DESCRIÇÃO DE ESPÉCIES INVASORAS DA FAMÍLIA 1

CARACTERIZAÇÃO MORFOLÓGICA DE SEMENTES DE ESPÉCIES INVASORAS DA FAMÍLIA CONVOLVULACEAE Juss.1

DORIS GROTH2

ABSTRACT - The deficiency of specialized bibliography had caused great difficulties and changes in the identification of the , in the procedures of a Seed Testing Laboratory. Therefore, through the morphological characteristics of Convolvulaceae weed seed species, which occur in Brazil and in other tropical and temperate regions, it was elaborated a taxonomical key and drawings in order to help the seed identification of 36 species. It was also presented the main synonyms and some considerations about the species, as identification problems and others. The taxonomical key were based mainly on the colour and pilosity of the seed coat (surface) and the hilum area, the size and the outline of the hilum, which are very stable characteristics. It was used also the size (length, wideness and thickness) of the seeds. The shape of the Convolvulaceae seeds, vary with the amount of seeds developed in the fruit and is not a consistent characteristic that can be used in identification. Index terms: seed identification, weed seeds, Convolvulaceae, Convolvulus, Dichondra, Jaquemontia, , Merremia.

MORPHOLOGICAL CHARACTERIZATION OF CONVOLVULACEAE Juss. WEED SEED SPECIES

RESUMO - A deficiência de bibliografia especializada tem causado grandes dificuldades e incorreções na identificação das espécies. Baseando-se nas características morfológicas das sementes de espécies invasoras da família Convolvulaceae, que ocorrem no Brasil e em outras regiões de clima tropical e temperado, foi elaborada uma chave dicotômica e desenhos para auxiliar na identificação de sementes de 36 espécies. São apresentados,também, os principais sinônimos e algumas considerações sobre a espécie, como problemas na identificação e outros. Na elaboração da chave dicotômica foram utilizadas a coloração e a pilosidade da superfície do tegumento e da área hilar, o contorno e o tamanho do hilo, que são características muito estáveis. Foram utilizadas, também, as dimensões das sementes. A forma das sementes das Convolvulaceae, varia muito em função da quantidade de sementes que se desenvolvem no fruto e não é uma característica muito consistente. Termos para indexação: identificação de sementes, espécies invasoras, Convolvulaceae, Convolvulus, Dichondra, Jaquemontia, Ipomoea, Merremia.

INTRODUCTION directly or as allergens; they obstruct lakes, irrigation ditches and drainage-ways; they reduce support capacity of natural species are considered weeds when they interfere or cultivated pastures; they reduce the availability of fish and with man’s activities or his welfare. Such grow where wildlife; they make roadsides utility rights-of-way and they are not wanted. They reduce yield and quality of crop landscape plantings unsightly; some of them are toxic and and forage species; they poison livestock and man either can compromise animal live (United States, 1970; Koehn, 1977). Another point is the development of the seed trade,

1 Aceito para publicação em 22.09.2001; trabalho apresentado no 25th Congress of the International Seed Testing Association., em Pretória, África 2 Engª Agrª, Drª, Profª Titular da FEAGRI/UNICAMP; Av. Papa Pio XII, do Sul, de 15 a 24 abril 1998. 99 apt.61, 13066-710, Campinas-SP; e-mail: [email protected]

Revista Brasileira de Sementes, vol. 23, nº 2, p.1-13, 2001 2 D. GROTH survival and dissemination with crop seeds. The spread of MATERIAL AND METHODS weed seeds by the combine is another problem, because it The seeds used in this study were obtained at the field disseminates most of the smaller and lighter seeds ripe at the and from Seed Testing Laboratory. The description were done time of harvest (Harper, 1960). No plant is absolutely a weed. On the other hand, plants almost universally desired may be utilizing methods described by Gunn (1969), Groth (1980, occasionally regarded as weed. 1984a), Groth et al. (1979, 1983) and Koehn (1977), that consider the shape, the outline, the size (length, wideness and The seed is one of the distinctive features of the thickness) of the seeds, the colour and pilosity of the seed spermatophytes which sets them apart from the so called lower coat (surface) and hilum area, the size and the outline of the plants. Seed identification is a necessary part of seed testing, hilum. At minimum of 10 fruits and 20 seeds were studied. crop improvement, wildlife management, archaeology, paleobotany and . The seed characteristics are The size rank was mentioned, but in exceptional conditions remarkably stable, under varying environmental conditions; can appear seeds very shorter or longer and this size was indicated in parenthesis. The drawings, made with the stereo- therefore, they provide reliable criteria for positive microscope ”Zeiss” and the light camera, are accompained identification of unknown seeds. with a milimeter scale, that indicate the increase utilized. The Convolvulaceae is a tropical family and seeds of several genus of this family frequently occur in lots of commercial seeds, such as clover, line, lespedeza, lucern, RESULTS AND DISCUSSION soybean, maize, wheat, common beans, rice and sorghum. On the other hand, weeds of most genus, that occur in crop Seeds of the Convolvulaceae family were described by plants have clambing habit that produce some problems on Groth (1980, 1984a - b; 1991, 1997a - b, 1998), Groth et al. cultural practices, harvest and industrial processing, leading (1979), Groth & Boaretto (1985), Kissmann & Groth (1992, to decreased seed yield and product quality. 1999) and Koehn (1977). The Convolvulaceae seeds may be There are many different types of identification keys and recogniz by their wedge shape (in cross section) and basal various methods, which can be used in developing them. hilum. Occasionally, the seeds are flattened or spheroid. There Larger the number of contrasting features that exist between are three hila types: the ipomoea-type hilum is large, nearly the material being treated the easier it is to develop and use circular in outline, usually emarginate at the base and usually the key. surrounded by a flat space and a hilar ridge; the convolvulus- The objective of these work was based on the type hilum is small, wider than long and is not surrounded by morphological characteristics of some Convolvulaceae weed a ridge; the cuscuta-type hilum is a short slit or a whitish line seed species, which occur in Brazil and in other tropical and or a raised point. The embryo is axile linear (anular or temperate regions, to elaborate a taxonomical key and spiralled) and without cotyledons in Cuscuta; axile folded drawings in order to help the correct seed identification of 36 and with obovate cotyledons in Convolvulus, or elliptic species and to present the main synonyms and some cotyledons in Dichondra, or subkidney-shaped to orbicular considerations about the species, as identification problems cotyledons in Jaquemontia or with bilobed cotyledons in and others. Ipomoea and Merremia.

Key to seed identification

1. Hilum a short slit or whitish line or a raised point; seed less than 1,25mm in diameter or 0,5-1,25mm in diameter; seed more than 1,25mm in diameter or 1,25mm long, 1,25-2,0mm wide, 1,8-2,0mm thick ...... Cuscuta spp. (Figure 1) 1. Hilum without these characteristics. 2. Convolvulus-type hilum. 3. Seed coat roughened. 4. Seed more than 3,0mm long and seed coat with numerous blunt tubercles or short thick wavy lines. 5. Seed coat dull, ochre to light greyish-brown; seed 3,0-4,0mm long, 2,0-3,0mm wide, more rounded than sectoroid-shaped ...... Convolvulus arvensis L. (Figure 1)

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Cuscuta indecora Cholsy

Convolvulus arvensis L. Convolvulus crenatifolius Ruiz et Pav.

Ipomoea amnicola Morong Dichondra microcalyx Hall.f.Fabris.

Ipomoea alba L.

Ipomoea asarifolia Jacq ssp. (Desr.) Roem.& Schult. fistulosa (Mart.ex Choisy) D.Austin

Ipomoea cairica (L.) Sweet.

Ipomoea coccinea L. Ipomoea ramosissima (Poir.) Choisy

Ipomoea indica (Burm.f.) Merr. FIG. 1. Ventral view of Convolvulaceae weed seeds.

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5. Seed coat dull, black at the maturity; seed 3,0-3,8mm long, 2,5-3,2mm wide, broadly-ellipsoid to obovoid- sectoroid-shaped ...... Convolvulus crenatifolius Ruiz et Pav. (Figure 1) 4. Seed less than 3,0mm long; seed coat without tubercles. 6. Margins narrow winged; seed coat with numerous blister-like protuberances or short irregularly wavy lines. 7. Wing and seed coat yellowish to yellowish-brown; seed about 2,0mm long, 1,4-1,7mm wide, 1,2-1,3mm thick ...... Jacquemontia densiflora Hall.f. (Figure 3) 7. Wing yellowish and seed coat yellowish to brown; seed 2,5-3,2mm long, 1,8-2,3mm wide, 1,5-1,7mm thick ...... Jacquemontia velutina Choisy (Figure 3) 6. Margins not winged; seed coat only with numerous blister-like protuberances; seed 2,3-2,7(-3,0)mm long, 1,8-2,2mm wide, 1,5-1,6mm thick ...... Jacquemontia tamnifolia (L.) Griseb. (Figure 3) 3. Seed not roughned, subglobose or ovoid-shaped, 1,5-2,2mm in diameter or long and wide, about 1,5mm thick ...... Dichondra microcalyx (Hall.f.) Fabris (Figure 1) 2. Ipomoea-type hilum. 8. Hilum glabrous and surrounded by a semihalo above. 9. Semihalo glabrous. 10. Semihalo black (more visible in the dark-grey coloured seeds); hilum brownish-black; seed 7,0-9,0mm long, 6,0-7,2mm wide, 5,0-5,5mm thick; seed coat from dark-gray (immature) to dull-black (mature), with a light brown scruffy covering ...... Merremia dissecta (Jacq.) Hall.f. (Figure 3) 10. Semihalo and hilum whitish; seed 4,5-4,7(-5,0)mm long, 4,2-5,0mm wide, 3,8mm thick; seed coat from orange (immature) to dark-reddish-brown (mature) ...... Merremia aegyptia (L.) Urban (Figure 3) 9. Semihalo pilose and dark. 11. Semihalo with light tawny hairs; hilum black; seed with 4,0-4,6mm long, 3,5-4,0(-4,5)mm wide, 2,5-3,5mm thick; seed coat from orange (immature) to dark-brown (mature), covered with a fine waxlike layer (lighter) and numerous, simple and fasciculated white-translucent hairs...... Merremia cissoides (Lam.) Hall.f. (Figure 3) 11. Semihalo with white-translucent hairs; hilum brownish-black; seed 3,8-4,2mm long and 3,7-3,9mm in diameter; seed coat from dark-greyish-brown (immature) to black (mature), with numerous, simple and tiny white- translucent hairs ...... Merremia macrocalyx (Ruiz et Pav.) O’Donell (Figure 3) 8. Hilum without these characteristics. 12. Seed coat glabrous. 13. Seed 10-13(-14)mm long, 8,0-10,0mm wide, 6,0-9,0mm thick, from subglobose to ovoid-shaped; seed coat monochrome light ochre, brown, or blackish-brown; hilum 2,5-3,0mm in diameter ...... L. (Figure 1) 13. Seed 10mm or less in length. 14. Hilum about 0,8mm or more in diameter. 15. Hilum glabrous. 16. Seed 8,5-9,5mm long, 6,5-7,5mm wide, 4,5-5,5mm thick; hilum 1,2-1,5mm, partially sunken, dark-brown or black ...... Ipomoea turbinata Lag. (Figure 3) 16. Seed 4,0-5,0mm long, 3,8-5,0mm wide, 3,0-3,2mm thick; hilum 0,8-1,0mm, not sunken, reddish or lighter in colour than the seed coat...... L. (Figure 2) 15. Hilum 0,8mm in diameter and concealed by a pad of tawny hairs; seed 3,8-4,8mm long, 3,8-4,0mm wide, 2,8-3,0mm thick ...... Ipomoea martii Meissn. (Figure 2) 14. Hilum seldom as much as 0,5mm in diameter. 17. Seed with obtuse apex. 18. Seed coat brown to nearly black; hilum yellowish-orange or lighter than the seed coat ...... L. (Figure 2) 18. Seed coat light or dark-yellowish-brown and irregularly dark mottled; hilum dark-reddish- brown ...... Ipomoea tiliacea (Willd.) Choisy (Figure 2)

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Ipomoea fimbriosepala Ipomoea triloba L. Choisy

Ipomoea floribunda G.Don

Ipomoea tiliacea (Willd.) Choisy (L.) Jacq.

Ipomoea hederifolia L.

Ipomoea martii Meissn.

Ipomoea indivisa (Vell.) Hall.

Ipomoea lacunosa L.

Ipomoea pes-caprae (L.) R.Br. (L.) Roth

Ipomoea pandurata (L.) G.F.W.Meyer

FIG. 2. Ventral view of Convolvulaceae weed seeds.

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Ipomoea rubiflora O´Donell (L.) Roth

Ipomoea quamoclit L.

Ipomoea turbinata Jacquemontia densiflora Lag. Hall.f.

Ipomoea wrightii A.Gray

Jacquemontia tamnifolia (Jacq.) Hall.f. Merremia aegyptia (L.) Urban.

Jacquemontia velutina Choisy

Merremia cissoides Merremia macrocalyx (Lam.) Hall.f. (Ruiz et Pav.) O´Donell

Merremia dissecta (Jacq.) Hall.f. FIG. 3. Ventral view of Convolvulaceae weed seeds.

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17. Seed with acute apex; seed coat yellowish-brown to brown; hilum yellowish-orange or reddish- brown ...... Ipomoea ramosissima (Poir.) Choisy (Figure 1) 12. Seed coat pubescent. 19. Seed 5,5-6,0mm long, 4,0-4,5mm wide, 3,0mm thick; seed coat with lanate hairs (10mm or more in length) resticted to a small cluster near the apex, on each margin; hilum 0,9-1,0mm in diameter...... Ipomoea wrightii A.Gray (Figure 3) 19. Seed without lanate hairs. 20. Seed with tawny hairs. 21. Tawny fringed hairs (10-13mm long) restricted to the apex and along 2/3 of the margins; seed 5,0-6,0mm long, 3,5-4,0mm wide, 2,0-2,5mm thick ...... Ipomoea floribunda G.Don (Figure 2) 21. Hairs not fringed. 22. Tawny hairs restricted to an attachment area. 23. Hairs restricted to the margins and occasionally to the keel. 24. Seed coat with densely hirsute pale tawny hairs adpressed from the base to the apex and hairs about 13mm long restricted to the attachment area; seed 5,0-6,5mm long 3,0-3,3mm wide, 1,8-2,2mm thick, obovate or elliptic in outline, acute apex and base; hilum 0,5mm in diameter, concealed by a pad of pale tawny hairs ...... Morong (Figure 1) 24. Seed coat with densely, tiny and short tawny hairs and numerous soft hairs (4-5mm long) restricted to the attachment area; seed 5,0-7,0(-9,0)mm long, (3,0-)4,0-5,0mm wide, 3,0- 4,0(-5,0)mm thick, obovate in outline; hilum 1,0-1,5mm in diameter and glabrous ...... (L.) G.F.W.Meyer (Figure 2) 23. Hairs concealed at the hilum. 25. Hilum 1,0-1,1mm long, 0,6-0,7mm wide, with simple tawny hairs; seed (4,0-)4,5-6,0mm long, 3,0-3,5(-4,0)mm wide, (3,0-)3,5-4,0mm thick; seed coat with black tomentum and diminute tawny hairs ...... Ipomoea nil (L.) Roth (Figure 2) 25. Hilum about 1,0mm in diameter; seed 4,2-5,0mm long, 2,8-3,5mm wide, 3,2-3,5mm thick; seed coat with black tomentum and short silvery hairs ...... (Burm.f.) Merr. (Figure 1) 22. Hairs 5,0-7,0mm or more in length, covered the seed coat except the keel area; seed 8,5-9,5mm long, 5,5-6,0mm wide, 4,8-5,2mm thick ...... Ipomoea carnea Jacq. ssp. fistulosa (Mart.ex Choisy) D.Austin (Figure 1) 20. Seed without tawny hairs. 26. Tomentum cupper coloured; soft and long (5,0mm or more in length) white hairs restricted to the margins; seed 4,4-4,8(-6,0)mm long, 4,8-5,0(-6,0)mm wide, 4,0-5,0mm thick ...... (L.) Sweet. (Figure 1) 26. Tomentum brown to black coloured. 27. Tomentum velvety medium brown and with densely short pale hairs; hilum 2,5-3,0mm in diameter; seed 6,5-7,0mm long and wide, 4,0-5,0mm thick ...... Ipomoea pes-caprae (L.) R.Br. (Figure 2) 27. Tomentum not velvety and hilum less than 2,5mm in diameter. 28. Margins delimited. 29. Margins with a narrow line; seed 4,7-5,5mm long, 3,0-4,0mm wide, 3,0-3,5mm thick, with black tomentum and simple, diminute and sparse translucent hairs, which give a mottled or grey-black appearance ventral faces without wrinkles ...... Ipomoea hederacea (L.) Jacq. (Figure 2) 29. Margins usually with a ridge; seed 3,5-4,5mm long, 2,5-3,0mm wide, 3,0-3,5mm thick, with black tomentum and simple, diminute and sparse translucent hairs, which give a grey-black appearance, ventral faces usually with one or two transversal wrinkles ...... Ipomoea purpurea (L.) Roth (Figure 3)

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28. Margins not delimited. 30. Hilum glabrous and without tomentum. 31. Hilum yellow; seed with black tomentum, 5,0-5,7mm long, 4,2-4,5mm wide, 4,0-4,2mm thick ...... Ipomoea fimbriosepala Choisy (Figure 2) 31. Hilum yellowish-brown; seed with reddish-brown tomentum on the faces and dark- brown on the keel, margins and dorsal side, 6,0-7,6(-8,0)mm long, 4,0-5,0(-5,4)mm wide, (3,2-)3,5-4,5mm thick .... (Desr.) Roem. & Schult. (Figure 1) 30. Hilum pilose or with black tomentum. 32. Hilum with black tomentum; seed 5,0-5,5mm long, 2,1-2,3mm wide, 1,9-2,1mm thick, with black or dark-brown tomentum and diminute whitish-translucent hairs, in small clusters, which give the roughned and irregular grey-mottled appearance ...... L. (Figure 3) 32. Hilum pilose. 33. Seed mottled, 3,5-4,0mm long, 3,0mm wide, 2,2-2,5mm thick, with light greyish-brown and light reddish-brown tomentum; hilum 0,8mm in diameter, light reddish-brown and with long yellowish-translucent hairs ...... Ipomoea rubriflora O’Donell (Figure 3) 33. Seed not mottled. 34. Hilum with long adpressed and densely, simple, thick, tawny-translucent hairs, from the margin to the seed base; seed 3,5-3,8mm long, 3,0-3,5mm wide, 2,0- 2,8mm thick, with black tomentum ...... L. (Figure 2) 34. Hilum without these characteristics. 35. Seed 3,0-3,5(-4,1)mm long, (2,5-)3,0-3,5mm wide, 2,0-2,8mm thick, with an uniform dark-brown to black tomentum and sparse short translucent hairs ...... L. (Figure 1) 35. Seed 3,0-4,0mm long and wide, 2,0-2,5(-3,0)mm thick, with dark-brown to black tomentum, margins and dorsal lengthwise areas with long copper- translucent hairs, faces and keel with irregular copper or yellowish- translucent hairs ...... Ipomoea indivisa (Vell.) Hall. (Figure 2)

Consideration about the species (Synonym: Dichondra repens Forst & Forst var. microcalyx Hall.f.) y Convolvulus crenatifolius Ruiz et Pav. (Figure 1) Annual or perennial creeping herb; in Brazil this specie is (Synonyms: Convolvulus montevidensis Spreng.; Convolvulus widely known as Dichondra repens Forst & Forst; montevidensis Spreng. var. megapotamicum Meissn.) description of fruit and seed in Groth (1997b) and Kissmann Perennial herb that trail on the ground or climb on other & Groth (1992; 1999). plants or fences; description of fruit and seed in Groth y Ipomoea alba L. (Figure 1) (1997b); occur in cultivated areas in south of Brazil. (Synonyms: Convolvulus aculeatus L.; Ipomoea bona-nox y Convolvulus arvensis L. (Figure 1) L.; Convolvulus bona-nox (L.) Spreng.; Calonyction (Synonym: Calystegia arvensis L.) aculeatum (L.) Bojer) Perennial herb; plant very aggressive; description of fruit Perennial herbaceous plant, climbing habit; very similar and seed in Groth (1984b) and Kissmann & Groth (1992; specie to I. turbinata Lag.; seeds of the two species are also 1999). very similar; the fruit is a consistent characteristic to y Cuscuta spp. (Figure 1) distinguish the two species, which is erect in I. alba and Voluble herb; description of fruit and seed in Groth (1984b) reflex in I. turbinata; description of fruit and seed in Groth and Kissmann & Groth (1992; 1999). (1997a) and Kissmann & Groth (1992; 1999); occur in the y Dichondra microcalyx (Hall.f.) Fabris (Figure 1) Northeastern Regions of Brazil.

Revista Brasileira de Sementes, vol. 23, nº 2, p.1-13, 2001 DESCRIÇÃO DE ESPÉCIES INVASORAS DA FAMÍLIA CONVOLVULACEAE 9 y Ipomoea amnicola Morong (Figure 1) consistent difference between the two species is the seed, Annual vining herb; when the plant did not present mature therefore, exist a great uncertainness for the correct fruits it is very similar to I. ramosissima (Poir.) Choisy (Si- identification of this specie in Brazil; description of fruit mão-Bianchini, 1998); the most consistent characteristic and seed in Groth (1997a) and Kissmann & Groth (1992; between the two species is the seed; description of fruit and 1999); weed in annual cultures. seed in Groth (1997b); weed in cultivated areas. y Ipomoea fimbriosepala Choisy (Figure 2) y Ipomoea aristolochiaefolia (H.B.K.) G.Don (Synonym: Aniseia hastata Meissn.) (Synonym: Convolvulus aristolochiaefolius Kunth) Annual vining herb; very close specie to I. setifera Poir.; Annual vining herb; very similar specie to I. ramosissima fruit and seed was described as Aniseia hastata Meissn. in (Poir.) Choisy; in relation to the identification of plants Groth & Boaretto (1991); weed in cultivated areas, as and seeds of I. aristolochiaefolia, in Brazil, have a great pastures. confusion and were mentions as if they were from this y Ipomoea floribunda G.Don (Figure 2) specie (Groth, 1984b; Koehn, 1977; Leitão-Filho et al., Annual herbaceous plant; description of fruit and seed in 1972; Lorenzi, 1982), but in reality they were from I. Groth (1998). triloba L. y Ipomoea hederacea (L.) Jacq. (Figure 2) y Ipomoea asarifolia (Desr.) Roem. & Schult. (Figure 1) (Synonym: Convolvulus hederaceus L.) (Synonym: Convolvulus asarifolius Desr.) Annual herbaceous plant; very similar specie to I. nil (L.) Perennial herbaceous plant, climbing habit; very similar Roth; I. hederacea was confound erroneously with I. specie to I. subtomentosa (Chodat & Hassl.) O’Donell and acuminata (Vahl) Roem.et Schult.; description of fruit and to I. maurandioides Meissn. (Simão-Bianchini, 1998); the seed in Groth (1984a-b) and Kissmann & Groth (1992; leaf blade of the plant is very similar to the of I. pes-caprae 1999); weed in annual fields, as maize and other crops. (L.) Sweet. (see comments of this specie); the most y Ipomoea hederifolia L. (Figure 2) consistent characteristic to distinguish this two species is (Synonyms: Quamoclit hederifolia (L.) G.Don; Ipomoea the seed; description of fruit and seed in Groth (1998) and coccinea var. hederifolia (L.) A.Gray; Ipomoea angulata Kissmann & Groth (1992; 1999); occur from the littoral Lam.; Quamoclit angulata (Lam.) Bojer) North of Brazil to the state of São Paulo. Annual herbaceous plant; very close specie to I. indivisa y Ipomoea cairica (L.) Sweet. (Figure 1) (Vell.) Hall.; in the United States it was considered as a (Synonym: Convolvulus cairicus L.) variety of I. coccinea L.; the most consistent characteristic Perennial herbaceous plant, climbing; description of fruit to distinguish the two species is the seed; description of and seed in Groth (1984b), Groth & Boaretto (1985) and fruit and seed in Groth (1984b) and Kissmann & Groth Kissmann & Groth (1992; 1999); the most common weed (1992; 1999). in cultivated areas, as cotton, sugar-cane, pastures, citrus, y Ipomoea indica (Burm.f.) Merr. (Figure 1) coffee and vine. (Synonyms: Convolvulus acuminatus Vahl; Ipomoea y Ipomoea carnea Jacq. (Figure 1) congesta R.Br.; Ipomoea mutabilis Lindl.; Ipomoea (Synonyms: Convolvulus carneus (Jacq.) Spreng.; Ipomoea acuminata (Vahl) Roem. et Schult.) fistulosa Mart.ex Choisy; Ipomoea carnea spp. fistulosa Perennial vining herb; stoloniferous; polymorphic specie; (Mart.ex Choisy) D.Austin) during a long-time plants of I. nil (L.) Roth was identified Perennial shrubs; in Brazil the specie is known as I. fistulosa erroneously as I. acuminata (Leitão-Filho et al., 1972; Mart.ex Choisy and with this designation fruit and seed were Koehn, 1977). Nowadays the valid name is I. indica described in Groth (1991) and as I. carnea in Kissmann & (Burm.f.) Merr. (Simão-Bianchini, 1998); the seeds Groth (1992; 1999). According to Austin (1977) this specie presented consistent characteristics to distinguish I. nil from is only a subspecie of I. carnea; weed in cultivated areas, I. indica; the description of the seed was made as I. congesta as pastures. R.Br. in Groth (1998); weed in annual cultures. y Ipomoea coccinea L. (Figure 1) y Ipomoea indivisa (Vell.) Hall. (Figure 2) (Synonym: Quamoclit coccinea (L.) Moench) (Synonyms: Convolvulus indivisus Vell.; Ipomoea coccinea Annual vining herb; very similar specie to I. indivisa (Vell.) auct.non L., Meissn.; Quamoclit indivisa (Vell.) Hall.f.) Hall. and usually confused with this specie; the most Annual herbaceous plant, climbing habit and with twining

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branches; in Brazil this specie had been identified incorrectly plants identified as I. pes-caprae looks like I. asarifolia as I. coccinea L., because the plants of the two species and this evidence put in doubt the identification of this specie presented similar characteristics, but the more consistent in Brazil. However, the characterization of I. pes-caprae difference is the seed; description of fruit and seed in Groth seeds, identified in the United States and compared with (1997a) and Kissmann & Groth (1992; 1999); weed in the description of Gunn (1969), shows that exist a consistent annual and perennial cultures. difference between seeds of the two species. It was also y Ipomoea lacunosa L. (Figure 2) observed that seeds collected in Northeast Regions are not (Synonyms: Convolvulus lacunosus Spreng.; Quamoclit from I. pes-caprae, in reality they are I. asarifolia. lacunosa Rafin.) Description of fruit and seed in Groth (1997a) and Kissmann Annual herbaceous plant, climbing habit; description of fruit & Groth (1992; 1999); commonly planted for fixation of and seed in Groth (1997a); weed in cultivated and non the sand dunes and weed in cultures near the coast. cultivated areas. y Ipomoea purpurea (L.) Roth (Figure 3) y Ipomoea martii Meissn. (Figure 2) (Synonyms: Convolvulus purpureus L.; Pharbites purpurea (Synonym: Rivea cordata Choisy) (L.) Voigt.) Annual herbaceous plant in the beginning and after ligneous, Annual herbaceous plant, climbing habit; very close specie climbing habit; description of fruit and seed in Groth to I. nil (L.) Roth and I. indica (Burm.f.) Merr. and they are (1997b); weed in cultivated areas, as pastures. easy separated by the sepals; description of fruit and seed y Ipomoea nil (L.) Roth (Figure 2) in Groth (1984b), Groth & Boaretto (1985) and Kissmann (Synonyms: Convolvulus nil L.; Pharbitis nil (L.) Choisy; & Groth (1992; 1999); occur as weed in annual e perennial Ipomoea longicuspis Meissn.) cultures in al Brazilian country and are more frequent in Annual herbaceous plant, climbing or decumbent habit; very the Center-West Region of Brazil. close specie to I. hederacea (L.) Jacq.; in Brazil I. nil had y Ipomoea quamoclit L. (Figure 3) been identified incorrectly as I. acuminata (Vahl) Roem.et (Synonyms: Quamoclit vulgaris Choisy; Quamoclit pinnata Schult. (Groth, 1984b; Lorenzi, 1982) and as I. longicuspis (Desr.) Bojer (Simão-Bianchetti); Quamoclit quamoclit (L.) Meissn. (Leitão-Filho et al., 1972; Groth, 1991); description Britton.) of fruit and seed in Groth (1997a) and Kissmann & Groth Annual herbaceous plant, climbing habit; description of fruit (1992; 1999), as I. acuminata (Vahl) Roem.et Schult. in and seed in Groth (1984a-b) and Kissmann & Groth (1992; Groth (1984b) and as I. longicuspis in Groth (1991); one of 1999). the most frequent weed after I. triloba L.; weed in cultivated y Ipomoea ramosissima (Poir.) Choisy (Figure 1) areas, as soybean, maize, pastures and coffee. (Synonyms: Convolvulus ramosissima Poir.; Ipomoea y Ipomoea pandurata (L.) G.F.W.Meyer (Figure 2) cynanchifolia Meissn.) Perennial herb; description of fruit and seed in Groth Annual herbaceous plant, climbing habit; during some time (1997b); weed in cultivated areas. this specie was identified as I. cynanchifolia Meissn. (Lei- y Ipomoea pes-caprae (L.) R.Br. (Figure 2) tão-Filho et al., 1972; Groth, 1984b; Kissmann & Groth, (Synonyms: Convolvulus pes-caprae L.; Quamoclit pes- 1992); Simão-Bianchini (1998) based on the analyzed caprae (L.) Maza; Convolvulus brasiliensis L.; Ipomoea characteristics, in the study of the species of Ipomoea, affirm brasiliensis (L.) G.F.W.Meyer) that “there exist no consistent characteristics that permit Cosmopolitan, perennial herbaceous plant, climbing habit the recognition of two species (I. cynanchifolia and I. and sometimes creeping; very close specie to I. asarifolia ramosissima), neither even varieties”; the seeds are very (Desr.) Roem.et Schult. (Simão-Bianchini, 1998) and the similar and for that reason are confused with seeds from I. most important difference between them is the leaf shape, triloba; seedlings are very similar to I. triloba L. and the which is suborbicular or oblong in I. pes-caprae, with deeply difference is the deep re-entrance between the lobs and if bilobed apex (spp. pes-caprae), remembering a crowbar, the external margin of the leaf is straight (I. ramosissima) and with emarginate apex (spp. brasiliensis); in I. asarifolia or inclined (I. triloba) to the outside; description of fruit is heart-shaped with roundish, acute and rarely acuminate and seed in Kissmann & Groth (1999) and as I. apex. It was observed that seeds collected at the littoral cynanchifolia Meissn. in Groth (1984b), Groth & Boaretto Northeast Regions of Brazil, by Kurt G. Kissmann, from (1985) and Kissmann & Groth (1992).

Revista Brasileira de Sementes, vol. 23, nº 2, p.1-13, 2001 DESCRIÇÃO DE ESPÉCIES INVASORAS DA FAMÍLIA CONVOLVULACEAE 11 y Ipomoea rubriflora O’Donell (Figure 3) Kissmann & Groth (1999), as I. aristolochiaefolia (Synonyms: Ipomoea hederifolia auct. non L.; Ipomoea (H.B.K.) G.Don in Groth (1984b) and as I. grandifolia cocccinea auct. non L.) (Dammer) O’Don. in Kissmann & Groth (1992); is the Annual herbaceous plant, climbing habit; description of fruit most important weed of the genus Ipomoea that occur and seed in Groth (1997a) and Kissmann & Groth (1992; in soybean cultures. 1999); weed in cultivated and non cultivated areas. y Ipomoea turbinata Lag. (Figure 3) y Ipomoea tiliacea (Willd.) Choisy (Figure 3) (Synonyms: Convolvulus muricatus L.; Ipomoea muricata (Synonym: Convolvulus tiliaceus Willd.) (L.) Jacq.; Calonyction muricatum (L.) G.Don) Annual herbaceous plant, with polymorphic aspect and leaf Annual herbaceous plant, climbing habit; specie very simi- shape with great diversity; in Brazil species of I. tiliacea lar to I. alba L.; many authors had used the name I. muricata and I. batatas (L.) Lam. are very different (Simão-Bianchini, (L.) Jacq. for the identification this specie (Simão-Bianchini, 1998); the description of the seed was done as I. hirta in 1998); the fruit presented consistent characteristics for the Groth (1998) and description of fruit and seed in Kissmann separation of the two species, which is erect in I. alba and & Groth (1999). reflex in I. turbinata; description of fruit and seed in Groth y Ipomoea triloba L. (Figure 3) (1997a); weed in cultivated areas, as soybean. (Synonyms: Quamoclit triloba (L.) G.Don; Jacquemontia y Ipomoea wrightii A.Gray (Figure 3) grandifolia Dammer; Ipomoea grandifolia (Dammer) (Synonyms: Convolvulus heptaphyllus Roxb.; Ipomoea O’Donell) heptaphylla (Roxb.) Voigt; Ipomoea pulchella Roth.; Annual herbaceous plant, climbing habit; in Brazil the specie Ipomoea spiralis House) was identified as I. aristolochiaefolia (H.B.K.) G.Don (Lei- Annual herbaceous plant, climbing habit; remember I. tão-Filho et al., 1972; Groth, 1984b; Koehn, 1977 e Lorenzi, cairica (L.) Sweet. and frequently so identified erroneously; 1982), establishing great confusion; but this specie is more a consistent difference between the two species are in the slender; after, based on the leaf shape it was identified as I seeds; description of fruit and seed in Groth (1997a) and grandifolia (Dammer) O’Donell, according to O’Donell Kissmann & Groth (1992; 1999). (1952) and Austin (1978). According to Burkart (1979), I. y Jacquemontia densiflora Hall.f. (Figure 3) grandifolia is not the synonym of I. triloba and Austin (1978) Annual herbaceous plant, climbing habit; description of seed have the same opinion and affirm that “various authors in Groth (1991) and Kissmann & Groth (1992; 1999); weed confound the two species”. O’Donell (1952) at that time in cultivated and non cultivated areas. had already refer this confusion and suggest that the species y Jacquemontia tamnifolia (L.) Griseb. (Figure 3) of I. triloba mentioned for South America will be “called” (Synonyms: Ipomoea tamnifolia L.; Convolvulus tamnifolia I. grandifolia. The difference was based, mainly, on the (L.) Meyer) shape and size of the sepals, which are bigger in I. Annual vines or herbs, climbing habit, very aggressive; grandifolia than in I. triloba. Simão-Bianchini (1998), description of fruit and seed in Groth (1997b) and Kissmann analysed Table 1 from O’Donell (1952), concluded, in his & Groth (1992; 1999). study about the Ipomoea from southeast of Brazil, that are y Jacquemontia velutina Choisy (Figure 3) a transition between the characteristics utilized in the Annual herbaceous plant, climbing habit; description of seed separation of the two species and so decided to consider in Groth (1991) and Kissmann & Groth (1992; 1999); weed the two as only one species, with cosmopolitan distribution. in cultivated areas. According to O’Donell (1952), I. grandifolia has bigger y Merremia aegyptia (L.) Urban (Figure 3) (3,5-4,0mm) seeds than I. triloba, while Austin (1992), (Synonyms: Ipomoea aegyptia L.; Convolvulus mentioned smaller sizes for I. grandifolia (2,0-2,5mm) than penthaphyllus L.; Ipomoea penthaphylla (L.) Jacq.; Ipomoea for I. cynanchifolia Meissn. (= I. ramosissima). Austin pilosa Cav.) (1978) mentioned for the fruit of I. triloba 5-6mm long and Annual herbaceous plant, climbing habit; description of fruit for I. grandifolia 6-7mm. I. triloba is narrowly related to I. and seed in Groth (1991) and Kissmann & Groth (1992; cordatotriloba Dennst. (Simão-Bianchini, 1998) and some 1999); weed in annual and perennial cultures. species presented intermediate characteristics with I. y Merremia cissoides (Lam.) Hall.f. (Figure 3) ramosissima; description of I. triloba fruit and seed in (Synonyms: Convolvulus cissoides Lam.; Convolvulus

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calycinus H.B.K.; Convolvulus guadalupensis Steud.; GROTH, D. Identificação botânica de plantas e sementes de espécies Ipomoea potentiloides Meissn.; Merremia potentiloides invasoras na cultura da soja. Revista Brasileira de Sementes, Brasília, v.2, n.3, p.59-95, 1980. (Meissn.) Hall.f.) GROTH, D. Caracterização morfológica das plântulas e das sementes Annual herbaceous plant, climbing habit; in Brazil occur de três espécies daninhas da cultura de soja. In: SEMINÁRIO two varieties: subsessile and viscid; seeds of the two varieties NACIONAL DE PESQUISA DE SOJA, 3, Campinas, 1984. does not present differential characteristics; description of Anais. Campinas: EMBRAPA/Centro Nacional de Pesquisa fruit and seed in Groth (1991) and Kissmann & Groth (1992; de Soja, 1984a. p.575-86. 1999); weed in annual and perennial cultures. GROTH, D. Unidades de dispersão e plântulas de espécies de plantas invasoras. Campinas: Instituto de Biologia/ y Merremia dissecta (Jacq.) Hall.f. (Figure 3) UNICAMP, 1984b. p.426-459. (Tese Doutorado). (Synonyms: Convolvulus dissectus Jacq.; Ipomoea sinuata GROTH, D. Morphological characterization of seeds and seedlings Ortega; Ipomoea dissecta (Jacq.) Pursh.; Operculina of seven weed species of Convolvulaceae occurring in dissecta (Jacq.) House; Ipomoea dissecta (Jacq.) Pers.) agricultural seeds in Brazil. Iheringia, Sér.Bot., Porto Alegre, Annual vines or herbs, climbing habit; in Brazil occur two n.41, p.83-99, Jun. 1991. varieties: typica and edentata (Meissn.) O’Donell; seeds of GROTH, D. Caracterização morfológica das sementes e frutos de the two varieties does not present differential characteristics; nove espécies invasoras de Ipomoea (Convolvulaceae). Revista Brasileira de Sementes, Brasília, v.19, n.2, p.303-314, 1997a. description of fruit and seed in Groth (1997b) and Kissmann & Groth (1992; 1999); weed in cultivated areas, as coffee, GROTH, D. Caracterização morfológica das sementes e frutos de oito espécies invasoras da família Convolvulaceae. Revista citrus and pastures. Brasileira de Sementes, Brasília, v.19, n.2, p.361-368, y Merremia macrocalyx (Ruiz et Pav.) O’Donell (Figure 3) 1997b. (Synonyms: Convolvulus glaber Aublet; Convolvulus GROTH, D. Morphological characterization of four weed seeds of macrocalyx Ruiz et Pav.; Ipomoea macrocalyx (Ruiz et Pav.) Ipomoea L. (Convolvulaceae). Revista Brasileira de Choisy; Merremia glabra (Aublet) Hall.f.) Sementes, Brasília, v.20, n.1, p.75-79, 1998. Annual vines or perennial herbs, depending from the GROTH, D. & BOARETTO, M.R. Estudo morfológico das environmental; climbing habit; description of fruit and seed sementes, frutos e plantas de quatro espécies invasoras do gênero Ipomoea L. Revista Brasileira de Sementes, Brasília, in Groth (1997b) and Kissmann & Groth (1992; 1999); weed v.7, n.3, p.79-97, 1985. in annual cultures, as crops and perennial, such as coffee. GROTH, D.; SILVA, H.T & WEISS, B. Caracterização botânica de plantas de espécies invasoras e respectivas sementes na cultura da soja (Glycine max (L.) Merrill.) no Rio Grande do Sul. In: CONCLUSIONS SEMINÁRIO NACIONAL DE PESQUISA DE SOJA, 1, Londrina, 1978. Anais. Londrina: Centro Nacional de Pesquisa The shape of Convolvulaceae weed seeds species can not de Soja - EMBRAPA, 1979. p.187-202. be used as a differential characteristic because it depends of the GROTH, D.; BOARETTO, M.R. & SILVA, R.N. Morfologia de number of mature seeds developed in the fruit. The more stable sementes, frutos e plantas invasoras em algumas culturas. Revista Brasileira de Sementes, Brasília, v.5, n.3, p.151-82, morphological characteristics which permited the identification 1983. of the species are the colour and the pilosity of the seed coat GUNN, C.R. Seeds of the United States noxious and common weeds (surface), the hilum area and the outline of the hilum. in the Convolvulaceae excluding the genus Cuscuta. Proceedings AOSA, Geneva, v.59, p.101-115, 1969. GUNN, C.R. Seed collecting and identification. In: KOZLOWSKY, REFERENCES T.T. Seed biology. New York, Academic Press, 1972. v.3, p.55-69. AUSTIN, D.F. The Ipomoea batatas complex - I. Taxonomy. Bull.Torrey Bot.Club., v.105, n.2, p.114-129, 1978. HARPER, J.L. The biology of weeds. Oxford: Blackwell Scientific Publications, 1960. p.1-9. AUSTIN, D.F. Ipomoea carnea Jacq. vs. Ipomoea fistulosa Mart.ex Choisy. Taxon, v.26, n.2/3, p.235-238, 1977. KISSMANN, K.G. & GROTH, D. Plantas infestantes e nocivas. São Paulo: BASF Brasileira, 1992. tomo 2, p.504-609. AUSTIN, D.F. Seeds in some poorly known species of Ipomoea section Batatas (Convolvulaceae). Bull.Torrey Bot. Club., KISSMANN, K.G. & GROTH, D. Plantas infestantes e nocivas. v.119, n.2, p.142-144, 1992. 2.ed. São Paulo: BASF Brasileira, 1999. tomo 2, p.617-738. BURKART, A. Flora ilustrada de Entre Rios (Argentina). Buenos KOEHN, D. Identificação de algumas invasoras encontradas em Aires: INTA, 1979. pt.5, tomo 6 e 5, p.148-194. sementes das principais espécies forrageiras, produzidas no Rio

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Grande do Sul. Boltim Técnico do IPAGRO, Porto Alegre, Archivos do Museu Paranaense, Curitiba, v.4, p.207-244, v.1, p.3-96, 1977. 1952. LEITÃO-FILHO, H.F.; ARANHA, C. & BACCHI, O. Plantas SIMÃO-BIANCHINI, R. Ipomoea L. (Convolvulaceae) no sudeste invasoras de culturas no estado de São Paulo. São Paulo: do Brasil. São Paulo: Instituto de Biociências/USP, 1998 476p. HUCITEC, 1972. p.39-74. (Tese Doutorado). LORENZI, H. Plantas daninhas do Brasil. Nova Odessa: H. UNITED STATES. Department of Agriculture. Selected weeds of Lorenzi, 1982. p.115-129. the United States. Washington: Agricultural Research Service, O’DONELL, C.A. Convolvulaceas americanas nuevas o criticas III. 1970. p.1-3. (Agriculture Handbook, 366).

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