Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
Alismatales from the upper and middle Araguaia river basin (Brazil)
SAMANTHA KOEHLER1, 3 and CLAUDIA PETEAN BOVE2
(received: October 30, 2002; accepted: March 25, 2004)
ABSTRACT – (Alismatales from the upper and middle Araguaia river basin (Brazil)). The present study deals with a survey of
the order Alismatales (except Araceae) in the upper and middle Araguaia River region located between the states of Mato
Grosso and Goiás, Brazil. Field expeditions were carried out during the rainy and dry seasons. The route covered approximately
2,000 km and 41 aquatic environments were visited. Thirteen taxa, representing the families Alismataceae (nine), Hydrocharitaceae
(three) and Najadaceae (one) were identified. Keys for the identification of families and species in field, brief diagnoses,
schematic illustrations and relevant comments were elaborated based on field observations as well as on the analysis of the
specimens collected.
Key words - Alismatales, aquatic plants, Araguaia river, Brazil, hydrophytes
RESUMO – (Alismatales da bacia do alto e médio rio Araguaia (Brasil)). Realizou-se o levantamento de espécies da ordem
Alismatales (exceto Araceae) ocorrentes na região do alto e médio rio Araguaia, entre os estados de Mato Grosso e Goiás,
Brasil. As expedições para coleta de material, ocorridas tanto na época de chuvas quanto na seca, totalizaram cerca de 2.000 km
percorridos, abrangendo 41 ambientes aquáticos. Foram identificados treze táxons pertencentes às famílias Alismataceae
(nove), Hydrocharitaceae (três) e Najadaceae (uma). Foram elaboradas chaves para identificação em campo das famílias e
espécies, descrições breves, ilustrações esquemáticas e comentários relevantes, baseados em dados levantados em campo e
através da análise do material coletado.
Palavras-chave - Alismatales, Brasil, hidrófitas, plantas aquáticas, rio Araguaia
Introduction
Brazil has the largest hydrographic net of the world,
estimated as 6,950 km2, representing the largest hydric
supply of South America (Tundisi & Barbosa 1995).
Plants encountered growing in permanent or seasonally
wet environments, whose photosynthetic parts are
constantly or at least for several months each year in
contact with water are defined by Cook (1996) as
vascular aquatic plants. The aquatic flora plays an
important role on the ecology of aquatic ecosystems. It
affects the water chemistry by the assimilation of
nutrients and noxious substances (Esteves & Camargo
1986) and provides favorable conditions to innumerous
organisms, such as fishes, aquatic arthropods, mollusks
and periphyton (Esteves 1988, Pott & Pott 1997). Aquatic
plants also constitute the base of an extensive food chain
(Esteves 1988, Niering 1985) and some species may
even be used as ornamental plants, soil fertilizers or for
1.
2.
3.
Universidade Estadual de Campinas, Instituto de Biologia,
Departamento de Botânica, Caixa Postal 6109, 13083-970
Campinas, SP, Brasil.
Universidade Federal do Rio de Janeiro, Museu Nacional,
Departamento de Botânica, Quinta da Boa Vista, 20940-040
Rio de Janeiro, RJ, Brasil. cpbove@ibpinet.com.br
Corresponding author: koehler@unicamp.br
cattle nourishment (Berg 1984, Joyce 1990). However,
the excessive growth of some species may interfere
with the functioning of hydroelectric turbines, favor the
proliferation of mosquitoes, reduce water oxygen
concentration, and prevent navigation (Esteves 1988,
Palombo & Pereira 1992).
Despite the great diversity of hydrophytes in Brazil
(Hoehne 1948), there is still a lack of studies on the
ecological and systematic aspects of Brazilian aquatic
plants (Pott et al. 1992). Besides the work of Hoehne
(1948) on aquatic plants of Brazil, there are studies,
generally taxonomically and/or geographically restricted,
developed for the Amazonian region (Albuquerque 1981)
and for the states of Goiás (Wanderley 1989), Mato
Grosso (Prado et al. 1994, Schessl 1997) Mato Grosso
do Sul (Guimarães 1999; Pott 1993, Pott & Pott 1997,
Pott et al. 1989, 1992, Pott & Cervi 1999, Pott & Pott
2000 [Pantanal]), Minas Gerais (Pedralli et al. 1993a,
b), Rio de Janeiro (Henriques et al. 1988, Bove et al.
2003), Rio Grande do Sul (Almeida-Rego 1988, Irgang
et al. 1984, Irgang & Gastal 1996, Pedralli, 1981), Santa
Catarina (Pedersen & Klein 1976) and São Paulo (Abreu
1981, Faria 1998).
The order Alismatales, with roughly 3,320 species,
consists mostly of aquatic or wetland plants and is
represented in Brazil by approximately 32 genera and
11 families (Haynes & Holm-Nielsen 1985, Judd et al.
440
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
1999, Les & Haynes 1995). A generic treatment of the
order in the Neotropics has been published by Haynes
& Holm-Nielsen (1985), besides surveys and taxonomic
treatments on the families Alismataceae (Almeida Rego
1988, Guimarães 1999, Haynes & Holm-Nielsen 1994,
Rataj 1978), Hydrocharitaceae (Cook 1985, Cook &
Urmi-König 1984a, b), Limnocharitaceae, included in
Alismataceae in this paper (Haynes & Holm-Nielsen
1992), and Najadaceae (Lowden 1986).
Despite the fact most studies of aquatic plants
published so far for the central region of Brazil are
geographically and/or taxonomically restricted, there are
still innumerous aquatic habitats in this region that are poorly
known. This assertion was corroborated by the discovery
of a new species of Hydrocharitaceae, Egeria
heterostemon S. Koehler & C.P. Bove, during the
development of this study (Koehler & Bove 2001). The
purpose of this study is to contribute to the knowledge of
the Brazilian aquatic flora presenting a survey of the order
Alismatales of the upper and middle Araguaia River region.
Material and methods
The study area is located on the Araguaia - Tocantins
depression, in the states of Mato Grosso and Goiás, Brazil.
The climate, warm and wet, is composed of two main seasons:
a dry one from June to November followed by a rainy season
from December to May (IBGE 1989). The vegetation is
characterized mainly by the Cerrado type, rather devastated,
and by riparian forests and aquatic ecosystems, hitherto well
preserved. Field expeditions were undertaken in 1997
(May-June and October) and in 1999 (November), covering
the rainy and dry seasons. Specimens were collected in 41
different aquatic environments along the roads indicated in
figure 1, covering an extension of approximately 2,000 km.
The sites visited consisted mainly of permanently and
temporally undrained floodplains.
25
55
o
52
I
o
50 o
I
o
Água
Boa
Rio Araguaia
Nova
Xavantina
Rio
das Mortes
Cocalinho
Peixe
Mozarlândia
Aruanã
Araguapaz
Britânia
Barra do Aragarças
Garças
Goiás
__ 16 o
Jussara
Iporã
Caipônia
Santa RIta
do Araguaia
Figure 1. Geographical location of the study area.
Species identifications were based on the taxonomic
treatments available and were complemented by comparisons
with herbaria material. The material was deposited in the
herbarium of the Museu Nacional do Rio de Janeiro (R) and
duplicates were sent to the to herbaria of the Universidade
de São Paulo (SPF) and to Embrapa – Pantanal (CPAP). Keys
for the identification of the families and species in the field,
brief diagnoses, schematic illustrations and relevant
ecological and morphological comments were elaborated
based on an analysis of specimens collected, as well as on
field observations.
Results and Discussion
In the treatment that follows thirteen taxa were
recognized.
Key to families of Alismatales of the upper and middle Araguaia river region
1. Submersed plants
2. Leaves sessile, 1-veined, margins serrulate
3. Leaves with sheathing base, margins with visible serrulations; flowers lacking a perianth,
inconspicuous .......................................................................................................................... Najadaceae
3. Leaves without sheathing base, margins without visible serrulations; flowers conspicuous,
with white perianth ....................................................................................................... Hydrocharitaceae
2. Leaves petiolate, multi-veined, margins entire
4. Leaves up to 10 cm long; flowers white .............................................................................. Alismataceae
4. Leaves longer than 10 cm; flowers yellow ................................................................... Hydrocharitaceae
1. Emersed plants ............................................................................................................................... Alismataceae
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
Alismataceae Vent.
Herbaceous, annual or perennial, glabrous or
pubescent, lactiferous. Stems corm-like, rhizomatous or
stoloniferous. Leaves submersed, floating or emergent,
basal or alternate, petiolate with a sheathing base;
submersed leaves linear, sessile; emergent and floating
leaves petiolate, the blade linear to lanceolate, ovate to
rhomboid, pluriveined, with or without pellucid marks,
the margins entire. Inflorescences verticillate, paniculate,
racemose, spicate or umbellate, without a subtending
spathe, with whorled or separate bracts. Flowers
441
hypogynous, perfect or imperfect, pedicellate, perianth
actinomorphic, of 6 free segments in 2 series: sepals 3,
green, persistent; petals 3, larger than sepals, white to
yellow sometimes with purple spots at the base, delicate,
deciduous; stamens 3 or numerous, free, staminodes
occasionally present; carpels 3, 6 or more, free or basally
fused, 1-loculed, ovules 1-2 or numerous of basal or
laminar placentation, styles terminal or lateral sometimes
very short or absent, stigma linear. Fruits achenes or
follicles, numerous, compressed or terete, often winged,
longitudinally costate and glandular, sometimes with a
terminal or horizontal beak. Seeds U-shaped.
Key to species of Alismataceae of the upper and middle Araguaia river region
1. Floatting leaves sagittate ........................................................................................... 5. Sagittaria guayanensis
1. Emersed leaves linear, lanceolate or ovate
2. Plants delicate, to 25 cm tall
3. Pellucid lines on leaf blades present .............................................................. 1. Echinodorus bolivianus
3. Pellucid lines on leaf blades absent .................................................................... 4. Echinodorus tenellus
2. Plants stout, taller than 40 cm
4. Ovules of basal placentation
5. Floral bracts subulate, usually longer than the pedicels ............................. 3. Echinodorus subalatus
5. Floral bracts lanceolate, usually shorter than the pedicels
6. Submersed leaves absent; flowers perfect ...................................... 2. Echinodorus longipetalus
6. Submersed leaves present; flowers, at least the lower, imperfect ......... 6. Sagittaria rhombifolia
4. Ovules of laminar placentation
7. Blades ovate to orbicular, as long as wide ............................................... 7. Hydrocleys nymphoides
7. Blades at least twice longer than wide
8. Leaves floating; carpels 3; staminodes 3 ............................................... 8. Hydrocleys parviflora
8. Leaves erect; carpels and staminodes numerous .................................... 9. Limnocharis laforestii
1. Echinodorus bolivianus (Rusby) Holm-Niels.,
Brittonia 31(2):276. 1979. Type: BOLIVIA: Reyes,
White 1540 (lectotype NY, fide Haynes & HolmNielsen 1994; isolectotypes, GH, K, NY, US).
Figure 2A.
Plants delicate, annual, glabrous, stoloniferous, to
15 cm tall. Submersed leaves present; emersed leaves
petiolate, 10 cm long, 1.0-1.5 cm wide, the blade
lanceolate with a sheathing base, pellucid markings
present as lines. Inflorescence umbelliform with only
one floral whorl or racemose with two whorls, erect,
up to 25 cm long, without vegetative proliferations.
Bracts triangular, basally united, coarse with thin
margins, 0.5 cm long, 0.1-0.2 cm wide. Flowers perfect,
2-6, 1 cm diam.; sepals appressed, 0.3 cm long; petals
clawed, 0.3-0.4 cm long; stamens 9, 0.1 cm long, carpels
15-20. Fruits not observed.
Common name: “dwarf amazon swordplant”, “erva do
pântano” (Haynes & Holm-Nielsen 1994, Pott & Pott 2000).
Specimens examined: BRAZIL: G OIÁS : Estr.
Jussara-Aragarças a 100 km de Jussara, 9-X-1997 (fl.),
C.P. Bove et al. 238. (R, SPF).
Habitat: amphibious or submersed plant, growing in
still or running water up to 50 cm deep. Rare.
Distribution: from Central America to southern South
America. In Brazil, in Distrito Federal, Goiás, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Pará,
Paraná, Rio Grande do Sul, Rio de Janeiro, Santa
Catarina and São Paulo (Haynes & Holm-Nielsen 1994).
2. Echinodorus longipetalus Micheli, Monogr. Phan.
3:60. 1881. Type: PARAGUAY: Cordillera de Peribebuy,
Patiño Cué, Balansa 570 (lectotype K, fide Rataj 1969;
isolectotypes, G, K).
Figure 2B.
442
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
Plants stout, perennial, glabrous, to 200 cm tall.
Submersed leaves absent; emersed leaves petiolate,
40-120 cm long, 10-15 cm wide, with a sheathing base
5-15 cm long, the blade lanceolate, pellucid markings
reticulate. Inflorescence racemose, erect or decumbent,
overtopping the leaves, without vegetative proliferations,
2-10 whorls with 3-15 flowers each. Bracts lanceolate
with an attenuate apex, coarse, 2 cm long. Flowers
perfect; sepals erect, covering the mature fruit, 1-1.5 cm
long, 0.5-1 cm wide; petals 2 cm long, 1.5-2 cm wide;
stamens numerous, 0.5 cm long, carpels numerous. Fruits
flattened, dorsally keeled, 0.3 cm long, the beak lateral,
horizontal.
Common name: “narrow-leaved Amazon
swordplant”, “chapéu de couro” (Haynes & HolmNielsen 1994, Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Aragarças, a 100 km de Jussara, 26-V-1997
(fl.), C.P. Bove et al. 158, 159 (R); Estrada AruanãBritânia, a 33 km de Aruanã, 14-X-1997 (fl.), C.P. Bove
et al. 295 (R); Estrada Jussara-Britânia, ca. de 40 km
do trevo, 14-XI-1999 (fl., fr.), C.P. Bove et al. 577
(R, SPF). MATO GROSSO: Estrada Água Boa-Cocalinho,
a 144 km de Água Boa, 12-X-1997 (fl., fr.), C.P. Bove
et al. 272 (R, SPF).
Habitat: emergent plant, growing in still water up
to 50 cm deep. Commonly found around the edges of
marshes, usually presenting a high local abundance.
Distribution: Suriname and from central Brazil
(states of Goiás, Mato Grosso, Mato Grosso do Sul,
Minas Gerais and São Paulo) to eastern Paraguay
(Haynes & Holm-Nielsen 1994, Pott & Pott 2000).
3. Echinodorus subalatus (Mart.) Griseb. subsp.
subalatus, Cat. pl. Cub. p.218. 1866. ≡ Alisma
subalatum Mart., Syst. veg. 7(2):1609. 1830. Type:
BRAZIL: s. loc., C.F.P. Martius 150 (lectotype M, fide
Rataj 1971; isolectotype MO).
Figure 2C-D.
Plants stout, perennial, glabrous, 40-100 cm tall.
Submersed leaves absent; emersed leaves petiolate,
23-55 cm long, 1-8 cm wide, with a sheathing base
6-15 cm long, the blade lanceolate, pellucid markings
present as lines. Inflorescence racemose or paniculate,
erect, 50-60 cm long, with or without vegetative
proliferations, 5-6 whorls with 5-10 flowers each. Bracts
subulate, coarse, with delicate margins, 0.5-2 cm long.
Flowers perfect; sepals reflexed, 0.5 cm long, 0.5 cm
wide; petals 1 cm long, 0.5-1 cm wide; stamens 20,
0.4 cm long, carpels numerous. Fruits terete, unkeeled,
glandular, 0.1-0.2 cm long, the beak terminal, erect.
Common name: “longstyled toadspoon”, “chapéu
de couro miúdo” (Haynes & Holm-Nielsen 1994, Pott
& Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Aragarças, a 3 km de Jussara, 9-X-1997 (fl.,
fr.), C.P. Bove et al. 246 (R, SPF); Estrada AruanãBritânia, a 10 km de Britânia, 14-XI-1999 (fl.), C.P. Bove
et al. 586 (R); Estrada para Peixe (GO-334), a 39 km
da GO-164, 16-XI-99 (fl.), C.P. Bove et al. 624 (R).
MATO GROSSO: Estrada Água Boa-Cocalinho, a 100 km
de Água Boa, 12-X-1997 (fl., fr.), C.P. Bove et al. 264
(R).
Observations: we found specimens with vegetative
proliferating inflorescences (C.P. Bove et al. 246), a
character not recorded in previously taxonomic
treatments of this species (Haynes & Holm-Nielsen
1994, Rataj 1978).
Habitat: emergent plant, growing in still water up
to 50 cm deep; however, it may be found in the middle
of rivers. Commonly found around the edges of marshes
and as a dominant species in polluted environments.
Distribution: from Central America to south-central
Brazil, states of Amazonas, Bahia, Ceará, Goiás,
Maranhão, Mato Grosso, Mato Grosso do Sul, Minas
Gerais, Paraná, Rio Grande do Norte, Rio de Janeiro,
Roraima, Santa Catarina and São Paulo (Haynes &
Holm-Nielsen 1994, Pott & Pott 2000).
4. Echinodorus tenellus (Mart. ex Schult. & Schult.
f.) Buchenau, Abh. Naturwiss. Vereines Bremen 2:21.
1868. ≡ Alisma tenellum Mart. ex Schult. & Schult. f.,
Syst. veg. 7(2):1600. 1830. Type: BRAZIL: Buritihaes,
Contendas, C.F.P. Martius a56 (lectotype M, fide Rataj
1975).
Figure 2E.
Plants delicate, annual, glabrous, stoloniferous, to
6 cm tall. Submersed leaves present, 5 cm long, 0.2 cm
wide, with a sheathing base 2 cm long; emersed leaves
petiolate, 1.5-4.5 cm long, 0.2 cm wide, the blade linearlanceolate with a sheathing base 1 cm long, pellucid
markings absent. Inflorescence umbelliform with only
one floral whorl, erect, up to 6 cm long, without
vegetative proliferations. Bracts triangular, basally united,
coarse with delicate margins, 0.3 cm long, 0.1-0.2 cm
wide. Flowers perfect, 2-28, 0.6-1 cm diam.; sepals
appressed, 0.3 cm long; petals clawed, 0.2-0.4 cm long,
0.1-0.3 cm wide; stamens 9, 1 cm long, carpels 15-20.
Fruits not observed.
Common name: “pigmy chain-sword”, “dwarf
Amazonian sword plant”, “erva do pântano” (Haynes
& Holm-Nielsen 1994, Pott & Pott 2000).
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Aragarças, a 100 km de Jussara, 26-V-1997
(fl.), C.P. Bove et al. 161 (R); 9-X-1997 (fl.), C.P. Bove
et al. 240 (R); Estrada Jussara-Britânia a 59 km de
Jussara, 14-XI-1999 (fl.), C.P. Bove et al. 581 (R);
Estrada Jussara-Britânia (via Jacilândia), a 59 km de
Jussara, 12-XI-1999 (fl.), C.P. Bove et al. 550 (R, SPF);
443
Estrada para Peixe (GO-334), a 21 km da GO-164,
16-XI-1999 (fl.), C.P. Bove et al. 622 (R, SPF). MATO
GROSSO: Estrada Água Boa-Cocalinho, a 58 km de Água
Boa, 11-X-1997 (fl.), C.P. Bove et al. 255 (R, SPF).
Habitat: amphibious or submersed plant, growing
in still water up to 50 cm deep or in muddy soil.
Specimens collected in dry soil were intensively
Figure 2. Species of Echinodorus from the upper and middle Araguaia basin. A. E. bolivianus, habit. B. E. longipetalus, habit.
C. E. subalatus, habit. D. E. subalatus, flower. E. E. tenellus, habit (A: Bove et al. 238, B: Bove et al. 158, C-D: Bove et al. 264,
E: Bove et al. 255).
444
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
flowering compared to flowering specimens found in
wet environments. Frequent and usually locally abundant.
Distribution: northeastern U.S.A. to southern Brazil,
states of Amazônia, Bahia, Minas Gerais, Goiás, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Rio
de Janeiro, Roraima, Santa Catarina, São Paulo (Haynes
& Holm-Nielsen 1994, Pott & Pott 2000).
Habitat: floating-leaved plant, growing in still water
50-100 cm deep. A very common species in the
Araguaia river region, usually locally abundant.
Distribution: from southern U.S.A. to Argentina and
Brazil, states of Acre, Amazonas, Ceará, Goiás,
Maranhão, Mato Grosso, Mato Grosso do Sul, Minas
Gerais, Pará, Piauí, Rondônia and Roraima (Haynes &
Holm-Nielsen 1994, Pott & Pott 2000).
5. Sagittaria guayanensis Kunth subsp. guayanensis,
Nov. gen. sp. 1:250. 1816. Type: COLOMBIA: Guainia,
Humboldt s.n. (holotype P?; fragment at MO; lectotype
TCD, Rataj 1972; isolectotypes LE, W).
Figure 3A-B.
6. Sagittaria rhombifolia Cham., Linnaea 10:219. 1835.
Type: BRAZIL: s. loc., Sellow s.n. (lectotype E, fide
Rataj 1972; isolectotype K, LE).
Figure 3C-E.
Plants perennial, glabrous or pubescent, to 50 cm
tall. Stems present as corms. Submersed leaves present,
0.5-1 cm long; emersed leaves floating, the peciole to
30 cm long, with a sheathing base to 10cm long, the
blade sagittate, 1-4 cm long, 0.5-5 cm wide, pellucid
markings absent. Inflorescence racemose, floating,
rarely erect, 1-20 cm long, with 1-5 whorls, 3 flowers
each, without vegetative proliferations. Bracts linearlanceolate, free, delicate, 0.3-0.5 cm long. Flowers
imperfect; sepals erect, 0.5-1 cm long, 0.5 cm wide;
petals clawed, 0.5-1 cm long, 0.6 cm wide; male flowers
with sterile carpels, stamens 6, 0.3 cm long; female
flowers with a ring of sterile stamens, carpels numerous.
Fruits oblanceolate, keeled, without glands, the margins
echinate, 0.2 cm long, the beak lateral, erect.
Common name: “lagartixa”, “largatissa”, “corazón
de agua” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Araguapaz-Aruanã, a 11 Km de Araguapaz, 13-X-1997
(fl., fr.), C.P. Bove et al. 281 (R, SPF); 13-X-1997
(fl., fr.), C.P. Bove et al. 282 (R); Estrada JussaraAragarças, a 100 km de Jussara, 25-V-1997 (fl.), C.P.
Bove et al. 136, fl. (R); 25-V-1997 (fl.), C.P. Bove
et al. 150 (R); 26-V-1997 (fl.), C.P. Bove et al. 187
(R); 26-V-1997 (fl.), C.P. Bove et al. 189 (R);
27-V-1997 (fl., fr.), C.P. Bove et al. 193 (R, SPF);
Estrada Jussara-Aragarças, a 97 km de Jussara,
27-V-1997 (fl.), C.P. Bove et al. 196 (R); 11-XI-1999
(fl., fr.), C.P. Bove et al. 513 (R); Estrada JussaraBritânia a 39,6 km de Jussara, 12-XI-1999 (fl., fr.),
C.P. Bove et al. 561 (R, SPF); Estrada JussaraBritânia, a 54,4 km de Jussara, 28-V-1997 (fl., fr.),
C.P. Bove et al. 200 (R, SPF); Estrada JussaraBritânia (via Jacilândia), a 78,2 km de Jussara,
12-XI-1999 (fl.), C.P. Bove et al. 536 (R). M ATO
G ROSSO : Estrada Paranatinga-Canarana km 151,
12-II-1993 (fl., fr.), C.P. Bove et al. 95 (R).
Plants perennial, glabrous, 50-100 cm tall.
Submersed leaves present, ca. 50 cm long, 0.5 cm wide;
emersed leaves erect, petiolate, 10-45 cm long, with a
sheathing base to 5-10 cm long, the blade linearlanceolate, 5-15 cm long, 0.2-2.3 cm wide, pellucid
markings absent. Inflorescence racemose, erect, rarely
floating, 20-40 cm long, with 2-10 whorls with 3 flowers
each, without vegetative proliferations. Bracts
lanceolate, united at base, delicate to coarse, 0.5-2.5 cm
long. Flowers imperfect; male flowers with spreading
pedicels, 2-4 cm long, sepals erect, 1 cm long, 0.5 cm
wide; petals clawed, 2 cm long, 1 cm wide; stamens
9-12, 0.5 cm long, sterile carpels present; female flowers
with erect pedicels, spreading and enlarged in fruit,
1.5-2 cm long, appressed in flower and fruit, 1.5-2 cm
long, 1-2 cm wide, petals clawed, 1.5-3 cm long, 0.5 cm
wide, without sterile stamens, carpels numerous. Fruits
oblanceolate, keeled, without glands, 0.1 cm long, the
beak lateral, horizontal.
Common name: “lagartixa” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Britânia, a 62 km de Jussara, córrego Píndaíba,
14-XI-1999 (fl.), C.P. Bove et al. 582 (R); Estrada
Jussara-Aragarças, a 100 km de Jussara, 25-V-1997
(fl.), C.P. Bove et al. 143 (R); 25-V-1997 (fl., fr.), C.P.
Bove et al. 144 (R); 25-V-1997 (fl.), C.P. Bove et al.
145 (R); 25-V-1997, C.P. Bove et al. 146 (R);
25-V-1997 (fl.), C.P. Bove et al. 147 (R); 25-V-1997
(fl.), C.P. Bove et al. 148 (R); 25-V-1997 (fl.), C.P.
Bove et al. 149 (R); 26-V-1997 (fl., fr.), C.P. Bove
et al. 162 (R); 26-V-1997 (fl.), C.P. Bove et al. 163
(R); 26-V-1997 (fl., fr.), C.P. Bove et al. 164 (R);
26-V-1997 (fl.), C.P. Bove et al. 165 (R); 9-X-1997
(fl.), C.P. Bove et al. 236 (R, SPF); Estrada JussaraAragarças, a 111 km de Jussara, 11-XI-1999 (fl., fr.),
C.P. Bove et al. 526 (R); Estrada Jussara-Britânia (via
Jacilândia), a 42,2 km de Jussara, 12-XI-1999 (fl.), C.P.
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
Bove et al. 560 (R); Estrada Caiapônia-Iporã, a 54 km
do trevo, 19-XI-99 (fl., fr.), C.P. Bove et al. 670 (R,
SPF). M ATO GROSSO : Paranatinga, 39 km ao sul,
9-II-1993 (fl., fr.), C.P. Bove et al. 89 (R).
Habitat: emergent plant, growing in still or running
water 50-100 cm deep. A very common species in this
region.
Distribution: from southern Costa Rica to Argentina.
In Brazil in the states of Bahia, Distrito Federal, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Pará,
Paraná, Piauí, Rio de Janeiro, Santa Catarina and São
Paulo (Haynes & Holm-Nielsen 1994, Pott & Pott 2000).
7. Hydrocleys nymphoides (Willd.) Buchenau, Index
Crit. Butom. Alism. Juncag. p. 9. 1868. ≡ Stratiotes
nymphoides Willd., Sp. pl. 4(2):821. 1806. Type:
VENEZUELA: s. loc., Humboldt & Bonpland s.n.
(holotype B-W).
Figure 4A-C.
445
Plants to 50 cm tall, stoloniferous. Floating leaves
7-20 cm long, 2-6 cm wide, with an ovate-orbicular
blade, petiolate, with a sheathing base 5cm long.
Inflorescences with 1-5 flowers, scape 5-10 cm long.
Bracts elliptic, 1-2 cm long, 0.5 cm wide. Sepals with
midvein inconspicuous and hialine margins, 3 cm long,
1 cm wide; petals reflexed, pale yellow to white with a
yellow base, 2.5 cm long, 3 cm wide; stamens 20-30 in
2 series or more, 1 cm long, staminodes numerous;
carpels 7, 1.5 cm long. Fruits not observed.
Common name: “lagartixa”, “lagartissa”, “amapolade-água”, “water poppy” (Pott & Pott 2000).
Specimens examined: BRAZIL: MATO GROSSO:
Estrada Mozarlândia-Nova Crixás km 18, 13-X-1997
(fl.), C.P. Bove et al. 276 (R, SPF); Estrada
Mozarlândia-Nova Crixás a 18,6 km do trevo, 15-XI-1999
(fl.), C.P. Bove et al. 606 (R, SPF).
Observations: flowers closed early in the morning,
opening around 10:00 am.
Figure 3. Species of Sagittaria from the upper and middle Araguaia basin. A-B. S. guayanensis subsp. guayanensis. A. Habit;
B. Male flower. C-E. S. rhombifolia. C. Habit; D-E. Leaves (A-B: Bove et al. 200, C-D: Bove et al. 148, E: Bove et al. 670).
446
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
Figure 4. Species of Alismataceae and Najadaceae from the upper and middle Araguaia basin. A-C. Hydrocleys nymphoides. A.
Habit; B. Flower; C. Calix. D-F. Hydrocleys parviflora. D. Habit; E. Fruit; E. Calix. G-H. Limnocharis laforestii. G. Habit; H. Fruit.
I-J. Najas microcarpa. I. Habit; J. Leaf. (A-C: Bove et al. 276, D-F: Bove et al. 201, G-H: Bove et al. 618, I-J: Bove et al. 260).
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
Habitat: plants with floating leaves, growing in still
water 50-100 cm deep. Sporadic.
Distribution: tropical and subtropical North America,
Central America, and South America. In Brazil in the
states of Bahia, Ceará, Mato Grosso, Mato Grosso do
Sul, Pará, Paraíba, Pernambuco, Piauí, Rio Grande do
Norte, Rio de Janeiro and Santa Catarina (Haynes &
Holm-Nielsen 1992, Pott & Pott 2000).
8. Hydrocleys parviflora Seub. in Mart. & Eichler, Fl.
bras. 3(1):117. 1847. Type: BRAZIL: s. loc., Martius
s.n. (lectotype M, fide Haynes & Holm-Nielsen 1992).
Figure 4D-F.
Plants to 100 cm tall, stoloniferous. Floating leaves
with an elliptic blade, 5 cm long, 1 cm wide, petiole up
to 100 cm long with a sheathing base 10 cm long.
Inflorescences with 2-10 flowers, scape 100 cm long.
Bracts lanceolate, 2 cm long, 1 cm wide. Sepals with
conspicuous midvein, 2 cm long, 1 cm wide; petals white,
1.5 cm long, 1 cm wide; stamens 5 in one whorl, 0.3 cm
long, staminodes 3; carpels 3, 0.4 cm long. Fruits 1.2 cm
long, beak 0.1 cm long; seeds numerous, glandular
pubescent, 1 mm long.
Common name: “lagartixa” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Britânia, a 54 km de Jussara, 28-V-1997 (fl.),
C.P. Bove et al. 201 (R, SPF).
Habitat: plants with floating leaves, growing in still
water 50-100 cm deep. Rare in the Araguaia river region.
Distribution: southern Mexico to Costa Rica, northern
Colombia, Venezuela and Bolivia to east-central Brazil, in
the states of Goiás, Mato Grosso do Sul, Minas Gerais and
Rio de Janeiro (Haynes & Holm-Nielsen 1992).
9. Limnocharis laforestii Griseb., Bonplandia 6(1):11.
1858. Type: PANAMA: s. loc. Duchassaing et al. s.n.
(lectotype GOET, fide Novelo 1987; isolectotype
GOET).
Figure 4G-H.
Plants to 50 cm tall, rhizomatous. Leaves erect,
petiolate, with an elliptic to ovate blade, 15-25 cm long,
1-2 cm wide with a sheathing base 5 cm long.
Inflorescences with up to 5 flowers. Bracts elliptic-linear,
447
0.5-1.5 cm long, 0.3-1 cm wide; pedicels 1 cm long,
1 cm wide. Sepals ovate, 1cm long, 0.7 cm wide; petals
yellow, ovate to orbicular, 1.5 cm long, 0.8 cm wide;
stamens 15-20, 0.6 cm long, staminodes numerous;
carpels 20-30, 0.4 cm long. Fruits 0.1 cm long; seeds
numerous, 0.1 cm long.
Common name: “camalote” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Britânia-Aruanã, a 5 km de Jussara, 27-V-1997 (fl., fr.),
C.P. Bove et al. 205 (R). MATO GROSSO: Estrada Água
Boa-Cocalinho, a 144 km da GO-164, 12-X-1997 (fl., fr.),
C.P. Bove et al. 268 (R, SPF).
Observations: the specimens examined presented
the peduncles shorter than the petioles and the pedicels
not conspicuously winged. The staminodes were
numerous, like in Limnocharis flava (L.) Buch. The
number of staminodes, however, is not considered a
diagnostic character according to the last taxonomic
treatment available for the genus (Haynes & HolmNielsen 1992).
Habitat: emergent plant, growing in still water
50-100 cm deep. Sporadic.
Distribuição: Mexico to Bolivia, Argentina and
Brazil, in the states of Bahia, Goiás, Maranhão, Mato
Grosso, Mato Grosso do Sul, Paraíba, and Piauí (Haynes
& Holm-Nielsen 1992, Pott & Pott 2000).
Hydrocharitaceae Juss.
Herbaceous, annual or perennial, glabrous, not
lactiferous. Stems erect, rhizomatous or stoloniferous,
elongated or contracted. Leaves submersed, floating or
partially emersed, basal, alternate or whorled, sessile or
petiolate, margins entire or serrulate, the blade linear to
eliptic-lanceolate, 1- or pluriveined, without pellucid
markings. Inflorescences complex with 1-many flowers
or flowers solitary, with subtending spathe, composed
of 1-2 free or united bracts. Flowers epigynous, perfect
or imperfect, perianth actinomorphic, of 6 free segments
in 2 series: sepals 3, green, persistent; petals 3, larger
than sepals, white or yellow, delicate, deciduous; stamens
3, 6 or 9, free; carpels 3, fused, ovary 1-loculed,
prolongated in a free hypanthium, numerous ovules of
parietal or laminar placentation, the styles flattened. Fruit
a capsule with numerous fusiform seeds.
Key to the species of Hydrocharitaceae of the upper and middle Araguaia river region
1. Stems elongated; leaves sessile; flowers white
2. Flowers perfect; stamens 3 ................................................................................ 10. Apalanthe granatensis
2. Flowers imperfect; stamens 9 ................................................................................ 11. Egeria heterostemon
1. Stems contracted; leaves petiolate; flowers yellow ....................................................... 12. Ottelia brasiliensis
448
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
10. Apalanthe granatensis (Bonpl.) Planch., Ann. Mag.
Nat. Hist., Sér. 2 (1):87. 1848. ≡ Elodea granatensis
Bonpl. in Humbolt & Bonpland, Pl. aequinoct. 2(16):150,
pl.128. 1813. Type: COLOMBIA: “in aquis Novas
Granatea, prope urbem Guaduas”, “entre Honda et
Cune”, elev. 1,150 m, Humboldt & Bonpland s.n.
(holotype, plate in Humbolt & Bonpland, Pl. aequinoct.
2 (16), pl. 128. 1813).
Figure 5A-B.
Plants submersed or partially emersed. Stems
elongated, terete, irregularly branched. Prophylls paired
at the base of branches, deltate; leaves alternate or
whorled, sessile, 3-7 in each node, margins serrulate,
1-veined, the blade linear and gradually attenuated at
the apex, 1-1.5 cm long. Subtending spathe consisting
of two bracts, sessile, axillary, urn-shaped, 0.5 cm long,
0.1 cm wide. Flowers solitary, perfect, sessile, emersed;
sepals ovate, reflexed, 0.2 cm long, 0.1cm wide; petals
obovate, white, delicate, 0.3 cm long, 0.2 mm wide;
stamens 3, 0.2 cm long; styles 3, flattened, 2- or
3-divided, with nectaries at the base, 0.2 cm long; ovary
0.5 cm long, attenuate above in an elongated free
hypanthium to 5 cm long. Capsule ellipsoidal, irregularly
dehiscent, 0.5 cm long; seeds up to 30, ca. 1 mm long,
covered with hairs.
Common name: “lodinho branco” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Aruanã-Goiás 8,8 km de Aruanã, 29-V-1997 (fl.), C.P.
Bove et al. 224 (R, RB); Estrada GO-334, a 47,8 km da
GO-164, 16-XI-1999 (fl.), C.P. Bove et al. 632 (R);
Brasilândia, Tupiratins, Rio Feio, 20-III-1976 (fl.),
G. Hatschbach & R. Kummrov 38497 (MBM). MATO
GROSSO. Estrada Água Boa-Cocalinho, a 148 km de Água
Boa, 12-X-1997 (fl., fr.), C.P. Bove et al. 273b (R, RB);
Estrada Água Boa-Rio das Mortes, 20-II-1993, C.P. Bove
et al. 114 (R, CPAP); Córrego do Rosário, Forquilha do
Rio, III-1918 (fr.), J.G. Kuhlmann 1620 (R).
Habitat: submersed plant, growing in still or running
water up to 50 cm deep. Sporadic.
Distribution: tropical South America, in Brazil in the
states of Amazonas, Bahia, Goiás, Mato Grosso, Mato
Grosso do Sul, Maranhão, Minas Gerais, Pará,
Pernambuco, Piauí, Rio de Janeiro, Roraima, São Paulo
(Cook 1985).
11. Egeria heterostemon S. Koehler & C.P. Bove, Novon
11:63. 2001. Type: BRAZIL: GOIÁS, Estrada AruanãGoiânia, 19 km from Aruanã, C.P. Bove et al. 225
(holotype RB; isotypes CPAP, HUEFS, MBM, MO, R).
Figure 5C-I.
Plants submersed or partially emersed. Stems
elongated, terete, irregularly branched. Prophylls paired,
deltate; leaves opposite or whorled, sometimes alternate
near the base, sessile, 4-10 in each node, frequently
recurved, linear, gradually attenuated to an acute apex,
0.6-0.8 cm long, margins and apex bearing unicellular,
thick-walled prickle-hairs. Intravaginal squamules
usually 2 in each leaf axil, ovoid to orbicular, 0.1-0.2 mm
long. Spathe sessile, axillary, ovoid, 1.0 cm long, with 2
apical teeth. Male flowers: 2 in each spathe, rarely more,
pedicels up to 4.0 cm long, emersed; sepals ovate and
boat-shaped, 0.2 cm long, 0.1 cm wide; petals
suborbicular, white, delicate, 0.2-0.4 cm long, 0.2-0.3 mm
wide; stamens 9, in alternate whorls of 3; outer and
median whorls 0.1 cm long; filaments glabrous; stamens
in the outer whorl usually smaller than those of median
whorl, although similar in form. Inner whorl: 0.15 cm
long, filaments strongly pilose distally; nectary 1, central,
3-lobed, 0.2 mm wide. Female flower: 1 in each spathe,
sessile, emersed; sepals ovate and boat-shaped, 0.2 cm
long, 0.1 cm wide; petals suborbicular, white, delicate,
0.3 cm long, 0.2 mm wide; staminodes 3, strongly
papillose distally; 0.15 cm long, styles 3, two-thirds
divided along length; ovary up to 0.3 cm long, ovoid,
attenuate above in an elongated free hypanthium to
0.35 cm long. 30-40 ovules with parietal placentation.
Fruit unknown.
Common name: unknown
Specimens examined: BRAZIL: GOIÁS: Estrada
Aruanã-Goiás, 19 km de Aruanã, 29-V-1997 (fl.), C.P.
Bove et al. 225 (CPAP, HUEFS, MBM, MO, R, RB);
MATO GROSSO: Estrada Água Boa-Cocalinho, a 148 km
de Água Boa, 12-X-1997 (fl.), C.P. Bove et al. 273a
(R, RB).
Habitat: submersed plant, growing in still or running
water up to 50 cm deep along stream margins and in
swampy environments. Rare.
Distribution: Median Araguaia river in the states of
Mato Grosso and Goiás (Koehler & Bove 2001).
12. Ottelia brasiliensis (Planch.) Walp., Ann. Bot. Syst.
3(3):510. 1852. ≡ Damasonium brasiliensis Planch.,
Ann. Sci. Nat. Bot. (Paris) 11:81, t.14. 1849. Type:
BRAZIL: MINAS GERAIS, Gardner 5191 (holotype K).
Figure 5J-K.
Plants submersed. Stems contracted, rhizomatous.
Juvenile leaves linear, sessile, ephemerous; adult leaves
petiolate, up to 50 cm, with a sheathing base, blade
membranaceous, elliptic-lanceolate, margins entire or
undulate, 10-40 cm long, 3-5 cm wide. Subtending spathe
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
449
Figure 5. Species of Hydrocharitaceae from the upper and middle Araguaia basin. A-B. Apalanthe granatensis. A. Habit;
B. Leaf. C-I. Egeria heterostemon. C. Habit; D. Male flower; E. Female flower; F. Stamen of outer whorl; G. Stamen of inner
whorl; H. Style; I. Staminode. J-K. Ottelia brasiliensis. J. Habit. K. Flower. (A-B: Bove et al. 114, C-I: Bove et al. 225,
reprodution of original publication in Novon, J-K: Bove et al. 194).
450
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
consisting of two bracts, emersed, membranaceous,
urn-shaped, 5 cm long, 1 cm wide. Flowers solitary,
perfect, pedicels 2 cm long; sepals obovate-lanceolate,
with delicate margins, 1-2 cm long, 0.5 cm wide; petals
obovate-orbicular, yellow, delicate, margins undulate,
2 cm long, 2 cm wide; stamens 6, 1cm long; styles 3,
flattened, linear, 2- or 3-divided, 3 cm long, with nectaries
at the base, ca. 0.2 cm long; ovary 0.5 cm long, anthers
elongated with a mucronate connective. Fruits not
observed.
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Aragarças, a 100 km de Jussara, 27-V-1997 (fl.),
C.P. Bove et al. 191 (R, SPF); 9-X-1997 (fl., fr.), C.P.
Bove et al. 239 (R, SPF); Estrada Jussara-Aragarças,
a 97 km de Jussara, 27-V-1997 (fl.), C.P. Bove et al.
194 (R); 11-XI-1999 (fl.), C.P. Bove et al. 503 (R, SPF);
Estrada Jussara-Britânia, via Jacilândia, a 42 km de
Jussara, 12-XI-1999 (fl.), C.P. Bove et al. 556 (R);
Estrada Jussara-Britânia, via Jacilândia, a 59 km de
Jussara, 12-XI-1999 (fl.), C.P. Bove et al. 551 (R).
Habitat: submersed plant growing in running water
50-100 cm deep. Sporadic.
Distribution: southeastern Brazil, in the states of
Minas Gerais, Mato Grosso, Mato Grosso do Sul, and
Paraná.
Najadaceae Juss.
Herbaceous, submersed, glabrous, not lactiferous.
Stems elongated with short to long internodes, irregularly
branched, smooth to spinulose, easily fragmented.
Leaves alternate to subopposite or apparently whorled,
sessile, sheathed, blade linear to lanceolate, 1-veined,
flattened to recurved, margins with inconspicuous to
conspicuous, unicellular to multicellular serrulate
projections. Flowers hypogynous, imperfect,
inconspicuous, solitary or not, perianth absent. Male
flowers shortly pedunculate, subtended by an involucral
bract, bearing 1 stamen with a sessile anther; female
flowers sessile with involucral bract rarely present
1-carpelate, 1-loculed, 1 ovule, stigma 3. Fruit a nutlet.
Seeds elliptic, ovate or fusiform, ridged, testa pitted with
many aureolae of distinct shapes.
13. Najas microcarpa K. Schum. in Mart., Eichler &
Urban, Fl. bras. 3(3):727. 1894. Type: PARAGUAY:
Rio Paraguay, Weddell 3289 (lectotype P).
Figure 4I-J.
Stems elongated with short internodes, smooth,
leaves concentrated in the terminal region. Leaves
subopposite or apparently whorled, recurved with
conspicuous, multicellular serrulate margins, 0.1-0.2 cm
long, and ciliated sheaths bearing microscopic
projections. Seeds fusiform to obovate, 0.1-0.2 cm long,
areolate with square to rectangular ornamentations.
Common name: “lodo”, “najad” (Pott & Pott 2000).
Specimens examined: BRAZIL: GOIÁS: Estrada
Jussara-Aragarças, a 3 km de Jussara, 11-XI-1999
(fl., fr.), C.P. Bove et al. 496 (R, SPF). MATO GROSSO:
Estrada Água Boa-Cocalinho, a 59 km de Água-Boa,
11-X-1997 (fl., fr.), C.P. Bove et al. 260 (R, SPF).
Habitat: free or attached on the substrate,
submersed, growing in a small brackish pond as a
dominant species in association with Cabomba furcata
Schultes & Schultes f. Sometimes also captured
between rocks of rivers. Sporadic.
Distribution: South America in French Guiana,
Venezuela, Colombia, Paraguay and Brazil, in the states
of Amazonas, Maranhão, Mato Grosso and Mato Grosso
do Sul (Lowden 1986, Pott & Pott 2000).
Albeit its biological importance, the diversity of
tropical aquatic ecosystems remains poorly known
(Crow 1993). The elaboration of checklists followed by
taxonomic studies of aquatic plants constitute a basic
need for Brazilian aquatic ecosystems today, especially
in regions that are poorly studied, such as the northern
and central regions. Unfortunately Brazilian aquatic
ecosystems have been continuously destroyed or altered
by the deviation of natural water courses, by local direct
and indirect pollution of the water as well as by the
destruction of the Cerrado vegetation and Riparian
forests, as observed during the development of this study.
Acknowledgments – We are grateful to Wilson Costa for
suggesting the exploration of the interesting and still poorly
known aquatic flora of the Araguaia basin; to André Gil,
Claudio Moreira, Jane Morrey-Jones and Renato Lisbôa for
their help in the field and to Eduardo Hicköfel for the
illustrations; to the journal Novon (Missouri Botanical Garden
Press) for permission to reproduce the illustrations of Egeria
heterostemon; and to the curators of the herbaria R, BHCB,
CPAP, HUEFS and MBM for sending material for comparative
studies. This manuiscript was greatly improved by the
comments of Fabio de Barros, Renato Mello-Silva, Robert
Haynes, Volker Bittrich, Wilson Costa and an anonymous
reviewer. Material was collected with authorization
02001.004264/98-16-AC from Ibama.
References
ABREU, L.C. 1981. Flora fanerogâmica da reserva do Parque
Estadual das Fontes do Ipiranga (São Paulo, Brasil).
102-Alismataceae. Hoehnea 9:119-120.
Revista Brasil. Bot., V.27, n.3, p.439-452, jul.-set. 2004
ALBUQUERQUE, B.W.P. 1981. Plantas forrageiras da
Amazônia. I. Aquáticas flutuantes e livres. Acta
Amazonica 11:457-471.
ALMEIDA-REGO, S.C. 1988. Alismataceae Ventenant do Rio
Grande do Sul. Dissertação de mestrado, Universidade
Federal do Rio Grande do Sul, Porto Alegre.
BERG, M.E.V.D. 1984. Formas atuais de aproveitamento das
espécies nativas e exóticas do Pantanal mato-grossense.
In Anais do 1o Simpósio de Recursos Naturais e Sócioeconômicos do Pantanal. Embrapa-DDT-CPAP,
Corumbá, p.131-136.
BOVE, C.P., GIL, A.S.B., MOREIRA, C.B., ANJOS, R.F.B. 2003.
Hidrófitas fanerogâmicas de ecossistemas aquáticos
temporários da planície costeira do estado do Rio de
Janeiro, Brasil. Acta Botanica Brasilica 17:119-135.
COOK, C.D.K. 1985. A revision of the genus Apalanthe
(Hydrocharitaceae). Aquatic Botany 21:157-164.
COOK, C.D.K. 1996. Water plants of the world. SPB Academic
Publishing, Amsterdam.
COOK, C.D.K. & URMI-KÖNIG, K. 1984a. A revision of the
genus Egeria (Hydrocharitaceae). Aquatic Botany
19:73-96.
COOK, C.D.K. & URMI-KÖNIG, K. 1984b. A revision of the
genus Ottelia (Hydrocharitaceae). 2. The species of
Eurasia, Australasia and America. Aquatic Botany
20:131-177.
CROW, G.E. 1993. Species diversity in aquatic angiosperms:
latitudinal patterns. Aquatic Botany 44:229-258.
ESTEVES, F.A. 1988. Fundamentos de Limnologia. Editora
Interciência, Finep, Rio de Janeiro.
ESTEVES, F.A. & CAMARGO, A.F.M. 1986. Sobre o papel
das macrófitas aquáticas na estocagem e ciclagem dos
nutrientes. Acta Limnologica Brasiliensia 1:273-298.
FARIA, A.D. 1998. O gênero Eleocharis R. Br. (Cyperaceae)
no Estado de São Paulo. Dissertação de mestrado,
Universidade Estadual de Campinas, Campinas.
GUIMARÃES, E.S. 1999. Alismataceae da região sul do
Pantanal Mato-grossense. Dissertação de mestrado,
Universidade de São Paulo, São Paulo.
HAYNES, R.R. & HOLM-NIELSEN, L.B. 1985. A generic
treatment of Alismatidae in the neotropics with special
reference to Brazil. Acta Amazonica 15 (supl. 1-2):153-193.
HAYNES, R.R. & HOLM-NIELSEN, L.B. 1992. The
Limnocharitaceae. Flora Neotropica Monographs
56:1-32.
HAYNES, R.R. & HOLM-NIELSEN, L.B. 1994. The
Alismataceae. Flora Neotropica Monographs 64:1-112.
HENRIQUES, R.P.B., ARAÚJO, D.S.D., ESTEVES. F.A. &
FRANCO, A.C. 1988. Análise preliminar das comunidades
de macrófitas aquáticas da lagoa de Cabiúnas, Rio de
Janeiro, Brasil. Acta Limnologica Brasiliensia 3:783-802.
HOEHNE, F.C. 1948. Plantas Aquáticas. Secretaria da
Agricultura do Estado de São Paulo, São Paulo.
IBGE. 1989. Geografia do Brasil - região Centro-Oeste. v.1.
Fundação Instituto Brasileiro de Geografia e Estatística,
Rio de Janeiro.
451
IRGANG, B.E. & GASTAL, C.V.S. 1996. Macrófitas Aquáticas
da Planície Costeira do RS. Universidade Federal do Rio
Grande do Sul, Porto Alegre.
IRGANG, B.E., PEDRALLI, G. & WAECHTER, J.L. 1984.
Macrófitos aquáticos da Estação Ecológica do Taim,
Rio Grande do Sul, Brasil. Rossléria 6:395-404.
JOYCE, J.C. 1990. Pratical uses of aquatic weeds. In Aquatic
weeds: the ecology and management of nuisance aquatic
vegetation (A.H. Pieterse & K.J. Murphy, eds.). Oxford
University Press, Oxford, p.274-291.
JUDD, W.S., CAMPBELL, C.S., KELLOGG, E.A. & STEVENS,
P.F. 1999. Plant systematics. A phylogenetic approach.
Sinauer Associates, Sunderland.
KOEHLER, S. & BOVE, C.P. 2001. Hydrocharitaceae from
Central Brazil: a new species of Egeria and a note on
Apalanthe granatensis. Novon 11:63-66.
LES, D.H. & HAYNES, R.R. 1995. Systematics of the subclass
Alismatidae: a synthesis of approaches. In
Monocotyledons: systematics and evolution (P.J.
Rudall, P.J. Cribb, D.F. Cutter & C.J. Humphries, eds.).
Royal Botanical Gardens, Kew, p.353-377.
LOWDEN, R.M. 1986. Taxonomy of the genus Najas L.
(Najadaceae) in the Neotropics. Aquatic Botany
24:147-184.
NIERING, W.A. 1985. Wetlands. The Audubon Society, New
York.
NOVELO, A. 1987. Lectotipificación de Limnocharis laforestii
Duchass. ex Grieseb., (Limnocharitaceae). Boletín de la
Sociedad Botánica de México 47:92-93.
PALOMBO, C.R. & PEREIRA, M.D.B. 1992. Monitoramento
de plantas aquáticas por satélite. Ambiente 6:49-53.
PEDERSEN, T.M. & KLEIN, R.M. 1976. Limnocaritáceas . In
Flora Ilustrada Catarinense (R. Reitz, ed.). Herbario
Barbosa Rodrigues, Itajaí, fasc. LIMN, p.1-9.
PEDRALLI, G. 1981. A família Mayacaceae Kunth no Rio
Grande do Sul, Brasil. Iheringia, série Botânica 28:47-54.
PEDRALLI, G., STEHMANN, J.R., TEIXEIRA, M.C.,
OLIVEIRA, V.L. & MEYER, S.T. 1993a. Levantamento da
vegetação aquática (macrófitos) na área da EPDA-Peti,
Santa Bárbara, MG. Iheringia, série Botânica 43:15-28.
PEDRALLI, G., MEYER, S.T., TEIXEIRA, M.C., OLIVEIRA,
V.L. & STEHMANN, J.R. 1993b. Levantamento dos
macrófitos aquáticos e da mata ciliar do Reservatório de
Volta Grande, Minas Gerais, Brasil. Iheringia, série
Botânica 43:29-40.
POTT, V.J. 1993. A família Lemnaceae S. F. Gray no Pantanal
(Mato Grosso e Mato Grosso do Sul), Brasil. Dissertação
de mestrado, Universidade Federal do Paraná, Curitiba.
POTT, V.J. & CERVI, A.C. 1999. A família Lemnaceae no
Pantanal (Mato Grosso e Mato Grosso do Sul) Brasil.
Revista Brasileira de Botânica 22:153-174.
POTT, V.J. & POTT, A. 1997. Checklist das macrófitas aquáticas
do Pantanal, Brasil. Acta Botanica Brasilica 11:215-227.
POTT, V.J. & POTT, A. 2000. Plantas Aquáticas do Pantanal.
EMBRAPA, Brasília.
452
S. Koehler & C.P. Bove: Alismatales from the Araguaia river, Brazil
POTT, V.J., BUENO, N.C., PEREIRA, R.A.C., SALIS, S.M. &
VIEIRA, N.L. 1989. Distribuição de macrófitas aquáticas
numa lagoa na Fazenda Nhumirim, Nhecolândia, Pantanal,
MS. Acta Botanica Brasilica, suplemento 3:135-168.
POTT, V.J., BUENO, N.C. & SILVA, M.P. 1992. Levantamento
florístico e fitossociológico de macrófitas aquáticas em
lagoas da Fazenda Leque, Pantanal, MS. In Anais do
VII Congresso da Sociedade Botânica de São Paulo.
Sociedade Botânica de São Paulo, Campinas, p.91-99.
PRADO, A.L., HECKMAN, C.W. & MARTINS, R.F. 1994. The
seasonal sucession of biotic communities in wetlands
of tropical wet-and-dry climate zone. II. The Aquatic
Macrophyte Vegetation in the Pantanal of Mato Grosso,
Brasil. 1994. Internationale Revue der Gesamten
Hydrobiologie 79:569-589.
RATAJ, K. 1969. Echinodorus longipetalus Mich. and other
species with reticulate markings in the blades from
Central and South America. Folia Geobotanica &
Phytotaxonomica 4:435-442.
RATAJ, K. 1971. The taxonomy of Echinodorus palaefolius
(Nees et Mart.) MacBr. (Alismataceae) and related
species from Mexico, Central and South America. Preslia
43:10-61.
RATAJ, K. 1972. Revision of the genus Sagittaria. Part II.
The species of West Indies, Central and South America.
Annotationes Zoologicae et Botanicae 78:1-61.
RATAJ, K. 1975. Revizion [sic] of the genus Echinodorus
Rich. Akvárium-Terarium:1-155.
RATAJ, K. 1978. Alismataceae of Brazil. Acta Amazonica 8
(supl. 1):1-154.
SCHESSL, M. 1997. Flora und Vegetation des nördlichen
Pantanal von Mato Grosso, Brazilien. Floristiche
Zusammensetzung, Pflanzengesellschaften und
Vegetationsdynamic saisonal und permanent
überfluteter Standorte eines tropischen
Sedimentationbeckens. Martina Galunder-Verlag,
Wiel.
TUNDISI, J.G. & BARBOSA, F.A.R. 1995. Conservation of
aquatic ecossystems: present status and perspectives.
In Limnology in Brazil (J.G. Tundisi, C.E.M. Bicudo &
T. Matsumura, eds.). ABC/SBL, Rio de Janeiro,
p.365-376.
WANDERLEY, M.G.L. 1989. Xyridaceae. In Flora do Estado
de Goiás. Coleção Rizzo (J.A. Rizzo, coord.).
Universidade Federal de Goiás, Goiânia, v.11, p.1-81.