species, taxonomic studies of Podostemaceae in Brazil are still scarce, although there have been some important recent contributions (Philbrick & Bove 2008; Bove et al. 2011). Tulasne (1863), in the Flora brasiliensis, recognized 15 genera and 30 species. Van Royen (1951, 1953, 1954) did taxonomic treatments for the Americas. The taxonomic decisions of Van Royen represented higher levels of richness and grade of endemism for the South American taxa than those recognized by Philbrick & Bove (2010). Tavares (1997) studied Podostemaceae from some rivers in the Amazon, associating taxonomy with ecological concepts. Aona & Amaral (2006) studied the taxonomy of the family for the state of São Paulo. For the state of Santa Catarina Van Royen & Reitz (1971) indicated the ocurrence of five genera and 13 species. Furthermore, the studies of Pontirolli (1955) and Tur (1975, 1988, 1997, 1999) for Argentina and Paraguay are references for taxonomic studies in southern Brazil, especially because of shared species in the watersheds. New genera and species have being recently described, mainly for Brazil and Argentina (Tur 2003, Philbrick & Novelo 2003, Philbrick et al. 2005, Philbrick & Bove 2008). Bove (2010) listed 17 genera and 87 species for Brazil. The scarcity of collections, few field studies, and unrepresentative and poorly preserved exsicatae in the collections favor controversial opinions on the taxonomy of the family (Philbrick & Novelo 1995). However, some herbaria from southern Brazil have good collections of Podostemaceae, comprising the most of the states of Paraná, Santa Catarina, and Rio Grande do Sul. This study aims to present a taxonomic treatment of the species of Podostemaceae occurring in southern Brazil.

Results and Discussion
Taxonomic treatment
Podostemaceae L.C. Richard ex Kunth. nom. conserv. In: Humboldt & Bompland – Nova Genera et Species Plantarum. 1: 246-247. 1816. Aquatic herbs, submerged, fixed to the substrate by haptera, forming or not cenobia. Leaves with variable size and shape, sessile or petiolate, generally with sheath, linear, lanceolate, palmate, reniform, cordate, obovate, reduced to capillary filaments, or thalloid, margin entire or dichotomically subdivided; uninerved, penninerved ou palminerved, surface smooth, papilose or spinulose, phyllotaxy opposite, spiraled, distichous, tristichous, rosulate or fasciculate. Inflorescence a spiciform monochasium, flowers fasciculate or solitary, axillary, basal, terminal, or extra-axillary, conspicuous or tiny, bisexual, zygomorphic or actinomorphic, 1 to many, generally enveloped by a membranous spathella, rare coriaceous, opening irregularly or wrapped by leaves. Perigone (1)3– 5(many) tepals, sometimes reduced, free, partial or totally fused, triangular, linear, filiform or cordiform. Stamens 1-many, usually alternating with floral segments, free, partial or totally adelphic, forming andropodium or not, filament entire, differently branched, 1 or 2 whorls complete or incomplete, sometimes on one side of the flower, persistent or not; anthers sagittate or rounded, apex emarginate or entire, dehiscence longitudinal, basifixed ou dorsifixed. Ovary superior, gamocarpelar, 1–3 locules, sessile or forming gynophore, ribbed or smooth, pluriovulate, anatropous ovule; stylus 1– 3, free to slightly fused; stigma entire, retuse or branched. Capsule 1–2 locules, septifragal or septicidal, valves equal or unequal, smooth or ornamented. Seeds 1 to many, ovoid to ellipsoid, exalbuminate; embryo straight.

Material and Methods
For the taxonomic treatment we followed classical methodology, including a review of the specialized literature, herbarium material, and field expeditions from 2008 to 2010, in which the main rivers of the all important watersheds in southern Brazil were visited. We collected 60 specimens which were included in the collection of the herbarium of Universidade Federal de Santa Catarina, FLOR. About 300 exsicatae were examined from different herbaria: FLOR, FURB, HAS, HBR, HURG, ICN, LP, MBM, PACA, UPCB, HUCS, from southern

Rodriguésia 62(4): 867-885. 2011

......I.... fr... Bull.. 5.. stamens 2..... stamens 1-many..... subulate. The collection L.... 1.S. 8... (ICN 127....1964. in the states of Paraná... G.... flowers basal. Smith & R..... ending with dichotomous divisions... 11: 90. Marathrum 4’.. solitary . confirmed A....... Fabris & J.. lobes irregular..... Tur 2138 (LP). 5...S..2010..1 Apinagia yguazuensis Chodat et Vischer... São José do Oeste........ flowers always surrounded by spathellae before anthesis...1998... 9: 241... fl. fucoides and A.. 1917..A.IX.. filiform and tiny.2010. leaves without papillae.. somewhat flabellate or pinnatilobed.......I... 4. Santa Rodriguésia 62(4): 867-885...........1964. Smith & R... F. Apinagia yguazuensis was treated by Van Royen & Reitz (1971) as A.606 A).. Nuernberg 584... Rio Uruguai. Soc......... analysis of specimens from Paraguay and Argentina.. Flowers solitary..XII... 3.... Plants acaulescent. Tur 2028 (LP)..... Concórdia.. 19108 (MBM)... phyllotaxy alternate..Podostemaceae in Southern Brazil Key to the genera of Podostemaceae occurring in southern Brazil 1. Tur 2109 (LP). filaments linear. in the present study only A. 28... fr.. 11. fucoides in Santa Catarina. Geneve.. stamens 2 supported by an andropodium .... Borgo et al... 869 Plants with tristichous phyllotaxy.... riedelii (Van Royen 1954..2004.S... fr.. varying in size and shape of leaves... Silva F. Iguaçu and Uruguay rivers.. rudimentary....... Flowers solitary or fasciculate. tepals 3. Mello & A. ovary oblong. Rio Chapecó... 3 ribs per valve.... 1’. fr.... The literature indicates the occurence of Apinagia yguazuensis. 7. inconspicuous....... Guaira.. never tristichous... . Ser.. SANTA CATARINA: Abelardo Luz.. inserts on the same side.. Fruit pedunculate. 584a (FLOR).. 23. Ribas et al... 10 mm long. not forming a whorl.. s. Céu Azul.IV. camping del Nandu.. Apinagia yguazuensis can be considered a rare species in southern Brazil.. Leaves pinnate... associated with new collections from the state.III.. similar to the ovary. Bove 2010) in southern Brazil... Leaves flabellate or pinnatilobed.......... 3. A..... 2. A. two lateral sutures.... Hatschbach 20537 (MBM). Crisci 7146 (LP)...... 1–10 cm long.1996. H. without sheaths.. Salto do Arroyo Tabay. 2a-c Herbs 2–10 cm tall.1998.. This species is distributed in the basins of the Paraná.. XII. O.. leaves rough due to the presence of papillae on the adaxial surface ...... fasciculate ...... terminations dichotomous. 1849..... tepals 2 to many. Podostemum 3’.. A.... smaller leaves with yellow to light green coloration.. 6... Iguazú...IX.... Estância Hermínio e Maria. ser....... 2011 ...... A.XII.........S.. However..... Flowers solitary or in fascicules... 4.... membranous.. Foz del Iguazú.... Flowers always solitary. A. nerves inconspicuous.. ribs indistinct. 710 (MBM)... flowers never surrounded by spathellae. Haas 15881 (MBM).1967. 3 tepals forming a welldeveloped perigone . 3.. according to IUCN criteria.1999.. decurrent. yguazuensis lives in streams of low to medium depth and has narrow stems....... never in spiciform monochasium..... Mello & A... Rio Marmeleiro. Reitz 13927 (HBR) has wider stems and leaves.. 6............ flowers axillary or terminal. 24.... cuneiform. 19.. L. forming whorls complete or not. Observations on populations occurring in the states of Santa Catarina and Rio Grande do Sul showed that A.... Selected material : PARANÁ: Braganey.......... Klein 13890 (HBR)... Guimarães. G......... 1. slightly asymmetrical.IV..M.II...2010.... Flowers arranged in spiciform monochasium. Plants with developed stem. 5.... 1999)...... Leaves 1.V..IX. Figs....... Hatschbach & J.... being proposed as vulnerable (VU) by Philbrick & Bove (2010)....... Rio Iguaçú..... Tristicha Plants with varied phyllotaxy...... Sete Quedas do Rio Paraná. fr. 6106 (MBM)... rudimentary..... However. 2... 6. 3. Sci.. Nuernberg 591 (FLOR)...2000.. phyllotaxy rosulate.. 2.... Mello & A. Parque Nacional del Iguazú. roots simple.. Bot. Stems erect.. fucoides for the state of Santa Catarina and suggested the occurrence of A.. Nat.. Mourera 2’.1968.1967... Wettsteiniola 5’...XII... G.. Additional examined material : ARGENTINA.. MISSIONES: Dep. never forming andropodium. Dep San Ignácio...IX. 1a-e.... Apinagia Tul.. inconspicuous in young fruit........ bivalve. filiform. Parque Nacional do Iguaçu. Bot.. axillary or terminal.I. yguazuensis was confirmed for southern Brazil.. Hatschbach & O. RIO GRANDE DO SUL: Barracão.... nerves conspicuous .. 8.. Van Royen & Reitz (1971) cited A...2–7 cm long. Apinagia 1... in Ann.. riedelli. yguazuensis as the taxon occurring in southern Brazil.. Nuernberg 591 (FLOR). 1. adhered to the substrate forming a basal disk.n... e fr.. guairaensis (Tur. The species shows high phenotypic plasticity.. Smith & Klein 13927 (HBR)... 21.. This morphological variability was also observed by Fontana (2008) within a population...1968...

Catarina and Rio Grande do Sul. Apinagia yguazuensis – a. Hatcshbach & O. & Bonpl. forming laciniae. 2. 25. Rio Chapecó. Leaves opposite or subopposite.2010. b. fl. Hickenia 3(38): 153 (151-156. 3 mm long. 6106 (MBM)). 2 mm long. flower. R. finely flat. recently. Fig. 1808. The genus was previously recognized as restricted to Central America and the Amazon region (Van Royen 1954) and. Arroyo Chimiray.5 mm long. persistent. Flowers solitary. d G. c.IX. and in Argentina. anthers introrse. Marathrum Humb. cuneate. A. d. 7. Rodriguésia 62(4): 867-885.. 2 valves equal.. Pl. 1:39. habit with fruit. y pro 2 cm long.1983. up to 2 cm tall. habit of vegetative specimens. 2d-e Herbs acaulescent. Mello & A.S. apex spatulate. tepals 3. del Apostoles.. 3. had its occurrence registered for austral South America with the description of Maratrhum azarensis by Tur (2003). Tavares. Selected material: SANTA CATARINA: Abelardo Luz. stamens 2. 1 mm 1 mm 5 mm c b a 5 mm 5 mm d e Figure 1 – a-e. Guimarães 20237 (MBM). pedicels 4 mm long. Illustrations: a-b Maria Virgilia. (a-b A. c-e Fatima Zagonel. A.S. Additional examined material : ARGENTINA. in the basins of the Uruguay and Paraná rivers. Roots thalloid.5 mm long. Nuernberg 591. 0. 1. Aequinoct. c. 4 km ao norte do núcleo urbano.I. Nuernberg 584 (FLOR). & Trevisan. spathella when closed forming prominences on the surface before anthesis. Ribas et al. broken spathella funnel–shaped.S. Ovary ellipsoid.M Tur 965 (LP). Capsule 2 mm long. with conspicuous nerves. 2011 . membranous. c. e. each valve with 3 ribs and 2 longitudinal sutures. N. figs.7 mm long. N.V. rosulate.S.I.5–2 mm long. c-e O.. filaments 1.1996. 2. Paraná River. 0.M. fruit. Tur et al 2104 (Holótipo LP). A. MISSIONES: Dep. c. styli 2. 2003. free. base emarginate. axillary. fertile specimens.870 Mello. c. 1 mm wide. It also occurs in Paraguay.1 Marathrum azarensis Tur. 1-3). last leaf divisions 1–2 mm long. Mello & A.

2011 . Apinagia yguazuensis – a.Podostemaceae in Southern Brazil 871 a b d e c Figure 2 – a-c. flowering plants in habitat. Rodriguésia 62(4): 867-885. b-c. plant in habitat. Marathrum azarensis – d. d-e. e. plant with mature fruits. habit.

1992.n. and the only reference is an undated collection by Müller s. 5 mm × 2 mm.Herrera 83 (LP). For the genus Mourera we registered the occurrence of M. Guimarães 19106 (HBR.M. 3. Rio Iguaçú.1996. the genus has 12 species in south Brazil and Van Royen & Reitz (1971) cited the occurrence of seven species in Santa Catarina. In southern Brazil. Tur 2137 (LP). Although Van Royen & Reitz (1971) presented a taxonomic treatment and indicated its possible occurrence in the state of Santa Catarina. 185 (LP). 3. rutifolium Warm. pedicel 1–2 cm long. & Trevisan. nerves prominent on the adaxial face. Nat. 1999). del Ytuizaingó. irgangii Philbrick & Novelo. UPCB).S.III.S. A. N. Am. anthers extrorse. Y. This species was cited by Bove (2010) for the state of Paraná. Mourera aspera has the largest leaves and flowers of the southern Podostemaceae. Santa Catarina. 23.) and added P.5 cm long in ripe fruit. Additional examined material : ARGENTINA. Cabrera et al. rachis 5.5 mm long.Sci. 10. however it was not confirmed.IV. A. The most striking feature of the genus is the presence of an andropodium.H. Parque Nacional del Iguazú. M. Podostemum is the genus with the most species in southern Brazil. According to Van Royen (1954). the taxonomy is still controversial. and P. 1775.XI. far from the border with Brazil (Fontana 2007). 5–9 mm long. Those authors recognized only four species (Podostemum comatum Hicken. In contrast. Guaira. Labiack 3803 (UPCB). This species has a very restricted area of distribution.X. variable size. The monograph on Podostemum (Philbrick & Novelo 2004) used phylogenetic analysis based only on morphological characters and presented a new proposal for the treatments of Van Royen (1954). The collection A. Fl. Guiana.5–17. Foz do Rio Cotegipe. Butura s. Rodriguésia 62(4): 867-885. 2–3 mm. Umana 1 (LP). obtuse or emarginate. ser. Monochasium spiciform. Mourera Aubl. Boreal. Kunyoshi 4288 (MBM).1993.5–4. with remarkable papillae. It is considered to be vulnerable (VU) according to IUCN criteria by Philbrick & Bove (2010). Mourera aspera was collected in southern Brazil in only a few areas in northwestern Paraná. 3. 19. only in the basins of the Paraná and Iguaçú rivers. It is also found in Argentina and Paraguay. Mourera weddelliana Tul. 17.I. Cataratas del Iguazú. N. 11: 93. (UPCB 23990).2006. 1997. The most striking morphological features are the size and shape of leaves. Pl. Flowers numerous.1967. so far.M. in fact. Despite all available literature and the species of the genus being the most common and well reprepresented in collections in austral South America. simple ou branched. 8–10 ribbed. Tavares. Hatschbach & O.. 30.VII. MBM 38666). 9. Sete Quedas do Rio Paraná. basifixed. 3 a-d Herbs.. R.872 Mello. Minas Gerais.n. Ann. fl. pink flowers. 1803.1951.VIII.1 Mourera aspera (Bong. Tur 2124 (LP). G. Silva s.5–35 cm × 3–15 cm.. Espírito Santo. Foz do Iguaçú.M. P. Podostemum Mich. MISSIONES: Dep. this species occurs in the states of Bahia. Goiás.VI.1968. Hist. São Paulo. adaxial surface rough.1996.V. 1849..) Tul. CORRIENTES: Dep.n. which can not be analyzed. P. muelleri Warm. 21. Fr. S. 24. composed of a set of siliceous cells. restricted to the state of Paraná. dorsiventral. pedicel up to 3. Nuernberg 584 (FLOR) in the municipality of Abelardo Luz.2004.S. 4–23 cm long. Parque Nacional del Iguazú.S. In addition to Paraná. Leaves 5. Tur 2036 (LP). Hatschbach 15869 (MBM).1967. in Paraguay. was cited for Santa Catarina (Van Royen & Reitz 1971). rosulate. Parque Nacional do Iguaçú. 3218 (MBM. (MBM).5 cm long. Capsule 3. Mello & A. including five species. Mato Grosso..IV.1994.) Wedd. Dep. Labiak et al. fl. fl. in the inflorescence branches. local forms of species with greater distribution. tepals 5–10. a structure that bears the base of two stamens with anthers. with thick petiole. Marathrum azarensis is a new record for Brazil and it is certainly the rarest species of Podostemaceae in southern Brazil. abaxial face glabrous. A.I. endings filiform. aspera Tul.. 1: 582. although the material referred to by the author had been collected in the Paraná River. G. there are no herbarium records or field observations that confirm the occurrence of the species in that state. P. is in effect the first and only record of the species in Brazil. Iguazú. bracts c. Philbrick & Novelo (2004) justified that the species recognized by Van Royen (1954) are. in the western portion of the state.IV.. The leaves are flattened.. and the rough papillae that cover the adaxial surface of leaves. R. distichum (Cham. 2: 164.. spathella 5–8 mm. Fig.3. Iguazú. coloration generally light green and reddish margin. Van Royen & Reitz (1971) and Tur (1988.I. 11. P. inflorescence a spiciform monochasium. Bot. 2011 .H. 23. N. Marathrum azarensis is hardly recognizable in the field because of its small size and lack of stem. and Rio de Janeiro. 4. the species was collected in rivers of the watershed of the Paraná River.1980. Selected material : PARANÁ: Capitão Leônidas Marques. 21.

RIO GRANDE DO SUL: Cachoeira do Sul. MBM)...7 mm long......... Stipule of only one kind.. 5..... J...1979....... Santa Catarina and Rio Grande do Sul. Passo do Cortado.. Leaves 2–20 cm long.......M.... base of the leaves narrow. fertile 0. Iguazú.. andropodium 1–3 mm long. Parque Estadual do Turvo. J. 5267 (ICN)..1998. Baptista 1353 (ICN.. nat...T.. little branched. Mello & M...... In addition... Divisions of the leaves tridimensional.VIII. fl..... Divisions of the leaves in the same plane... 21(6):149... 4a-d..... visible on both faces of the leaves. The herbarium collections are limited and usually incomplete... but it led to a discussion concerning their taxonomic conclusions. del Apostoles. 3(–5) ribs on each valve......... Paraná.. C.. Podostemum comatum Hicken....266 S......5 mm long.. PARAGUAY. 54°16.5 mm.. Bordas 20242 (CTES. P... stipules amplexicaulis. especially the dimorphic stems.. stipules always with two equal teeth .. 5028 (ICN).. Rev. 2011 .....8–2 mm wide..X.2009... J.. Marmori 2049 (CTES). dichotomously divided...1998.... stigmas entire.... C..L. 58 (CTES). Rio das Cinzas.. Salto Apipé..... 1’.. often dichotomized or composed by leaflets in spiral arrangment. .... 25... 2 valves unequal....1998. stipules consisting of two or more teeth.1982.4... base of the leaves broad. 17. s. stipules with 2–7 teeth of different sizes and shapes ...... the first composed of two ear lobes...... Arroyo Chimiray 10 km al sur de Azara.... apex of endings elongate... C.. 28.... MISSIONES: Dep.....1980... vegetative up to 115 cm long.I... I...c... URUGUAY.... Chilena Hist... Flowers growing on short stalks arising from the roots. Plants with dimorphic stems.. stipules entire. Waechter 1654 (ICN...... Schinini & E. Bonifácio et al.... C.. Ponte sobre o rio Santa Rosa.. symmetric..VIII...1–2 cm long .. treated by Van Royen & Reitz (1971) 873 as Mniopsis Mart...... symmetrical.. Cabrera et al....n........ P.. Dep....L.. with extensions filiform simple or dichotomous.5 cm. 5030 (ICN).. Selected material: PARANÁ: Telêmaco Borba. arising one on each side of the stem.. 1935 (CTES). the second composed of a set of 3–9 teeth subequal..... 4. P.... requiring intensification of taxonomic studies...... not differentiating the vegetative from the reproductive.. Rio Lageado Grande.1917.... comatum 2’ Plants with monomorphic stems.. Philbrick et al.... 14..... with 2(–3) triangular teeth..S.. a population was found with stems over one meter in length. ovary ellipsoid.. AMABAY: Cerro Corá. 3. Stipules of two kinds.. with 2–7 teeth. divisions appearing in two dimensions.. P......... often collected separately and treated as distinct species. N...... linear to filiform.731 O.. 17. asymmetrical. Podostemum comatum is rare in southern Brazil.. Rio Aquidabán Nigui.... Arroyo Sepulturas. Parque Nacional del Iguazú... Capsule 2 mm long... a situation with which we do not agree. Tomazina.. endings of the leaves varied.. Roots 0..5–2 cm long when reproductive... In the municipality of Telêmaco Borba........... Stem dimorphic.. Leaves usually entire. Additional examined material : ARGENTINA..... Dep. A... the lateral asymmetric. A.. 0..... s. P. I.... distichum 1. requiring intensified new sampling. irgangii 3’......5–1..... Dom Pedrito.. 5a Herbs of variable size........ filiform. with a basal triangular tooth.... rounded to acute.. had no confirmed occurrence for southern Brazil. s. Derrubadas....... 1. the species of Crenias were transferred to Podostemum by Philbrik & Novelo (2004)... Tur 567 (LP). Key to the species of Podostemum occuring in southern Brazil 1..... Rodriguésia 62(4): 867-885..Podostemaceae in Southern Brazil This study was innovative. I.3. muelleri Leaves compound. twisted.. During field work it was not possible to collect this species in Santa Catarina.... Podostemum comatum is distributed in the states of Paraná.... 2–115 cm long when vegetative and 1....XI....... inserted above the petiole .. A.. Paraguay and Uruguay. with variable morphology..L..2– 1. 2. pedicel 0. Vila Lageado Grande.. Ituzaingó. The genus Crenias Spreng....1983.... Silva et al.. Pinheiro Machado.. 1–2.2007..... CORRIENTES: Dep. Philbrick et al.... petiole flattened. CTES).T. del Artigas.. C. 2.. a fact never reported before in the literature.... 31°13.T.......... Figs... It also occurs in the state of São Paulo and in Argentina..... 1–1.d. (LP). 3. HAS). ALTO PARANÁ: Caballero. Philbrick et al. endings of the leaves spatulate or rarely acute. despite its wide dispersion area in the three states. visible only on one face of the leaves ....... 1980. Waechter & L.... fr. Beretta 553 (ICN)... tepals 3. rutifolium 4’.. 6038 (MBM)..M.IX.

I. B. Sete Quedas do Rio Paraná. 26. 3–4 ribs per valve. Monomorphic stems. Rambo 54828 (HBR. G.4 mm long. Canela. A. hard and darkened in older stems. 1873. A. Sehnem 4946 (HUCS. 17: 73. S.d. 2011 . Osório. Silva et al. RIO GRANDE DO SUL: Bom Jesus.1998. Guimarães 20526 (MBM).1941. J.S. 7.S. 31. 2 lateral and 1 at the bifurcation of the andropodium. Salto Morato.I. Cascata dos Venâncio. A. stigmas entire.II.C. Prodromus. 17. Fazenda do Posto.3 mm long.XI.1942. Linha Campestre.) Wedd. Passo do Inferno. fr. 0.. 2 .2007. 10. (ICN 8663). 2 or 3 axes from which arise leaflets. Vila Seca. 0.472S. 6a-c Herbs. J. stem with leaves.XII. apex acute. persistent. when verticillate.S. 8877 (UPCB).2009.667’O.II. Silva 56124 (MBM).2–1. older rigid. & Trevisan. Cachoeira da Mariquinha.. papillae at the tip of the leaves. andropodium 0.9 mm wide. C. 50705074 (ICN). Montenegro. Roots prostrate. F. Tavares. Espumoso. Ponta Grossa. 14. 12. Hatschbach 42235 (MBM)).6–2.X. (PACA).1934. G.2008.F. S.M. 6–8 ribs. Irgang s.J. 52°42. Rambo 9027. 0. Hatschbach & J. General Carneiro. 50°14.XII. inserted perpendicularly on the stem axis. d.VI.3–5.X. b.V.n. 6. dark and twisted. PACA).. 0. last divisions of leaves in V or U shape. Illustrations: Fatima Zagonel.2006.2–15 cm long. R. 29°30. 2 cm 2 mm a b 3 cm 1 mm c d Figure 3 – a-d. PACA).J. fr.T. distichous. 16. Tibagi. Podostemum distichum (Cham. Cervi et al.171’S. s. 5b-d. C. Rodriguésia 62(4): 867-885. flower.T. Jaquirana. tepals 3. Kinnupp 2892 (ICN). ovary reddish. variable forms. 25. Cachoeira do Rio Azul. (G. 2181 (MBM).4–600 mm long. 2.1991. Guaira. Rio Butiá. A. 28°50. when dichotomous. Rio Nene. Mello & M.1968. Philbrick et al. compound. Mourera aspera – a.. triangular. Selected material: PARANÁ: Céu Azul. stipules emerging as an extension of the sheath navicular-shaped.556O.2 mm long. Estrada de Cambará para Jaquirana.1971. Capsule bivalvar. Beretta 555 (FLOR). 6342 (MBM). 2–8 dichotomic divisions emerging in three dimensions. Rio do Engenho Velho. 0. Leaves peciolate. Lindeman & B. 2–7 teeth of different sizes. lateral larger than the central. inflorescence.1950.III. c.I. Cachoeira Salto Morato. Lavras do Sul. 5187-5188 (ICN). Caxias do Sul.05–0. Rambo 4827 (PACA). Cerro do Umbú. flattened. Maquiné. V.M. A. Philbrick et al. Figs.874 Mello. Flowers 1–5 per stem.C. Hatschbach & O. Barbosa et al. corredeiras. 6.

J. (a A. A. fl.M.IX.846S. Herval do Oeste. 28°08°43000S.. 12. Klein 6110 (HBR). Rio do Oratório. Soledade. 12 km da cidade de São Joaquim. Sehnem 1339 (PACA). fl.1941. Klein 8491 (HBR).2010.L. Urubici. 9. SANTA CATARINA: Abelardo Luz. Rio do Sul.. b. J.1958. Rodriguésia 62(4): 867-885. São José do Cerrito. Mello & M. Mello & A. Nuernberg 573 (FLOR).III.416O.1965. 30. S. Nuernberg 587 (FLOR). d. L. L. flower. Nuernberg 582. Rambo 34957 (HBR. 5003-5007 (ICN). R..1998. Passos Maia. Bom Jardim da Serra.. e fr. A. Podostemum comatum – a. 5. 28°36.1963. Rio Gueira. 5055 (ICN).1956.000O. roots with fertile stems. Lauro Muller. 52°36.T. d Maria Virgilia. Vacaria.S. 2011 1 cm . Schinini 38352 (CTES). Passo dos Fernandes. I.1947. 5. b-d Silva et al.6036. Rambo 4748 (PACA).IX. Mello 557 (FLOR). Mello & A. A. C. Alto Matador. Corupá. Smith & R.1991.S. A. Campos Novos. Santa Cruz do Sul. S. roots with vegetative stems. Reitz & R.M. fl. fr.2010. A. R. fl.II. Illustrations: a-c Fatima Zagonel. Waechter 2508 (ICN.. Reitz 14333 (HBR). A. Vale do Sol.2010. 20. 17. terminal portion of the stem. Reitz & R.IX.J Rambo 1258 (PACA).I. 52°30.1965. Gasper 2089 (FURB). Vale Vêneto.M. Nuernberg 581.I.6038 (MBM)).2009.2009. Krapovichas & A. 581b (FLOR). 581a. 578b. 5. R. Fazenda da Ronda. 6. PACA).1959. A. 3. HAS).Podostemaceae in Southern Brazil 875 10 cm 1 cm a 2 mm c d b Figure 4 – a-d.L. Reitz 14189 (HBR). 10. Mello & A.II. S. 29°40. 3.2009. São Francisco de Paula. A.IX.III.2010. Philbrick et al. 582a (FLOR). Philbrick et al. A. Cascata 5. 8. 578a. 7.IV.J. Ponte Baixa. C. est.V. 4. Rio Irani. Lajes. Smith & R. Mello & A. São Joaquim. 49°37’06.I. Beretta 553 (FLOR).592O.. Klein 16177 (HBR).798S.T. Mello & A.S. c. Reitz & R.S.XII. 578c (FLOR). 13. Rio Francisco Alves.S.S.S.XII.IX. fl. Nuernberg 578.

leaves with verticillate leaflets. including morphological and molecular characters. verticillate or not and stems short or long. Podostemum comatum – population with stems up to 115 cm long. it is necessary to intensify work and it will probably be given new taxonomic positions. with the greatest dispersion area in southern Brazil.O. Salto Grande. 30. comatum. stems with short leaves. schenckii and P. N. Although we are accepting the proposal made by Moline et al. R.1967. with little. R. muelleri. & Trevisan.876 Mello.II.1975. Cainguás. distichum. P. However. It also occurs in Argentina. as well as São Paulo and Minas Gerais. CONCEPCIÓN: Rio Ypané.M.IX. the phylogenetic approach of Moline et al. 6 Km de Belen. Podostemum distichum – b. Podostemum distichum is widely distributed over the three southern states. P. Rodriguésia 62(4): 867-885.1985. 2011 . a b c d Figure 5 – a. It differs from other species by the leaf divisions arising in three dimensions. Salto las Golandrinas.S. Tur 955 (LP). Tavares. arguirense.n. ENTRE RIOS: Dep. (2006). Podostemum distichum is the most common species. A.. occurring preferentially in waterways on the South Brazilian Plateau. The analysis of collections and field work were crucial to note this complexity. (2006). indicated that the proposed complex should be carefully studied. Philbrick & Novelo (2004) synonymized P. c. 18. Wolf s. Zuloaga 559 (LP). flower with recently split spathella. In Brazil. A. rutifolium and Tristicha trifaria. in the basins of the Uruguay River and Paraná rivers. In field works we realized that it is common at higher altitudes.S. P. atrichum. P. Additional examined material : ARGENTINA. PARAGUAY. d. b-d. 10. distichum has the highest morphological variability within the genus. MISIONES: Dep.X. F. P. in Paraguay and Uruguay. (LP).or much-divided leaves. Concordia. warmingii under P. glaziovianum. P. forming dense communities with P.

hard. 0. Haas 1151 (HBR). 1–5 cm long. dark in older stems. persistent. andropodium 0. 6–11 leaflets per whorl.7 mm.IV. Podostemum irgangii C.5 mm. linear. Rio Marmeleiro. protruding 0.S.. A. 2001.7 mm × c. 26. Rio Canto do Galo. 5–14 whorls per leaf.. whorls uniformly arranged. RIO GRANDE DO SUL: Barracão. second type appearing above the insertion of the petiole. 6d-g.. e. 0. Mello & A. 1–42 mm long. Monomorphic stems.5 mm long. fl. e fr. d-g A. divided. 2 lateral.Illustrations: Fatima Zagonel 3. Roots prostrate. Mello Rodriguésia 62(4): 867-885.1968. (a-c A. G. 0. fruit. symmetric. perpendicular to the axis of the stem. Hatschbach 19182 (MBM).2 × 0. subamplexicaulis. Novon 11: 92. hard.4–0. stigmas 2.6 mm.7–1. fr.6 mm from the stem. entire. f. Capsule 1.2 mm long.2–0. 0. first type located at the base of the leaf insertion on the stem. Podostemum distichum – a.6–2mm long and 1 at the bifurcation of the andropodium.1– 7. 4. Ovary 0. with two lobes. Figs. 7a-b Herbs. Flowers 1–7 per stem. sometimes curved. T.1 mm long. roots with stems and leaves. J. Laranjeiras do Sul. divisions of the leaves verticillate.1 mm × 0. 2011 . solitary. rounded.4–1. leaves and flowers. Novelo. stipules in two forms. Leaves distichous.4 mm long. tepals 3. leaf. g.1 mm wide. Podostemum irgangii – d. stipule detail.Mello & A. c. d-g.IX. peduncle 1–6 mm long. 0. Nuernberg 581 (FLOR).Podostemaceae in Southern Brazil 877 2 cm c 2 mm 5 mm a b 2 mm e 2 mm 1 mm d f 5 mm g Figure 6 – a-c. teeth 4-9.4–1.2010. flower. roots with stems.S.2–0.2–2. 3. stems with leaves and verticillate leaflets. base symmetric.5 mm long.9–1. Nuernberg 587 (FLOR)). Selected material: PARANÁ: Clevelândia.S. 1.1966.8– 2. 3. fl. b.V. prostrate or erect.6 mm.2–2.8–1. Philbrick & A.C. Rio Chopim. apex acute. Petioles 0. Lindeman & J. 0.3–1.

left arrow line corresponds to leaf stipule. & Trevisan. Podostemum rutifolium – roots. leaves with characteristics broad-base. A. c-e Podostemum muelleri – c. two kinds of stipules..S. stem with leaves. f. population in the habit. b.878 Mello. a b c d e f Figure 7 – a-b. d. Rodriguésia 62(4): 867-885. Tavares. Podostemum irgangii – a. flower. 2011 . e. A. stems and leaves.S. R. right arrow line corresponds to stem stipule.

IX. W. some individuals had smooth stems and the stipules were inserted only above the petiole. Céu Azul. ser. C. Salto Grande. Podostemum irgangii usually has stems and leaves darker and harder.I. Philbrick et al..1998..S. 3. Maquiné.9 mm long and 1 at the bifurcation of the andropodium 0. Podostemum müelleri presents wide distribution in the southern states of Brazil. 480 (t. 25.7 mm × 0. Itapiranga. 6–8 ribs. Additional examined material : ARGENTINA. Nuernberg 583 (FLOR).. 5146-5148 (ICN). 4: 445. 8.T. 1888. e fr.1964.1998.1– 34 mm × 0. Podostemum müelleri is recognized by the enlarged base of the leaves and the stipules consisting of a tooth and only visible on one side. Rio Castelhanos. Smith & R. Santa Maria. stipule simple.. Podostem. 26. RIO GRANDE DO SUL: Bom Jesus. Tur 1309 (LP).XII.S. distichum may be grossly mistaken. A. Nuernberg 589 (FLOR). P. stipules and uniform verticillate division of the leaflets were not present in all populations analyzed. It was considered to be vulnerable (VU) by Philbrick & Bove (2010).2004.. Klein 11874 (HBR). VII.1998. G. G. 3–4 per valve. Quedas do Rio Chapecó. Prudentópolis.3 mm long. Nuernberg 579 (FLOR). Mello & A. Mello & A. Arroyo Garabi. fl. 3.1957. fl. C. Although some populations of Podostemum irgangii are easily identified. A. fr. Nuernberg 574 (FLOR).Podostemaceae in Southern Brazil & A. Passo do Rio Canoas. Nuernberg 580b (FLOR). A. entire or with 1–4 divisions dichotomic. Cataratas do Iguaçú. 1888 Warm. Mello & A.IX.IX.1998. However. Reitz 9290 (HBR). C. Klein 14432 (HBR). Philbrick et al.VII. L. Trepolo & S. Reitz 9888 (HBR). Parque Nacional do Iguaçú. Mello & A. Philbrick et al. fl. 5. identification of Podostemum irgangii and P. asymmetric.I. São José do Cerrito. Reitz & R.M. São Francisco de Paula.1956. 3. base of the leaves flattened.1–3 mm long. I. 16. Campos Novos. pl.2009. as wel as São Paulo and Goiás. 3. Podostemum irgangii is the only species of the genus endemic to southern Brazil. Nuernberg 577 (FLOR).J. Skr. Tiller 281 (MBM).V. SANTA CATARINA: Abelardo Luz. 26. A.2010.S.III. Mello & A. Nuernberg 574 (FLOR).2010.2010. Rio Irani. Rio Capetinga.IX.S. I.IX.9 mm. MISIONES: Dep Eldorado. Reitz & R. irgangii has two forms of stipules and its leaves are usually evenly verticillate. Rodriguésia 62(4): 867-885. indicates the need for increasing collections. O. 3: 1. São José do Cerrito.5–1. Ijuí. Tur 954 (LP). Nuernberg 576 (FLOR). Rambo 60293 (PACA. R. mainly in rivers when associated with Podostemum distichum. Ponta Baixa. forming a visible tooth only on one side. 16-17). 5. blades linear to spatulate. est. 9. 17.2010. Danske Vidensk. Riozinho.1998.2009.II. asymmetric.XII. A.2010. Rio Tafona. C. Arroyo Aguará Mimí. Mello & A.I. Philbrick et al.1972. G. C. fl. 22.S. 5200 (ICN). 19. tepals 3. 10.IV.1–1. 6– 8 ribs. Among the criteria used for diagnosis of the species.II. Hatschbach 31061 (MBM). Mello & A.I. 6. Selsk. Some populations have one or more divisions of the leaves in which the leaflets are inserted irregularly.I. S.9–1.T. Reitz & R. C. It occurs only in rivers located in the western portion of the Araucaria Plateau..3 × 0. It also occurs in Argentina. I.. Figs. Selected material: PARANÁ: Candói. ENTRE RIOS: Concordia. L. Guarani. C.. the species is in serious danger of extinction.5 mm. 0.M. 9. Joaçaba. wider than the blade. A. A.. Klein 15097 (HBR). Cambará do Sul. 5010 (ICN). andropodium 0. R. 2011 . I.T.S. Capsule 1–2. 5181.X. Arroio Salto Anita. Mello & A.1963. SANTA CATARINA: Abelardo Luz.8 mm. CORRIENTES: Dep.T. Quedas do Rio Chapecó. Foz do Iguaçú.T. Afd. Flowers 1–10 per stem. 2 latera 0. Rio Conceição. San Tome. 5.S.IX.3–8mm wide.R. Caçador. A. Leaves usually sessile.I.1998.S.. Rio dos Patos.1962.2010. Estancia Garruchos. divisions dichotomic emerging in two dimensions. e fr. in communities where both species occur together. A. Smith & R. 31. Nuernberg 586 (FLOR). Hatschbach 11219 (MBM). simple. Rio Irani. 0. Mello & A. 5208 (ICN).1070. Dep. Mello 569 (FLOR). Herval do Oeste.IV. Dep.S.S.IV. 5182 (ICN). Philbrick et al. A. Deginari 1145 (LP). 22. Pedersen 9236 (MBM). Ibicuí.1967.. Philbrick et al. the morphological variability found. est.2010. Roots prostrate. e fr. Monomorphic stems. stigmas entire.XII. Beira do Arroyo Yaboti.1962. fl. Herval do Oeste. Rio Irani. Prainha. Philbrick et al. 10. Math.1969. Another striking feature is the even distribution of the verticillate whorls. Fam. 5120-5130 (ICN). R. Nuernberg 592 (FLOR). Passos Maia. N. Keller 125 (CTES).M. 6133 (MBM).1998. Anita Garibaldi. Blumenau. Philbrick et al. Likewise. Due to the large loss of habitats and the small area of distribution. Rio das Antas. 3. In the present study it is recorded for the first time in Rio Grande do Sul. fl.2010. 3. HBR).8–2. C.1998. Montecarlo. entire or slightly emaginate flattened. Rio Vacacaí.2–1. Rio Jordão. flexible or hard. 7c-e Herbs. 5189-5191 (ICN).S. I. Passo dos Fernandes. Naturvidensk. Campo Erê. Rio Uruguai. ovary 0. Moreover.T.1999. persistent and hardened in older stems.T. 879 Podostemum irgangii is the unique species of the genus with two kinds of stipules.6–1. Rio Jacutinga.IV. obliquely inserted on the stem. Anitápolis. which compound the basins of the Iguaçú and Uruguay rivers. Kongel. 5050-5052 (ICN). 38.1957. Cascata do Arroio Conduto. Ribas et al. 3. Paraguay and Uruguay. 27.T. 17. Herval do Oeste. Rio da Prata. Rio Forqueta. Podostemum müelleri Warm. 4. 21. e fr. fl.

Capsule 1. Leaves distichous. Podostemum rutifolium Warm.S.7 mm long and 1 at the bifurcation of the andropodium 0.e. Podostemum rutifolium – a. obtuse or acute.1 mm long. 8– 10 ribs. tepals 3.S. 2 teeth triangular and flattened.4 mm long.6 mm. 7f. 0. hardened and darkened in older stems.B. leaf with stipules. apex acute.6–1. Selsk. e-f. small. Guimarães 345 (FLOR)). 8a-d Herbs. Tristicha trifaria .01–0. Figs. fruit. Skr. d. Monomorphic stems. 1. base of the leaves symmetric. 0. ovary reddish pink.1–94 mm long. Mello 4. persistent. leaf. 1 mm 1 mm a b 1 mm c 3 mm d 1 cm e 1 cm f Figure 8 – a-d. ser 6. 8–10 ribs.05 mm long. b. stipules 0.S. (FLOR). (a-d A.5–20 mm long. 9 (2): 127.2 long. 0.5 mm. 2 lateral 0. leaves and closed spathellae. f. Vidensk.1 × 0. T. Roots prostrate. spatulate. 4–5 per valve.880 Mello. terminal portion of the stem with flowers. 0. R. flattened. c. 5.4 mm wide. 1899. 2011 . & Trevisan. entire to 5 times dichotomized or lobed. apex rounded. Tavares. Flowers 1–5 per stem.2– 2. stigmas entire.4–1.9–2. roots with stems. A.8–1..4–2. 0. stem with flowers.2–3.2–1.3 mm × 0. Rodriguésia 62(4): 867-885. A.

dehiscence introrse. G. In the present study we do not consider any of the subspecies. N. 881 family. and Uruguay. Arroyo 11 vueltas. tricarpelar. Smith 9675 (HBR).1968.1997. 3.650S. A.1957. Rio Uruguai.XII.2004. mainly in rivers in the states of Rio Grande do Sul and Santa Catarina.XI.L. stamen 1.1973. Smith et al. L. 3220 (MBM).XII. Dionísio Cerqueira.1070. 13.1998. connate at the base. 53°59W. 5021 (ICN). 29. fl. Parque Nacional del Iguazú. Recent works on the biogeography of Tristicha (Kita & Kato 2004) suggested that this genus had originated in Africa and dispersed to other continents.I. Guarani.S. 50°02.580O. Concórdia. e fr. seeds minute. Herbarium colletions are very often mixed mainly with Rodriguésia 62(4): 867-885. G. Pouso Redondo.C.S. 5135.194S. 5. A. fr. Prainha. 26°52. Mello & A. Cabrera 10854B (LP). Passos Maia. 10. 22.n.IV. Rio Quaraímirim. 7. A. 5. Ibirama. São Lourenço do Sul. 5207 (ICN).. 27°17. Philbrick et al. 9. Deginian 1143 (LP). Tur 1318 (LP). 49°13. 5144. Additional examined material : ARGENTINA. Arroio Forquetinha. solitary or not. Branches prostrate.2–15 cm long. Santa Cruz do Sul. 7. inserted in opposite way along the roots. It also occurs in Argentina. 20. 50°26.1967. following a reverse path of the other species of the family. Nov. Hatschbach & Haas 15868 (MBM). Philbrick et al. ricciforme to Central America. Blumenau. Mello 5 (FLOR). 8e-f Herbs with branching roots.830O.I. O.1998. Tlacochistlahuaca.T.1976. 1806. AMAMBAY: Parque Nacional del Cerro Cora.1994. Riozinho. 5134. Nuernberg 578 (FLOR). rutifolium: P.. recognized by Engler (1930) and considered by many authors as a different Tristicha trifaria has received little attention from collectors.2010. blade spatulate. 1981.1956. Rio Moraes. The stems usually are diminute.II. J. (LP). 3.IV.1974. C. fr. Rio Timbó.1998. Paraguay. Madagascar. Rio Peperi-guaçu. laxly or densely arranged along the stem. Smith & Reitz 9913 (HBR).. Philbrick et al.426S. abajo del Salto Bosseti. I. Nuernberg 580b1 (FLOR). Ponte Baixa. occurring in austral South America and P. (ICN). 29. Dep.II.. P. 5025. apex obtuse. 24. RIO GRANDE DO SUL: Candelária. Irgangi s. ovoid to ellipsoid. fr. XI. est. SANTA CATARINA: Abelardo Luz. Rio Ponta Grossa. 19. Guaira. Veg. base obtuse. II.. 1824.IV. J. 16: 1-22. The stems and leaves are prostrate. 5026 (ICN). C.IX. 0. 21. 21.IX. Hatschbach 19130 (MBM).2009. 3219 (MBM). Labiack et al.1996. C.II. Additional examined material : ARGENTINA. Florentino Ameghino.n. est. fr. trilocular. J. rarely lobed.III. 5. 22. 13. because it is characterized by the absence of spathella before anthesis and by having whorls that involve all floral parts. I. stigmas 3. anthers oblong..2009. possibly due the diminute size and the appearance that resembles a bryophyte. numerous.XI. Rio Uruguay. Céu Azul. ENTRE RIOS: Concordia. rutifolium.1951.S. Corupá.1976. 5169-5176 (ICN). P. Parque Nacional do Iguaçú. membranous. GUERRERO: Ometepec. 17..n. RIO GRANDE DO SUL: Bom Jesus. Passo dos Fernandes.S. Federación. Rio Bruaca.1956. 29°29.c. Philbrick & Novelo (2004) propose two subspecies for P. Rio Iguaçú. XII. Sete Quedas. Salto grande. 22. (ICN). PARAGUAY.XII. Arroyo Yaboti 27°08’S.L.H. 7451 (MBM). Labiack et al. rutifolium subsp. MEXICO. Sete-quedas do Rio Paraná.T. 23. Both subspecies are differentiated only by leaf size and disjunct occurrence. 21. Flowers axillary or terminal.T. São José do Cerrito. fl. s. L.M. 4.1 Tristicha trifaria (Bory ex Willd.2010. 22°39’S. 19/III/2004.J.: 3. XII.I. Quaraí.T.I. Lajeado. 5160-5162. Tur 2127 (LP). Brack s. it forms dense populations. 50°03O. fl. Girardi & B. Philbrick et al. rutifolium subsp.. A. surrounded by membranous leaves. Waechter 227 (HAS). Solomon 7141 (LP).1956. Laguna dos Patos. Syst. Herval. Podostemum rutifolium is recognized by very small leaves. tepals 3.T.III. apex truncated. Philbrick et al.n. adhered to the substrate. hidrelétrica do Rio Timbó.) Spreng. Tristicha Thouars Gen. San Javier. Beretta 551 (FLOR). Mello & A. Arroyo Piray Guazu.IV.H. Smith & Reitz 9047 (HBR). s. paso Kalesplaner. Philbrick et al. 2011 .599O. Rio Iguaçu. ovary sessile. ed.2009. Selected material : PARANÁ: Capitão Leônidas Marques.1995. rarely acute. Nuernberg 588 (FLOR).T. Rio da Prata. trivalvar dehiscent. Iguazú.XI. Fig. C. Eldorado. Nuernberg 572 (FLOR). 7589 (HBR). C. Philbrick et al. entire. Passo do Rio Pardo para Candelária. Neiff s. In areas where it is dominant. since leaf size is not a significant character in the circumscription of taxa in this family.XI.2010.1998. sessile. 5091-5094 (ICN). Podostemum rutifolium is common in southern Brazil. Mello & A. 5141. P. 5140. Salto Yucumã.S. Rio Itajaí do Norte. 5183 (ICN). SANTA CATARINA: Biguaçú. (LP). Passo do Rio Pardo.Podostemaceae in Southern Brazil Selected material : PARANÁ: Capitão Leônidas Marques. Mello & M. A.IX. 7.II. Caçador. A. attached or floating. Ribas et al..1998. Capsule similar to the ovary.S. A. J.2004.I. Cerro Azul. 19. MISSIONES: Dep. The genus belongs to the subfamily Tristichoideae. Guaira. Mello & A. A. Leaves with phyllotaxy tristichous.S.1998. Derrubadas. A. 5145 (ICN). MISSIONES: Dep. Dep. Tur 2080 (LP). C.1982. Waechter 225 (ICN).

b.882 Mello. 2011 . leaf. Tavares. d. R. c. Rodriguésia 62(4): 867-885. habit. Wettsteiniola pinnata – a. flowers emerging in fascicle. & Trevisan. Illustration: Fatima Zagonel.. (a-d Hatschbach 42236. flower. 1 cm 1 mm b a 4 mm c 2 cm d Figure 9 – a-d.S. A.S. A. 15871 (MBM)).

The genus presents affinities with species of the section Hymenolacys of the genus Apinagia. The only collections for the genus Wettsteiniola were conducted in rivers with large waterfalls. Botanical Journal of the Linnean Society 139: 255-273.. Regni Veg.5 mm long. submersa (Podostemaceae: Podostemoideae) from Ghana. Leaves bipinnate. Selected material : BRAZIL. Bove.jbrj. Fig.L. irregular. a revision of the genus may include W. 6037 (MBM).). In addition.gov. 2011. accorsii and W. G. 2009. petiole 4–5 cm long. 9a-d Base or root thalloid. styli acute. Tristicha trifaria is widely distributed in southern Brazil. Spec.M. 6. rachis flattened. Brittonia. Flowers in fascicules. secondary divisions repeatedly furcated. C. pinnata. in Rio Paranapanema. we found specimens from the extinct “Sete-quedas” of the Paraná River. 1992.1 Wettsteiniola pinnata Süess. Salto Grande.G. Taxonomy. so the expedition was unsuccessful. Pfeifer E. stamens 1–4. Acesso em 21 Out 2010. Regni Veg. V.VIII. 1935. the Programa de PósGraduação em Biologia Vegetal of UFSC. In addition to the type. São Paulo. located in the Paraná River basin.I. peltate. northwestern Paraná. Bove. Salto Cavalcanti.. 39: 18. C. & Costa. Podostemaceae In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. W. anthers up to 1mm long. Philbrick. Brummit. membranous. Nov. (orgs. C.. Pp. which drains into the basin of the Paraná River. 24.2007. with 12 to 14 ribs.. 2 mm long. distribution and emended description of the Neotropical genus Lophogyne (Podostemaceae).S. first division of the sheet up to 1. In: Wanderley. occurring in southern Brazil only in the state of Paraná. 2010. Developmental morphology of Saxicolella amicorum and S. pedicels up to 3. 2 mm diam. 2006. pinnata and W. & Powell. 5–10 mm diam.K. References Ameka. 2002. occurs in the state of Paraná. stipules up to 3 mm wide. Wettsteiniola apipensis occurs only in the Salto Apipe. municipality of Tomazina. 2011 . Royal Botanic Gardens. Acknowledgements The authors thank the herbaria curators for the loan the exsicatae. up to 3mm long.. occurring also in Africa. Corrientes province. Rio das Cinzas. the first two are known only from the type locality. in the municipality of Piracicaba. Repert. M. Melhem. fl. Aona.5 cm long. 2–4 cm long.P. pinnata. The type of the species is indicated for the state of São Paulo. C. J. 17. Kew. 8–10 cm long. Capsule. This species is easily recognized in the fertile condition by large pedicels and three tepals involving the ovary. apipensis. Flora fanerogêmica do estado de São Paulo. R.. The species is the only one in the family that has a Pantropical distribution. Vol. Edusp. T. 3 mm long.. R. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III.M. c. arising from the thalloid base.P. Nov. municipality of Guaira and the collections Silva et al. Anelise Nuernberg and Mariane Beretta for photographs and CAPES for the scholarship granted to the first author.e fr. However. Wettsteiniola pinnata has the type indicated for the Paranapanema River.. occurring mainly in communities with species of Podostemum.E. Ovary c. Hatschbach & Haas 15871 (MBM). & Giulietti. In southern Brazil. APG III. 237-258.br/2010/ FB000194>.Y. G. Botanical Journal of the Linnean Society 161: 105-121. Spec. Wettsteiniola pinnata has a very restricted distribution. L. especially by arranging clusters of flowers at the thalloid base. biogeographic analyses (Kita & Kato 2004) indicate that there are possibly two different species. last divisions numerous. 39:18.5 cm long. Wettsteiniola Suess. 3–5mm diam. Fátima Zagonel and Maria Virgília for the illustrations. forming fascicles. Disponível em <http://floradobrasil. up to 0.1967. PARANÁ: Guaira..T.J. the field work in Salto Cavalcanti took place during periods of heavy rainfall. one confined to the African continent and the other to the Americas. Argentina.E. Rio das Cinzas. J. Tomazina. Unfortunately. Rodriguésia 62(4): 867-885.C. & Rutishauser.S. M. The tristichous phyllotaxy also assists in the identification. bivalvar.K. 63: 156-160. Authors of plant names. The Salto Cavalcanti is a large waterfall with high flow. only W. however this river is located on the border with the state of Paraná and new collections of the species were not done in São Paulo. state of São Paulo. tepals 3–5. Silva et al. Shepherd G. Sete Quedas do Rio Paraná. apex acute. Wettsteiniola accosii occurred in the former Salto de Piracicaba. 6 to 7 per valve. Podostemaceae. Brazil. 6. 1935 The genus is endemic to the basin of the Paraná River and composed by W.Podostemaceae in Southern Brazil specimens of Podostemum. 883 6034 and 6037 (MBM) from Salto Cavalcanti. São Paulo. & Amaral. Repert. accorsii as synonyms. filiform.

Fontana. A. A new species of Podostemum (Podostemaceae) from the states of Paraná and Santa Catarina. 1988. A. & Enricone.T. Z. R. 1955.A. New York Botanical Garden’s Virtual Herbarium.M. & Zuccarini. & Kato. E. 2008. R. Pp. C. C.T. Suzuki. New York. R. Moline. Santa Cruz do Sul 15: 7-12. 1975..T. Journal of Biogeography 31: 1605–1612. Philbrick. 18a: 1-68. Part I: The herbaria of the world. P.R. Adansonia 3: 223-262. 1990. 2004. C. Nueva espécie de Podostemaceae para Argentina Wettsteiniola apipensis. Novelo. L.H.. Pfeifer. Available in <http://sweetgum. Tavares. & Jäger-Zürn. & Novelo. 2006. 2011. Podostemaceas Argentinas.). Monograph of Podostemum (Podostemaceae). 427-476. Brazil. 2 vols.S. & Bove. B.M. Hicken. Universidade do Amazonas.T. 1822. J. Index Herbariorum: a global directory of public herbaria and associated staff. & Novelo.P. R. B. 1930.E. B. A. & Trevisan. 1-61. Fontana. Rutishauser. Manaus. O. A. A. & Novelo. C. L. Maeda. Flora brasiliensis 4: 229-275. M. A.. & Les. 1863. Philbrick. tab. Plant Biology 3: 156-163. A. & Novelo. 1997. Engler. Novelo. Caderno de Pesquisa Série Biologia. A. Xi. Bittrich. Phylogenetic relationships between disjunctly occurring groups of Tristicha trifaria (Podostemaceae). Podostemáceas Argentinas. Access on 18 Nov 2010.T. Tippery.T. M. New York Botanical Garden.H.B. Systematic Botany 29: 109-117. Lindley. C. Martius . 1262p.. H. 2. 2006. Botánica 6: 1-20. Mathew. A. A ocorrência inédita de uma Podostemaceae nas costas de uma laguna (Laguna dos Patos) no Rio Grande do Sul.P. C. R. C. Boletín de la Sociedad Argentina de Botánica. & Kato.T. C. C. 2004. Vol 3. 175-176. Brittonia 47: 210-222..44. Infrafamilial phylogeny of the aquatic angiosperm Podostemaceae inferred from the nucleotide sequences of the matK Gene. 164p. (ed. (ed. & Bove C. Sec. C.C. 126.. Cusset. L. & Irgang.H. Flora boreali Americana. In: Burkhart. J. 8th ed. Annals of the Missouri Botanical Garden 97: 425-456. Pontiroli.S.org/ih/>. Philbrick. Revista Chilena de Historia Natural 21: 148-151. Senna Gastal Jr. B. Holmgren. Tavares.S. Nova genera et species plantarum. An introduction to natural system of botany. A. 2005. 1995. tab.. 2001.T. Holmgren. Comparative morphology and molecular systematicas of Podostemum (including Crenias) – American river-weeds (Podostemaceae). figs. Endemism in neotropical Podostemaceae. Index Herbariorum. Bulletin Museum d’Historie Natural Paris.). Podostemaceae. 2004. N. Typis Caroli Crapelet.. Pp. Cronquist. C. M. Nova genera et species plantarum. Morphology and ecology of two new species of Polypleurum (Podostemaceae) from Kerala. Odnetz. 2010 [continuously updated].L. I. Nileena.Y. C. 1816. B.C. R. Bove.T. Michaux. & Davis. Bove.P..K. nybg. A. Rodriguésia 62(4): 867-885.. C.. 2008. Philbrick. Developmental morphology of foliose shoots and seedlings of Dalzellia zeylanica (Podostemaceae) with special reference to their meristems. Botanische Jahrbücher für Systematik. Gustafsson. Imaichi. 1988. India. Die Naturlichen Pflanzenfamilien. 2003. 16: 320-324. Instituto Nacional de Pesquisas da Amazônia. Philbrick. Candollea 62: 146-148. Rufhel.. Insula 35: 19-50. C. New world Podostemaceae: ecological and evolutionary enigmas. C. J. Tavares. Plant Systematic. 2003. Boletim Sociedad Argentina. Novon 11: 92-96. Columbia University Press. Philbrick. A. Vol. Kita. N. 1: 6. Tese de Doutorado. 1830..P.G.I. K. Leipzig. 2003.P. A. In: Humboldt & Bonpland. Flora Ilustrada de Entre Rios. V. 693p. Etudes sur les Podostemopsida repartition et evolution des Tristichaceae. C. D. A. & Cusset. Podostemaceae. Boletín Sociedad Argentina de Botánica 43: 269-272.. P. 1803. Kita. Tur.C.F. & Kato. Pflanzengeschichte und Pflanzengeographie. & Rutishauer. M.L. New York. Aspectos ecológicos y nuevas citas de Apinagia yguazuensis (Podostemaceae). & Barnett. 2007.T. C. Brazil. Philbrick. M... N.. 4 (10).. Richard. A família Podostemaceae em rios amazônicos e comunidades de insetos associados. In: Martius. An integrated system of classification of flowering plants. Systematic Botany Monogrphs 70: 1-72. 2011. Brasil. C. Evolution 237: 209-217. G. Podostemaceae. Botanical Journal of the Linnean Society 144: 289-302 Irgang.G.884 Mello. 1981. Philbrick. Systems caracteris plantarum . Marathrum (Podostemaceae). Systematic Botany 36: 105-118.. 1917. Two new genera of Podostemaceae from the state of Minas Gerais. Parisiis & Argentoraté. Novon 18: 94-98. Thiers. Systematics and phylogeny of neotropical Podostemaceae. 2011 . 2010. Brazil.. American Journal of Botany 98: 306-325. Tur. R. C. 73-76. Phylogeny of clusoid clade (Malpighiales): Evidence from the plastids and mitochondrial genomes.. Libraire greqcqueLaténe-Allemande 1: 246-247. J. A new species of Castelnavia (Podostemaceae) from Tocantins.R.. un nuevo gênero para el Paraguay.M. Podostemaceae de alguns rios de água preta do estado Amazonas.J.R.. Ser. A. N. Y. Philbrick. Tulasne.S. Philbrick.

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