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Accepted by A. Monro: 7 Aug. 2010; published: 31 Aug. 2010 1 PHYTOTAXA ISSN 1179-3155 (print edition) ISSN 1179-3163 (online edition) Copyright © 2010 Magnolia Press Phytotaxa 8: 124 (2010) www.mapress.com/ phytotaxa/ Article Climbing plants of a fragmented area of lowland Atlantic Forest, Igarassu, Pernambuco (northeastern Brazil) DIOGO ARAÚJO 1 & MARCCUS ALVES 2 1 Programa de Pós-Graduação em Biologia Vegetal, Dept. Botânica, Universidade Federal de Pernambuco. CEP: 50670-901, Recife, Pernambuco, Brazil; e-mail: [email protected] 2 Dept. Botânica, Universidade Federal de Pernambuco. CEP: 50670-901, Recife, Pernambuco, Brazil. Abstract This study was undertaken in fragments of lowland Atlantic Forest surrounded by sugar cane plantations in Igarassu, Pernambuco, Brazil. Amongst the 93 species recorded for the locality, 30 species have a climbing habit. Passiflora is among the most species-rich genera which is novel in a floristic study. Morphological characters such as the presence and position of a tendril, limb form and flower symmetry are the most important for species identification. Here we present a species checklist for the locality, annotated with distribution data. An illustrated key to the climbing species found in Igarassu is also provided. Key words: checklist, floristics, lianas, Passifloraceae Introduction Climbing plants are defined as plants incapable of autonomous vertical support once they reach a certain height (ca. 1 m tall) and depend on other plants for support in their natural environment (Gentry 1991). Lianas are defined as woody climbing plants (Putz 1984, Gerwing et al. 2006). Twining habit is a very common amongst climbing plants; this consists of the stems twisting around the branches or stems of a host with or without the aid of specialized structures such as tendrils (Darwin 1867, Gentry 1985). The climbing habit has arisen several times in the evolutionary history of Angiosperms, and this has resulted in a great taxonomic diversity of climbing plants (Gentry 1985). Families such as Smilacaceae, Menispermaceae, Passifloraceae, Cucurbitaceae and Convolvulaceae are essentially entirely composed of or dominated by species with a climbing habit. According to Gentry (1991), New World families with the highest diversity of climbing plants are Apocynaceae (esp. Asclepiadoideae), Convolvulaceae and Fabaceae. In the Brazilian flora, 86 of the 217 native families cited by Souza & Lorenzi (2008) have at least one species with a climbing habit. In his discussion of the climbing plants of Puerto Rico and the Virgin Islands, Acevedo-Rodríguez (2005) identifies lowland tropical forest as including the highest number of climbing species. According to Schnitzer & Bongers (2002) these species play an important role in vegetation dynamics, although the exact contribution of woody climbing plants in tropical forests biomass is currently unresolved. Hegarty & Cabalé (1991) estimated that up to 40% of the total leaf area and leaf productivity in a forest may be represented by climbers. Climbing species may also represent a significant component of the plant diversity of an area, accounting for ca 25% of the woody plant species (Gentry & Dodson 1987). The ecological importance of climbing plants, forming physical bridges for the fauna between individual trees in a forest was noted by Darwin (1867) and Hegarty (1991). Some climbers also have strict ecological relationships, for example, Heliconius butterflies and Passifloraceae species, Ithomiinae butterflies and Solanaceae species, and Buprestidae beetles with Bignoniaceae species (Odegaard 2000).

Climbing plants of a fragmented area of lowland Atlantic Forest, Igarassu, Pernambuco (northeastern Brazil)

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Page 1: Climbing plants of a fragmented area of lowland Atlantic Forest, Igarassu, Pernambuco (northeastern Brazil)

PHYTOTAXAISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2010 • Magnolia Press

Phytotaxa 8: 1–24 (2010) www.mapress.com/phytotaxa/ Article

Climbing plants of a fragmented area of lowland Atlantic Forest, Igarassu, Pernambuco (northeastern Brazil)

DIOGO ARAÚJO1 & MARCCUS ALVES2

1 Programa de Pós-Graduação em Biologia Vegetal, Dept. Botânica, Universidade Federal de Pernambuco. CEP: 50670-901, Recife, Pernambuco, Brazil; e-mail: [email protected] Dept. Botânica, Universidade Federal de Pernambuco. CEP: 50670-901, Recife, Pernambuco, Brazil.

Abstract

This study was undertaken in fragments of lowland Atlantic Forest surrounded by sugar cane plantations in Igarassu, Pernambuco, Brazil. Amongst the 93 species recorded for the locality, 30 species have a climbing habit. Passiflora is among the most species-rich genera which is novel in a floristic study. Morphological characters such as the presence and position of a tendril, limb form and flower symmetry are the most important for species identification. Here we present a species checklist for the locality, annotated with distribution data. An illustrated key to the climbing species found in Igarassu is also provided.

Key words: checklist, floristics, lianas, Passifloraceae

Introduction

Climbing plants are defined as plants incapable of autonomous vertical support once they reach a certain height (ca. 1 m tall) and depend on other plants for support in their natural environment (Gentry 1991). Lianas are defined as woody climbing plants (Putz 1984, Gerwing et al. 2006). Twining habit is a very common amongst climbing plants; this consists of the stems twisting around the branches or stems of a host with or without the aid of specialized structures such as tendrils (Darwin 1867, Gentry 1985).

The climbing habit has arisen several times in the evolutionary history of Angiosperms, and this has resulted in a great taxonomic diversity of climbing plants (Gentry 1985). Families such as Smilacaceae, Menispermaceae, Passifloraceae, Cucurbitaceae and Convolvulaceae are essentially entirely composed of or dominated by species with a climbing habit. According to Gentry (1991), New World families with the highest diversity of climbing plants are Apocynaceae (esp. Asclepiadoideae), Convolvulaceae and Fabaceae. In the Brazilian flora, 86 of the 217 native families cited by Souza & Lorenzi (2008) have at least one species with a climbing habit.

In his discussion of the climbing plants of Puerto Rico and the Virgin Islands, Acevedo-Rodríguez (2005) identifies lowland tropical forest as including the highest number of climbing species. According to Schnitzer & Bongers (2002) these species play an important role in vegetation dynamics, although the exact contribution of woody climbing plants in tropical forests biomass is currently unresolved. Hegarty & Cabalé (1991) estimated that up to 40% of the total leaf area and leaf productivity in a forest may be represented by climbers. Climbing species may also represent a significant component of the plant diversity of an area, accounting for ca 25% of the woody plant species (Gentry & Dodson 1987). The ecological importance of climbing plants, forming physical bridges for the fauna between individual trees in a forest was noted by Darwin (1867) and Hegarty (1991). Some climbers also have strict ecological relationships, for example, Heliconius butterflies and Passifloraceae species, Ithomiinae butterflies and Solanaceae species, and Buprestidae beetles with Bignoniaceae species (Odegaard 2000).

Accepted by A. Monro: 7 Aug. 2010; published: 31 Aug. 2010 1

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Climbing plants have often been neglected by plant collectors in the past (Gentry 1991), probably due to poor visibility and accessibility to the canopy. More recently there has been an increase in the number of floristic and taxonomic studies that have focused exclusively on species with a climbing habit (Solórzano et al. 2002, Reddy & Parthasarathy 2003, Acevedo-Rodriguez 2005). In Brazil, floristic and taxonomic studies that focus on or include climbing plants are restricted to seasonal semideciduous forest areas in the southeast (Hora & Soares 2002, Udulutsch et al. 2004, Rezende & Ranga 2005, Tibiriçá et al. 2006) and to date few studies have focused on climbing plants from the northeastern extension of the Brazilian Atlantic forests, an area critical for conservation.

The aim of this study is to provide the first floristic inventory of climbing plants for a lowland tropical rain forest in Brazil, as well as the first identification key to this group of plants for northeastern Brazil. It is hoped that a consequence of filling a gap in our taxonomic knowledge about this aspect of the local flora will encourage researchers of Neotropical forests to undertake more ecological and taxonomical studies of climbing plants.

Methods

The study areaIgarassu is located on the northern coast of the state of Pernambuco, Brazil. The major land use for the

area is plantations of sugar cane (CPRH 2003). Igarassu is included in the Mata Atlântica Biosphere Reserve an area of ’extreme biological importance’ (MMA 2000). Igarassu has an average annual temperature of 24.9°C, with minima of 21°C (May to June) and maxima of 25ºC (July to December). It receives an annual precipitation of 1444 mm (recorded 1998 and 2004), with the average monthly precipitation greater than 100 mm from January through August and less than 100 mm from September through December (Silva 2004). The terrain consists of hills with flattened tops intersected by floodplains and river terraces, slopes ranging from flat to greater than 30% where artificial lakes are frequently located (CPRH 2003) for water catchment and electricity production. The soil is predominantly a mixture of sandy-clay and non-consolidated sediments (CPRH 2003).

This study was undertaken in a fragmented forest landscape of 76.7 km2 (including secondary forest), where fragment size, shape and topography was variable (Table 1). Fragments (identified as Macacos, Pezinho, Piedade, Santa Helena, Vespas and Zambana), are surrounded by sugarcane plantations. The area is owned by the Usina São José factory (USJ), a processing plant that produces sugar and alcohol.

TABLE 1. Geographic data of the six selected forest fragments in the studied area.

Fragment Geographic coordinates Elevation (min–max) Total area (ha)

Piedade 7°49’12”, 7°50’55” S 35°0’35”, 34°59’21” W 30–113 m 305,787 ha

Zambana 7º41’29”, 7º43’09”S 35º0’8”, 34º58’38”W 20–115 m 387,851 ha

Macacos 7º46´02", 7º47’40” S 35º01´06", 34º59’42”W 40–155 m 356,926 ha

Pezinho 7°47’27”, 7°48’01” S 35°01’25”, 35°01’00”W 50–125 m 29,890 ha

Vespas 7º44’50”, 7º45’08” S 34º59’02”, 34º58’45”W 45–105 m 13,807 ha

Sta. Helena 7°46’59”, 7°47’16”S 34°56’41”, 34°56’29”W 20–85 m 11,854 ha

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Data collection Monthly excursions were made between February 2007 and May 2008. Each month 4–6 days of

fieldwork were undertaken, with at least 6 hours a day spent collecting. A total of 100 days of fieldwork were completed. Sample points were located with the aid of a GPS that also improved the re-location of sites to improve the sampling effort of all forest fragments. Larger forest fragments were sampled several times. The collected specimens were deposited at UFP with duplicates at IPA, CEPEC, ULM, NY and K.

Taxonomic treatment Plant identifications were undertaken by the authors and specialists of taxonomic groups to whom

duplicate samples were sent. Identification by the authors was based on herbarium specimens (IPA, UFP, PEUFR, CEPEC, HUEFS and RB), using morphological characters together with the relevant literature for each family. Duplicate specimens were sent to the following specialists, who all provided identifications in return: Asteraceae: Rita de Cássia Pereira (IPA); Sapindaceae: Marcondes Oliveira (UFP); Euphorbiaceae: Maria de Fátima Lucena (UFP); Fabaceae: Elizabeth Córdula (UFP), Marli PiresMorin (RB); Malpighiaceae: André Amorim (CEPEC); Convolvulaceae: Maria Teresa Vital (CEPEC); Passifloraceae: Teonildes Nunes (HUEFS). Other families were identified by the authors.

Species are assigned to families according to APG III (2009) and author names follow Brummit & Powell (1992). Herbarium acronyms follow Index Herbariorum (ht tp: / /sciweb.nybg.org/science2/IndexHerbariorum.asp). The order of country citation in the geographical distribution follows that of Flora Neotropica.

Results and discussion

The number of species found in the studied area is close to that recorded for southern Brazil (Lima et al. 1997,Udulutsch et al. 2004, Rezende & Ranga 2004, Tibiriçá et al. 2006). A total of 93 species of climbing plants were collected, of which 30 are considered lianas and 63 are herbaceous climbing plants. Fabaceae, Convolvulaceae, Passifloraceae and Cucurbitaceae were the most species-rich plant families represented. Combined, these families represent 50% of the total climbing plant diversity sampled. The remainder is split between 22 families.

Asteraceae, Bignoniaceae, Malpighiaceae and Sapindaceae were amongst the most commonly collected climbing plant families registered in inventories in southeast Brazil (Tibiriçá et al. 2006, Lima et al. 1997, Hora & Soares 2004, Udulutsch et al. 2004, Rezende & Ranga 2004). This agrees with results for inventories undertaken in other lowland forest formations below 1000 m elevation (Gentry 1991). The results of this inventory differ from the above study in the following respects: Fabaceae is the most diverse family with respect to in the number of genera (nine) recorded in Igarassu. According to Gentry (1991), this family is the third most species-rich climbing plant family in the Neotropics, especially in the subfamily Faboideae, with 15 species. The other two most diverse climbing plants families in the Neotropics, Asclepiadaceae (sensuCronquist 1981, now included in Apocynaceae) and Convolvulaceae are rarely dominant in Tropical Forests (Gentry 1991). In our study we found Convolvulaceae to be amongst the most representative. The low species richness of Apocynaceae can be explained by the preference of species in that family to shrubby, drier vegetation types, such as “caatinga” and “cerrado” of Central Brazil.

In our study Passiflora L. (9 species; Passifloraceae) is the most diverse genus among the 55 species found, followed by Ipomoea L. (5 species; Convolvulaceae), and Canavalia DC., Centrosema (DC.) Benth. (Fabaceae), Gurania (Schltdl.) Cogn. (Cucurbitaceae) and Paullinia L. (Sapindaceae; 3 species each). Six of these genera belong to the families identified by Gentry (1991) as richest in the Neotropics with respect to climbing species. Passiflora were found for the first time in a published inventory of climbing plants as the most species-rich genus in a Neotropical forest.

Even though Gentry (1991) stated that climbing plants are an important part of the botanical diversity in in tropical forests, recent studies have shown that climbing plants often remain poorly sampled in Brazil

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(Udulutsch et al. 2004). This under recording of climbing plant diversity results in a corresponding low number of herbarium specimens available for study and is accompanied by a corresponding paucity of taxonomic revisions of climbing plant families in Brazil (e.g. Cucurbitaceae, Dioscoreaceae and Menispermaceae).

The complete absence of published records of climbing plant diversity in northeastern Brazil explains the fact that the following abundant species are identified as new species records for this region: Aristolochia pappilaris Masters (1875: 100, Aristolochiaceae), Jacquemontia menispermoides Choisy (1838: 63, Convolvulaceae), Phryganocydia corymbosa (Vent.) Bureau ex Schumann (1894: 224, Bignoniaceae), Psiguria triphylla (Miq.) Jeffrey (1978: 353, Cucurbitaceae), and Sabicea grisea Chamisso & Schlechtendal (1829: 192, Rubiaceae).

At a global scale, latitude and seasonality in rainfall are often cited as explaining variations in the species composition of climbing plants floras in tropical forests (Gentry 1991, Campanello et al. 2007). According to Schnitzer & Bongers (2002), habitat diversity and history of human impact at a site are the most important factors determining climbing plant species richness. Alves-Araújo et al. (2008) published a checklist for Igarassu which includes plant habit. Based on this list the most species-rich fragment in Igarassu is Piedade with 49 species, followed by Macacos (41 species), Pezinho (34 species), Zambana (25 species), Santa Helena (7 species), and Vespas (4 species). Pezinho is among the most species-rich fragments yet is about a tenth of the size of Piedade. This result is possibly related to the diversity of habitats found in each fragment which in turn is a product of the history of human impact as suggested by Schnitzer & Bongers (2002) and Alves-Araújo et al. (2008) in addiction to the nearness of this little fragment to other big ones as Piedade and Macacos (Fig. 1B).

FIGURE 1. A. Map of northeastern Brazil, state of Pernambuco, Recife metropolitan region and town of Igarassu. B.Fragment disposition in the Usina São José plant’s territory. 1. Zambana. 2. Piedade. 3. Macacos. 4. Pezinho. 5. Vespas.6. Santa Helena.FIGURE 1. A. Map of northeastern Brazil, state of Pernambuco, Recife metropolitan region and town of Igarassu. B. Fragment disposition in the Usina São José plant’s territory. 1. Zambana. 2. Piedade. 3. Macacos. 4. Pezinho. 5. Vespas. 6. Santa Helena.

A

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Key to the identification of the climbing plants of Igarassu based on flowering materialNote that for species of the Sapindaceae flowering plant family, fruiting characters are included as these are essential for identification to species.

1. Tendrils present, but not associated with the inflorescence.......................................................................................... 21. Tendrils not present, or where present unbranched .................................................................................................... 342. Leaves opposite............................................................................................................................................................. 32. Leaves alternate ............................................................................................................................................................ 63. Leaves simple ................................................................................................................................ Strychnos parviflora3. Leaves compound. ........................................................................................................................................................ 44. Branches of the inflorescence flattened in cross-section, the calyx spathe-like… .............. Phryganocydia corymbosa4. Branches of the inflorescence circular in cross-section, the calyx campanulate .......................................................... 55. Glands of the calyx 5, discoid, present on the calyx outer surface (Fig. 2A); corolla yellow, anthers glabrous............

........................................................................................................................................... Adenocalymma hypostictum5. Glands absent from the outer surface of the calyx; corolla red to pink, anthers hairy .......................... Lundia cordata

B

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6. Leaves simple or 1-foliolate compound........................................................................................................................ 76. Leaves compound and with more than one leaflet ..................................................................................................... 267. Flowers zygomorphic ................................................................................................................................................... 87. Flowers actinomorphic ................................................................................................................................................. 98. Basal third of the leaflet connate; bracts reniform; petal outer surface completely hairy ............... Phanera outimouta8. Basal two-thirds of the leaflets connate; bracts linear-lanceolate; petals outer surface hairy only along the midvein ..

..................................................................................................................................................... Phanera trichosepala9. Tendrils opposite the leaves (Fig. 3A) ............................................................................................... Cissus verticillata9. Tendrils axillary to the leaves, either in the leaf sheath (appearing emergent from the petiole) or at the base of the

inflorescence (Fig. 3B, C, D, F, G) ............................................................................................................................. 1010. Tendrils 2, at the apex of the leaf sheath (appearing emergent from the petiole, Fig. 3G)................ Smilax syphilitica10. Tendril 1, in the leaf’s axils or borne at an acute angle to the petiole (Fig. 3D) ........................................................ 1111. Flowers unisexual ....................................................................................................................................................... 1211. Flowers bisexual ......................................................................................................................................................... 1712. Calyx urceolate (Fig. 2B); stamens 2.......................................................................................................................... 1312. Calyx infundibuliform or campanulate (Fig. 2C); stamens 3 ..................................................................................... 1513. Leaves 5-lobate; anthers straight ....................................................................................................... Psiguria umbrosa13. Leaves 1–3-lobate; anthers twice-folded from the base ............................................................................................. 1414. Lobes of the calyx longer than the lobes of the corolla ...................................................................Gurania acuminata14. Lobes of the calyx shorter than the lobes of the corolla ................................................................... Gurania spinulosa15. Flowers solitary............................................................................................................................Momordica charantia15. Flowers borne in clusters (racemes) ........................................................................................................................... 1616. Adaxial surface of the leaves with stiff recurved hairs; flowers grouped in lax, larger than 2cm long racemes, bearing

more than 3 flowers............................................................................................................................Cayaponia tayuya16. Leaves glabrous; flowers grouped in congested, up to 2cm, racemes bearing less than 3 flowers (Fig. 2D) ................

.....................................................................................................................................................Melothria fluminensis17. Ovary inferior .......................................................................................................................................Gouania virgata17. Ovary superior ............................................................................................................................................................ 1818. Plant hairy; petals absent ................................................................................................................ Passiflora suberosa18. Plant glabrous; petals 5 ............................................................................................................................................... 1919. Leaves entire ............................................................................................................................................................... 2019. Leaves lobate .............................................................................................................................................................. 22 20. Flowers grouped in racemes; styles 4 .................................................................................................. Passiflora ovalis20. Flowers solitary or in pairs; styles 3 ........................................................................................................................... 2121. Stem 4-angulate; petiole with 2–6 sessile glands ................................................................................. Passiflora alata21. Stem cylindrical; petiole with 2 stalked glands ............................................................................... Passiflora galbana22. Stems 2-angulate; leaves bearing 8–12 circular glands on the adaxial surface ..................................Passiflora misera22. Stems not angulate; leaves eglandular ........................................................................................................................ 2323. Leaves 5-lobate .............................................................................................................................Passiflora cincinnata23. Leaves 3-lobate ........................................................................................................................................................... 2424. Petiole with 2 sessile glands ................................................................................................................ Passiflora edulis24. Petiole with 2–6 stalked glands .................................................................................................................................. 2525. Apex of the leaves and stipules mucronate; bracts lanceolate, green… ..................................... Passiflora watsoniana25. Apex of the leaves and stipules truncate; bracts obovate, pink .....................................................Passiflora edmundoi26. Tendrils opposite to the leaves (Fig. 3A) ....................................................................................................Cissus erosa26. Tendrils in the axils of the leaves or at the base of the inflorescence (Fig. 3B, C, D, F, G)....................................... 2727. Flowers actinomorphic ............................................................................................................................................... 2827. Flowers zygomorphic ................................................................................................................................................. 2928. Calyx narrow in its length, geen, glabrous ........................................................................................ Psiguria triphylla28. Calyx wide at basis with narrowed entering, orange, hairy..........................................................Gurania bignoniacea29. Fruit divided into three samaras.................................................................................................................................. 3029. Fruit a simple capsule ................................................................................................................................................. 3130. Sepals with velvety margin (Fig. 2G) .......................................................................................Serjania subimpunctata30. Sepals entirely glabrescent or glabrous (Fig. 2H)........................................................................ Serjania salzmaniana31. Fruit without wings .............................................................................................................................Paullinia pinnata31. Fruit with wings .......................................................................................................................................................... 3232. Fruit with one wing...................................................................................................................Machaerium aculeatum32. Fruit with three wings ................................................................................................................................................. 3333. Leaf margin entire; wings of the fruit obovate, apiculate (Fig. 2E) ................................................Paullinia racemosa

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FIGURE 2. A Adenocalymma hypostictum, discoid glands of the calyx. B. Gurania bignoniacea, urceolate calyx. C. Cayaponia tayuya, campanulate calyx. D. Melothria fluminensis, short laxiflorous raceme. E. Paullinia racemosa, fruit with oblong and apiculate wings. F. Paullinia trigonia, fruit with elliptical wings and absent apicule. G. Serjania subim-punctata, fruit with oblong wings. H. Serjania salzmanianna, fruit with ovate wings. I. Stigmaphyllon salzmanii, fertile branch with fruit showing the wing expansion facing the base. J. Heteropterys nordestina, fertile branch with fruit show-ing the wing expansion facing the apex. K–M. Leaves and detail of the colleters, K. Blepharodon nitidum, L. Mandevilla moricandiana, M. Matelea maritima. N. Coccoloba ochreolata, ochrea.

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33. Leaf margin dentate; wings of the fruit elliptical, not apiculate (Fig. 2F)......................................... Paullinia trigonia34. Leaves opposite or verticillate .................................................................................................................................... 3534. Leaves alternate .......................................................................................................................................................... 5035. Flowers grouped into a capitate compound inflorescence...................................................................Mikania obovata35. Flowers grouped but not into a capitate compound inflorescences ............................................................................ 3636. Nectariferous disk conspicuous, surrounding the ovary............................................................................................. 3736. Nectariferous disk inconspicuous or absent............................................................................................................... 38 37. Plant with hairs that are rough to the touch ..............................................................................Prionostemma asperum37. Plant without hairs or with hairs that are not rough to the touch................................................Hippocratea vollubilis38. Petiole glandular, glands 2, discoid ............................................................................................................................ 3938. Petiole glandular or eglandular, where glandular glands not discoid ......................................................................... 4139. Glands 2 at the base of the petiole (Fig. 2J)............................................................................. Heteropterys nordestina39. Glands 2 at the apex of the petiole (Fig. 2I) ............................................................................................................... 4040. Surface abaxial of the leave hirsute ........................................................................................ Stigmaphyllon blanchetii40. Surface abaxial of the leave glabrous ......................................................................................Stigmaphyllon salzmanii41. Stipules borne between the petioles........................................................................................................ Sabicea grisea41. Stipules not borne between the petioles...................................................................................................................... 4242. Flowers actinomorphic, the number of stamens more than double the number of petals ......................................... 43 42. Flowers zygomorphic, the number of stamens double, equal, or less than number of petals .................................... 4843. Petiole twining; latex absent; stigma feather-like..................................................................................Clematis dioica43. Petiole not twining; latex present; stigma not plumose .............................................................................................. 4444. Plant glabrous ............................................................................................................................................................. 4544. Plants hairy ................................................................................................................................................................. 4645. Leaves oblong-elliptic; glands 2, broader at the base (Fig. 2K) ................................................... Blepharodon nitidum45. Leaves obovate; glands 2,not broader at the base (Fig. 2L) ................................................. Mandevilla moricandiana46. Glands absent from the leaf lamina .............................................................................................Temnadenia odorifera46. Glands 2–6, borne on the leaf lamina ......................................................................................................................... 4747. Glands 2–6, borne along the midvein ...............................................................................................Mandevilla scabra47. Glands 2–6, borne at the base of the midvein (Fig. 2M) ................................................................... Matelea maritima48. Corolla with wholly distinct petals; stamens 10 .......................................................................Niedenzuella acutifolia48. Corolla with wholly or partially fused petals; stamen 4 ............................................................................................ 4949. Plant herbaceous; petiole winged; flowers orange ............................................................................. Thunbergia alata49. Plant woody; petiole not winged; flowers white (Fig. 4I and J)…......................................... Mendoncia blanchetiana50. Ochrea present (Fig. 2N) .............................................................................................................Coccoloba ochreolata50. Ochrea absent.............................................................................................................................................................. 5151. Leaves compound ....................................................................................................................................................... 5251. Leaves simple or compound unifoliolate.................................................................................................................... 7152. Palms. Terminal leaflets reduced, stiff and hook-like (Fig. 4A)................................................................................. 5352. Not palms. Terminal leaflets not hook-like, foliaceus ................................................................................................ 5453. Raphe unarmed, leaflets unarmed or sometimes armed with thorns at the base of the leaflets; apex of bracts covered

in narrow thorns (Fig. 4B) ..................................................................................................... Desmoncus orthacanthos53. Raphe and abaxial surface of the leaflets armed with stiff and twisted thorns; apex of bracts covered in pustular

thorns (Fig. 4C) ...................................................................................................................... Desmoncus polyacanthos54. Inflorescences opposite to the leaves..........................................................................................................Cissus erosa54. Inflorescences never opposite to the leaves................................................................................................................ 5555. Leaves digitate .............................................................................................................................Merremia macrocalyx55. Leaves with 3 or more leaflets, never digitate ............................................................................................................ 5656. Prickles or thorns present................................................................................................................Senegalia tenuifolia56. Prickles or thorns absent ............................................................................................................................................. 5757. Calyx 2-lobate/divided................................................................................................................................................ 5857. Calyx 4 or 5 lobed....................................................................................................................................................... 6058. Surface abaxial of the leave glabrous ........................................................................................... Canavalia parviflora58. Surface abaxial of the leave glabrescent to pubescent................................................................................................ 5959. Calyx lobes of differing lengths..................................................................................................Canavalia braziliensis59. Calyx lobes of uniform length ......................................................................................................... Canavalia dictyota60. Calyx 5 lobed .............................................................................................................................................................. 6160. Calyx 4 lobed ............................................................................................................................................................. 68 61. Calyx including a spur ................................................................................................................................................ 6261. Calyx lacking a spur ................................................................................................................................................... 63

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FIGURE 3. Tendril disposition. A. Cissus erosa, opposite to the leaves. B. Gouania virgata, C. Passiflora alata,axillary. D. Psiguria umbrosa, in acute angle with the petiole E-F. Serjania subinpunctata, E. In the inflorescence base, F. detail G. Smilax syphilitca, on the apex of the sheath.

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FIGURE 4. A. Desmoncus polyacanthos, leaf. B–C. Bract and inflorescence, B. D. orthacanthos, C. D. polyacanthos. D. Mucuna pruriens, part of the inflorescence. E–G. Dalechampia species, bracts, E. D. convolvuloides, F. D. Brazilien-sis, G. D. pernambucensis. H. Davilla aspera, heteromorphic sepals, internal 2, bigger than the external 3. I–J. Mendon-cia blanchetiana, I. Branch with fruits surrounded by bracts and persistent stigma J. Detail of bract.

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62. Stem cylindrical in cross-section; calyx lobes unequal in size, the larger lobe twice the length of the calyx tube, flowers lilac................................................................................................................................ Centrosema pubescens

62. Stem angulate in cross-section; calyx lobes equal in size, the lobes not twice the length of the calyx tube; flowers yellow........................................................................................................................................... Centrosema plumieri

63. Bracteole caducous; calyx campanulate ..................................................................................................................... 6463. Bracteole persistent; calyx tubular.............................................................................................................................. 6564. Stems bearing lenticels, glabrous; leaflets lanceolate.................................................................... Connarus blanchetii64. Stem lacking lenticels, hairy; leaflets ovate............................................................................... Rynchosia pyramidalis65. Stigma straight ....................................................................................................................................... Clitoria falcata65. Stigma twisted............................................................................................................................................................. 6666. Corolla wings conspicually larger than the standard ............................................................ Macropitilium prostratum66. Corolla wings never larger than the standard ............................................................................................................. 6767. Plants glabrous; bracteoles stiff, adpressed to the rachis ...........................................................................Vigna luteola67. Plants hirsute; bracteoles membranaceous, free .....................................................................................Vigna vexillata68. Lobes of the calyx of differingt lengths (Fig. 4D) .............................................................................. Mucuna pruriens68. Lobes of the calyx of uniform length.......................................................................................................................... 6969. Plants herbaceous.....................................................................................................................Centrosema brazilianum69. Plants woody............................................................................................................................................................... 7070. Stem hairy; calyx glabrous .................................................................................................................... Dioclea virgata70. Stem glabrous; calyx hairy ..................................................................................................................Dioclea violacea71. Flowers grouped in a flower-like inflorescence ......................................................................................................... 7271. Flowers borne in racemes, panicles, glomerules or solitary ....................................................................................... 7572. Flowers arranged into a capitate head.................................................................................... Cyrtocymura scorpioides72. Inflroescence protected by two bright involucral bracts, flowers not arranged into a capitulum.73. Involucral bracts entire (Fig 4E) .......................................................................................Dalechampia convolvuloides73. Involucral bracts 3-lobate ........................................................................................................................................... 7474. Stipules linear-subulate; seeds rough, dark-brown (Fig 4F) .................................................. Dalechampia braziliensis74. Stipules reniform; seeds smooth, light-brown (Fig 4F) ..................................................Dalechampia pernambucensis75. Flowers unisexual ....................................................................................................................................................... 7675. Flowers bisexual ......................................................................................................................................................... 8076. Hairs urticating; flowers of both sexes borne in the same inflorescence.................................................................... 7776. Hairs not urticating, when present; male and female flowers borne in separate inflorescences ................................ 7877. Leaves 5-veined ....................................................................................................................................Tragia volubilis 77. Leaves 3-veined ................................................................................................................................Tragia lessertiana 78. Plants dioecious; nectariferous disk present; ovary inferior ........................................................Dioscorea marginata 78. Plants monoecious; nectariferous disk absent; ovary superior ................................................................................... 7979. Leaves peltate; bracts cordate; stamens fused at their base, sessile........................................Cissampelos glaberrima 79. Leaves not peltate; bracts filiform; stamens fused at their base, stalked .............................Cissampelos andromorpha80. Flowers zygomorphic ................................................................................................................................................. 8180. Flowers actninomorphic ............................................................................................................................................. 8581. Perianth composed of distinct sepals and petals ......................................................................................................... 8281. Perianth composed of tepals ....................................................................................................................................... 8482. Papilionoid corolla .....................................................................................................................Securidaca diversifolia82. Corolla not papilionoid ............................................................................................................................................... 8383. Basal third of the leaflets connate; bracts reniform; petals hairy over their entire surface ............. Phanera outimouta83. Apical third of the leaflets connate; bracts linear-lanceolate; petals hairy along the midvein only ...............................

..................................................................................................................................................... Phanera trichosepala84. Pseudo-stipules reniform; lip (expanded part of corolla tube) smooth....................................Aristolochia braziliensis84. Pseudo-stipules absent; lip (expanded part of corolla tube) papillose........................................Aristolochia papillaris85. Flowers with completely separate petals .................................................................................................................... 8685. Flowers with the petals fused...................................................................................................................................... 8986. Flowers borne in axillary or cauliflorous glomerules.................................................................. Doliocarpus dentatus86. Flowers borne in panicles or cymes............................................................................................................................ 8787. Plant glabrous; inflorescences borne opposite to the leaves............................................................. Cissus verticillada87. Plant with rough hairs; inflorescences terminal.......................................................................................................... 8888. Sepals unequal, inner two larger than the outer three, surrounding the flowers (Fig. 4H)..................... Davilla aspera88. Sepals equal, not surrounding the flowers ......................................................................................Tetracera breyniana89. Flowers solitary..............................................................................................................................Operculina altissima89. Flowers grouped in inforescences............................................................................................................................... 90

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90. Stigma globose............................................................................................................................................................ 9190. Stigma oblong ............................................................................................................................................................. 9691. Leaves pinnatifid.............................................................................................................................. Ipomoea quamoclit91. Leaves entire to 3–5 lobed .......................................................................................................................................... 9292. Leaves sagitate; sepals with liguliform appendix on their inner surface.......................................... Ipomoea bahiensis92. Leaves never sagitate; sepals without an appendix on their inner surface ................................................................. 9393. Leaves 3–5-lobed; sepals corniculate ............................................................................................ Ipomoea hederifolia93. Leaves entire; sepals not corniculate .......................................................................................................................... 9494. Leaves and sepals glabrous................................................................................................................. Ipomoea obscura94. Leaves and sepals hairy to velvety ............................................................................................................................. 9595. Leaves elliptic; hairs golden ..........................................................................................................Bonamia maripoides95. Leaves cordate; hairs white.................................................................................................................... Ipomoea martii96. Plants velvety, hairs stellate ...........................................................................................Jacquemontia menispermoides96. Plants glabrous to glabrescent, where glabrescent the hairs simple ............................................... Merremia umbelata

Key to the identification of the climbing plants of Igarassu based on fruiting material1. Tendrils present, but not associated with the inflorescence.......................................................................................... 21. Tendrils absent, or where present just a simple branch .............................................................................................. 342. Leaves opposite............................................................................................................................................................. 32. Leaves alternate ............................................................................................................................................................ 63. Leaves simple ................................................................................................................................ Strychnos parviflora3. Leaves compound ......................................................................................................................................................... 44. Leaflets bearing two dark glands at their base, branches of the inflorescence flattened in cross-section......................

............................................................................................................................................. Phryganocydia corymbosa4. Leaflets without dark glands, branches of the inflorescence cylindrical in cross-section............................................ 55. Fruits oblongoid, seeds not winged ................................................................................... Adenocalymma hypostictum5. Frutis elongated, seeds winged .............................................................................................................. Lundia cordata6. Leaves simple or 1-foliolate compound........................................................................................................................ 76. Leaves compound and with more than one foliole ..................................................................................................... 267. Tendrils hook-like ......................................................................................................................................................... 87. Tendrils twining ............................................................................................................................................................ 98. Basal third of the leaflet connate; fruit indehiscent ..................................................................... Phanera trichosepala8. Basal two-thirds of the leaflets connate; fruit dehiscent.................................................................. Phanera outimouta9. Tendrils axillary to the leaves ..................................................................................................................................... 109. Tendril not axillary to the leaves................................................................................................................................. 1910. Fruits winged, grouped in short (<20cm) racemes ...............................................................................Gouania virgata10. Fruits berries, solitary, in pairs or long (>20cm) racemes .......................................................................................... 1111. Leaves entire ............................................................................................................................................................... 1211. Leaves 2- or more lobbed ........................................................................................................................................... 1412. Fruits grouped in racemes up to 1.5 m in length ................................................................................. Passiflora ovalis12. Fruits solitary or in pairs............................................................................................................................................. 1313. Stem 4-angulate in cross-section; petiole with 2-6 sessile glands ........................................................ Passiflora alata13. Stem cylindrical in cross-section; petiole with 2 stalked glands ..................................................... Passiflora galbana14. Leaves 5-lobbed ............................................................................................................................Passiflora cincinnata14. Leaves 2- or 3-lobbed ................................................................................................................................................. 1515. Mature fruits never more than 15mm long ................................................................................................................. 1615. Mature fruits always more than 15mm long............................................................................................................... 1716. Stem hairy, flattened in cross-section .................................................................................................Passiflora misera16. Stem glabrous, cylindrical in cross-section .................................................................................... Passiflora suberosa17. Petiole with two sessile glands ............................................................................................................ Passiflora edulis17. Petiole with 2-6 stalked glands ................................................................................................................................... 1818. Apex of the leaves and stipules mucronate................................................................................. Passiflora watsoniana18. Apex of the leaves and stipules truncate........................................................................................Passiflora edmundoi19. Tendrils opposite to the leaves (Fig. 3A) ........................................................................................... Cissus verticillata19. Tendrils in the leaf sheath, at the basis of the inflorescence, or forming an acute angle with petiole (Fig. 3B, C, D, F,

G) ................................................................................................................................................................................ 2020. Tendrils two, in the apex of the leaf sheath (Fig. 3G) ....................................................................... Smilax syphilitica20. Tendril solitary, in the leaf’s axils, forming an acute angle with the petiole or at the basis of the inflorescence (Fig.

3D) .............................................................................................................................................................................. 21

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21. Leaves glabrous .......................................................................................................................................................... 2221. Leaves hairy................................................................................................................................................................ 2322. Latex watery; seeds arilate................................................................................................................. Psiguria umbrosa22. Latex absent; seeds without an aril ..............................................................................................Melothria fluminensis23. Mature fruits green or yellow striped ......................................................................................................................... 2423. Mature entirely yellow to orange................................................................................................................................ 2524. Petioles to 3 cm long; leaves scabrous on the adaxial surface.......................................................... Gurania spinulosa24. Petioles to 15 cm long; leaves pubescent on the adaxial surface..................................................... Guranis acuminata25. Mature fruit orange, dehiscent; seeds red ....................................................................................Momordica charantia25. Mature fruit yellow (rarely red), indehiscent; seeds white ................................................................Cayaponia tayuya26. Tendrils borne opposite to the leaves (Fig. 3A)..........................................................................................Cissus erosa26. Tendrils borne in the axils of the leaves or at the basis of the inflorescence (Fig. 3B, C, D, F, G)............................ 2727. Fruits fleshy ................................................................................................................................................................ 2827. Fruits dry..................................................................................................................................................................... 3228. Leaves with 2 or 3 foliolules....................................................................................................................................... 2928. Leaves with more than 3-foliolules ............................................................................................................................ 3029. Plants glabrous; fruits solitary ........................................................................................................... Psiguria triphylla29. Plants pubscent to scrabous; fruit borne in racemes .....................................................................Gurania bignoniacea30. Fruits without wings ...........................................................................................................................Paullinia pinnata30. Fruit 3-winged............................................................................................................................................................. 3131. Leaf margin entire; wings of the fruit obovate, apiculate................................................................Paullinia racemosa31. Leaf margin dentate; wings of the fruit elliptical, not apiculate ........................................................ Paullinia trigonia32. Stem flattened in cross-section; leaves pinnate ........................................................................Machaerium aculeatum32. Stem cylindrical in cross-section; leaves 3-foliolate or ternate .................................................................................. 3333. Fruit divided into three oblong-winged samaras .......................................................................Serjania subinpunctata33. Fruit divided into three ovate-winged samaras............................................................................ Serjania salzmaniana34. Leaves opposite or verticillate .................................................................................................................................... 3534. Leaves alternate .......................................................................................................................................................... 4835. Stipules borne between the petioles........................................................................................................ Sabicea grisea35. Stipules not borne between the petioles...................................................................................................................... 3636. Plants hairy ................................................................................................................................................................. 3736. Plants glabrous............................................................................................................................................................ 4237. Latex present............................................................................................................................................................... 3837. Latex absent ................................................................................................................................................................ 4038. Glands absent from the leaf limb.................................................................................................Temnadenia odorifera38. Glands present on the leaf limb .................................................................................................................................. 3939. Leaves scabrous, glands 2-6, distributed along the midvein ............................................................Mandevilla scabra39. Leaves not scabrous, glands 2-6 grouped at the base of the midvein................................................ Matelea maritima40. Leaf covered with long fine hairs; fruit divided into 3 samaras ........................................... Stigymaphyllon blanchetii40. Leaf covered with short thick hairs............................................................................................................................. 5041. Fruit a 3-valved capsule; seeds winged ....................................................................................Prionostemma asperum41. Fruits grouped into a capitate compound infrutescence ......................................................................Mikania obovata42. Fruit winged................................................................................................................................................................ 4342. Fruit not winged.......................................................................................................................................................... 4543. Wings less than three times the length of the fruit....................................................................Niendenzuela acutifolia43. Wings as long as or longer than three times the length of the fruit ............................................................................ 4444. Wings oblong-falcate (Fig. 2J) ................................................................................................ Heteropterys nordestina44. Wings asymmetrically obovate (Fig. 2I) .................................................................................Stigmaphyllon salzmanii45. Stigma feather-like.................................................................................................................................Clematis dioica45. Stigma not feather-like................................................................................................................................................ 4646. Fruit a 3-carpelar capsule; seeds winged ....................................................................................Hippocratea vollubilis46. Fruit a berry or a 2-carpelar capsule; seeds not winged ............................................................................................. 4747. Fruit a capsule, 2-carpelar................................................................................................................... Thunbergia alata47. Fruit a berry ............................................................................................................................ Mendoncia blanchetiana48. Ochrea present (Fig. 2N) .............................................................................................................Coccoloba ochreolata48. Ohcrea absent.............................................................................................................................................................. 4949. Leaves compound ....................................................................................................................................................... 5049. Leaves simple or compound unifoliolate.................................................................................................................... 6950. Palms. Terminal leaflets reduced, stiff and hook-like (Fig. 4A)................................................................................. 51

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50. Not palms. Terminal leaflets not hook-like, foliaceus ................................................................................................ 5251. Raphe unarmed, leaflets unarmed or sometimes armed with thorns at the basis of the leaflets; apex of bracts covered

in narrow thorns (Fig. 4B) ..................................................................................................... Desmoncus orthacanthos51. Raphe and abaxial surface of the leaflets armed with stiff and twisted thorns; apex of bracts covered in pustular

thorns (Fig. 4C) ...................................................................................................................... Desmoncus polyacanthos52. Inflorescences opposite to the leaves..........................................................................................................Cissus erosa52. Inflorescences never opposite to the leaves................................................................................................................ 5353. Leaves digitate .............................................................................................................................Merremia macrocalyx53. Leaves with 3 or more leaflets, never digitate ............................................................................................................ 5454. Prickles or thorns present................................................................................................................Senegalia tenuifolia54. Prickles or thorns absent ............................................................................................................................................. 5555. Calyx persistent, 2-lobed ............................................................................................................................................ 5655. Calyx persistent, 4 or 5 lobed ..................................................................................................................................... 5856. Abaxial surface of the leave glabrous........................................................................................... Canavalia parviflora56. Abaxial surface of the leave glabrescent to pubescent ............................................................................................... 5757. Calyx lobes of unequal lengths ...................................................................................................Canavalia braziliensis57. Calyx lobes of equal lengths ............................................................................................................ Canavalia dictyota58. Calyx 5 lobed .............................................................................................................................................................. 5858. Calyx 4-lobed.............................................................................................................................................................. 6659. Calyx including a spur ................................................................................................................................................ 6059. Calyx lacking a spur ................................................................................................................................................... 6160. Stem cylindrical; calyx lobes unequal in size, the larger lobe twice the length of the calyx tube..................................

................................................................................................................................................... Centrosema pubescens60. Stem angulate; calyx lobes of equal size ...................................................................................... Centrosema plumieri61. Bracteole caducous; calyx campanulate ..................................................................................................................... 6261. Bracteole persistent; calyx tubular.............................................................................................................................. 6362. Stems bearing lenticels, glabrous; leaflets lanceolate; calyx persistent in fruit............................. Connarus blanchetii62. Stem lacking lenticels, hairy; leaflets ovate; calyx not persistent ............................................. Rynchosia pyramidalis63. Seeds more than ten per fruit ...................................................................................................Macroptilum prostratum63. Seeds 4-6 per fruit....................................................................................................................................................... 6464. Legume obtusely quadrangular; seeds spherical ................................................................................... Clitoria falcata64. Legume oblong to linear ............................................................................................................................................. 6565. Plants glabrous; bracteoles stiff, adpressed to the rachis ...........................................................................Vigna luteola65. Plants hirsute; bracteoles membranaceous, free .....................................................................................Vigna vexillata66. Lobes of the calyx of unequal length (Fig. 4D); fruit with urticating hairs….................................... Mucuna pruriens66. Lobes of the calyx of equal length; fruit without urticating hairs .............................................................................. 6767. Plants herbaceous.....................................................................................................................Centrosema brazilianum67. Plants woody............................................................................................................................................................... 6868. Stem hairy; calyx glabrous .................................................................................................................... Dioclea virgata68. Stem glabrous; calyx hairy ..................................................................................................................Dioclea violacea69. Fruits grouped in a flower-like inflorescence ............................................................................................................. 7069. Fruits borne in racemes, panicles, glomerules or solitary........................................................................................... 7370. Fruits arranged into a capitate compound inflorescence ....................................................... Cyrtocymura scorpioides70. Fruits protected by two persistent, bright involucral bracts, not arranged into a capitulum ...................................... 7171. Involucral bracts entire (Fig 4E) .......................................................................................Dalechampia convolvuloides71. Involucral bracts 3-lobed ............................................................................................................................................ 7272. Stipules linear-subulate; seed rough, dark-brown (Fig 4F) ................................................... Dalechampia brasiliensis72. Stipules reniform; seeds smooth, light-brown (Fig 4F) ..................................................Dalechampia pernambucensis73. Plant bearing urticating hairs; fruit a 3-lobed capsule; a single seed per locule......................................................... 7473. Plant not bearing urticating hairs; fruit 3-lobed or not, where 3-lobed more than one seed per locule ..................... 7574. Leaves 5-veined .....................................................................................................................................Tragia volubilis74. Leaves 3-veined .................................................................................................................................Tragia lessertiana75. Fruit solitary................................................................................................................................................................ 7675. Fruit borne in racemes, panicles or glomerule............................................................................................................ 7776. Pseudo-stipules reniform .........................................................................................................Aristolochia brasiliensis76. Pesudo-stipules absent ..............................................................................................................Aristolochia pappillaris77. Fruit a pod................................................................................................................................................................... 7877. Fruit not a pod............................................................................................................................................................. 7978. Apical third of the leaflets connate; bracts linear-lanceolate....................................................... Phanera trichosepala

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78. Basal third of the leaflets connate; bracts reniform ......................................................................... Phanera outimouta79. Fruits winged .............................................................................................................................................................. 8079. Fruits not winged ........................................................................................................................................................ 8180. Fruits 3-winged; seeds numerous, flattened .................................................................................Dioscorea marginata80. Fruits 1-winged; one seed per fruit, spherical............................................................................Securidaca diversifolia81. Fruits fleshy ................................................................................................................................................................ 8281. Fruits dry..................................................................................................................................................................... 8482. Fruits born in a cime, opposite to the leaves....................................................................................... Cissus verticilata82. Fruits born in caulinar or terminal racemes ................................................................................................................ 8383. Leaves peltate; bracts cordate ..................................................................................................Cissampelos glaberrima83. Leaves no peltate; bracts filiform ........................................................................................Cissampelos andromorpha84. Stems not releasing latex when cut............................................................................................................................. 8584. Stems releasing white or colourless latex when cut ................................................................................................... 8685. Sepals persistent, unequal, internal 2 bigger than the external 3, surrounding the fruit (Fig. 4H); seeds with a white

aril ............................................................................................................................................................ Davila aspera85. Sepals persistent, equal in size, not surrounding the fruit; seeds without a white aril ............................................... 8786. Fruits solitary .................................................................................................................................Operculina altissima86. Fruits grouped in infrutescences ................................................................................................................................. 8887. Fruits born in terminal racemes ......................................................................................................Tetracera breyniana87. Fruits born in axillary or cauliflorous glomerules ....................................................................... Doliocarpus dentatus88. Leaves pinatifid................................................................................................................................ Ipomoea quamoclit88. Leaves entire to 5-lobed.............................................................................................................................................. 8989. Leaves sagitate; inner surface of sepals with liguliform appendix ................................................... Ipomoea bahiensis89. Leaves never sagitated; inner surface of sepals without appendix ............................................................................. 9090. Leaves 3–5-lobed; sepals ............................................................................................................... Ipomoea hederifolia90. Leaves entire; sepals not corniculate .......................................................................................................................... 9191. Leaves and sepals glabrous......................................................................................................................................... 9291. Leaves and sepals hairy .............................................................................................................................................. 9392. Fruits irregulary dehiscent; seeds triangular ................................................................................... Merremia umbelata92. Fruits regulary dehiscent; seeds with two surfaces flat, one convex .................................................. Ipomoea obscura93. Fruits 4-valved, each valve dividing into two................................................................Jacquemontia menispermoides93. Fruits 4-valved, valves not subdividing upon opening ............................................................................................... 9494. Leaves elliptic; hairs golden ..........................................................................................................Bonamia maripoides94. Leaves cordate; hairs white.................................................................................................................... Ipomoea martii

ACANTHACEAE1. Mendoncia blanchetiana ProficeBrazil (Pernambuco, Bahia). Humid areas in the Atlantic forest. Previously known only by the type specimen collected in the state of Bahia (Profice 1997).Selected Voucher:—Zambana, Araújo et al. 570 (UFP). 2. Thunbergia alata Bojer ex SimsTropical Africa, United States, Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Brazil (widely distributed ), Ecuador, Peru, Bolivia, Paraguay, Argentina. Selected Vouchers:—Piedade, Araújo et al. 468 (IPA, UFP).

APOCYNACEAE3. Blepharodon nitidum (Vell.) Macbr. (Fig. 1K)Venezuela, Brazil (widely distributed) (Pereira, 2005).Selected Voucher:—Pezinho, Alves-Araújo et al. 265 (IPA).4. Mandevilla moricandiana (A.DC.) Woodson (Fig. 1L)Brazil (Rio Grande do Norte, Paraíba, Pernambuco, Bahia).Selected Voucher:—Zambana, Alves-Araújo et al. 553 (UFP).5. Mandevilla scabra (Hoffmanns. ex Roem. & Schult.) K.Schum. in Engler & Prantl.Colombia, Venezuela, Guyana, Suriname, Brazil (widely distributed ), Ecuador, Peru, Bolivia. Selected Voucher:—Pezinho, Araújo et al. 610 (UFP); Piedade, Alves-Araújo et al. 192 (UFP); Macacos, Araújo et al. 428 (UFP).6. Matelea maritima (Jacquin 1760:17.) Woodson (1941:222) (Fig. 1L)

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Panama, Colombia, Venezuela, Guyanas, Brazil (from Pará to Espírito Santo). Restinga and Atlantic forest slopes.Selected Voucher:—Piedade, Araújo et al. 405 (UFP); Macacos, Alves Araújo et al. 601 (UFP), Araújo et al. 297 (UFP). 7. Temnadenia odorifera (Vell.) J.F.MoralesBrazil (from Ceará to Rio Grande do Sul). Restinga and slope Atlantic forest (Kinoshita 2005).Selected Vouchers:—Pezinho, Alves-Araújo & Marques 228 (IPA, UFP, ULM).

ARECACEAE8. Desmoncus orthacanthos Mart. (Fig. 4A, B)Mexico, Central America, Caribbean, Colombia, Venezuela, Guyana, Suriname, Brazil (Pernambuco, Espírito Santo, Rio de Janeiro), Ecuador, Peru, Bolivia. Atlantic forest, in disturbed areas, capoeiras and riverside (Henderson et al. 1995).Selected Voucher:—Piedade, Araújo et al. 180 (UFP). 9. Desmoncus polyacanthos Mart. (Fig. 4A, C)Colombia, Venezuela, Ecuador, Guyanas, Peru, Bolivia, Brazil (widely distributed). Atlantic forest (Henderson et al.1995). Selected Vouchers:—Piedade, Araújo et al. 392 (IPA, UFP).

ARISTOLOCHIACEAE10. Aristolochia brasiliensis Mart. & Zucc. Brazil. Only known in the state of Pernambuco.Selected Voucher:—Piedade, Melquíades 199 (PEUFR).11. Aristolochia papillaris Masters in Mart.Brazil (Pernambuco, Bahia, Espírito Santo, Rio de Janeiro, São Paulo, Santa Catarina).Selected Voucher:—Pezinho, Araújo et. al 602 (UFP); Macacos, Araújo & Alves-Araújo 503 (UFP)..

ASTERACEAE12. Cyrtocymura scorpioides (Lam.) H.Rob. (syn.: Vernonia scorpioides (Lam.) Pers.)Mexico, Central America, Caribbean, Venezuela, Brazil (Pernambuco, Bahia), Peru, Argentina. Selected Vouchers:—Macacos, Araújo et al. 292 (UFP, ULM); Zambana, Araújo et al. 340 (UFP).13. Mikania obovata DC. Brazil (Pernambuco, Bahia, Rio de Janeiro).Selected Voucher:—Zambana, 14 March 2007, Alves-Araújo et al. 887 (UFP).BIGNONIACEAE14. Adenocalymma hypostictum Bureau & K.Schum. in Mart. (Fig. 2A)Brazil (Pernambuco, Bahia, Espírito Santo, Rio de Janeiro).Selected Vouchers:—Piedade, Araújo et al. 380 (UFP, ULM).15. Lundia cordata (Vell. ) A.DC.Brazil (Ceará, Pernambuco, Alagoas, Bahia, Goiás, Espírito Santo), PeruSelected Voucher:—Zambana, 17 July 2007, Araújo et al. 329 (IPA); Macacos, Alves-Araújo et al. 276 (UFP), Araújo et al. 482 (UFP).16. Phryganocydia corymbosa (Vent.) Bureau ex K.Schum. in Engler & PrantlCentral America, Venezuela, Ecuador, Peru, Brazil (widely distributed). Semi-deciduous to perennial Atlantic forest (Gentry 1997).Selected Vouchers:—Piedade, Melo et al. 45 (IPA, UFP); Macacos, Albuquerque et al. 297 (IPA).

CELASTRACEAE17. Hippocratea volubilis L.United States, Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Guyanas, Bolivia, Brazil (widely distributed), Paraguay, Argentina. Atlantic forest and forest-cerrado(savannah-like vegetation) transition (Lombardi & Lara 2003).Selected Voucher:—Piedade, Araújo et al. 384 (UFP); Macacos, Araújo & Alves-Araújo 498 (UFP).18. Prionostemma asperum (Lam.) MiersCaribbean, Venezuela, Colombia, Guyanas, Bolivia, Brazil (Roraima, Amazonas, Acre, Pará, Pernambuco). Atlantic forest semi-deciduous to perennial, capoeiras, gallery Forest, lowlands (Mennega & Hedin, 1999).Selected Voucher:—Macacos, Araújo et al. 413 (UFP); Pezinho, Alves-Araújo et al. 301 (UFP).

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CONARACEAE19. Connarus blanchetii Planch.Brazil (Paraíba, Pernambuco, Bahia) (Forero 1983).Selected Voucher:—Piedade, Araújo et al. 585 (UFP).

CONVOLVULACEAE20. Bonamia maripoides Hallier f.Panama, Venezuela, Suriname, Brazil (Pernambuco, Bahia).Selected Voucher:—Zambana, Albuquerque 493 (IPA, ULM). 21. Ipomoea bahiensis Willd. ex Roem. & Schult.Brazil (widely distributed ) (Simão-Biachini 1993).Selected Voucher:—Pezinho, Alves-Araújo et al. 229 (UFP); Zambana, Alves-Araújo et al. 651 (UFP).22. Ipomoea hederifolia L. Tropical America (widely distributed ) (Simão-Biachini 1993). Selected Voucher:—Zambana, Albuquerque 542 (IPA, ULM); Pezinho, Alves-Araújo et al. 603 (UFP). 23. Ipomoea martii Meisner in Mart. Brazil (Pernambuco, Goiás, São Paulo).Selected Voucher:—Piedade, Melo et al. 296 (UFP).24. Ipomoea obscura (L.) Ker Gawl.Brazil (Pernambuco). Native from Tropical Africa, being the first register to Brazil’s northeast.Selected Voucher:—Piedade, Alves-Araújo et al. 565 (UFP).25. Ipomoea quamoclit L.Tropical America (widely distributed ) (Simão-Biachini 1993).Selected Vouchers:—Macacos, Alves-Araújo et al. 531 (IPA, UFP).26. Jacquemontia menispermoides ChoisyBrazil (Pernambuco). Atlantic forest and Caatinga.Selected Voucher:—Zambana, Alves-Araújo et al. 643 (UFP); Piedade, Albuquerque 356 (IPA, ULM).27. Merremia macrocalyx (Ruiz & Pav.) O’DonellVenezuela, Ecuador, Bolivia, Brazil (widely distributed ).Selected Vouchers:—Zambana, Albuquerque 525 (IPA, ULM); Piedade, Alves-Araújo & Araújo 723 (UFP).28. Merremia umbellata (L.) Hallier f.United States, Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Bolivia, Brazil (Amapá, Rondônia, Amazonas, Pará, Ceará, Pernambuco), Argentina.Selected Voucher:—Zambana, Alves-Araújo et al. 665 (UFP); Piedade, Alves-Araújo 561 (UFP).29. Operculina altissima (Mart. ex Choisy) Meisner in Mart.Brazil (Pernambuco).Selected Voucher:—Piedade, 21 November 2003, Bezerra 111 (UFP).

CUCURBITACEAE30. Cayaponia tayuya (Vell.) Cogn. (Fig. 1C)Ecuador, Bolivia, Brazil (widely distributed). Pluvial and dry Atlantic forest, restinga and caatinga (Klein 2000). Selected Voucher—Santa Helena, Sá e Silva 176 (UFP); Pezinho, Alves-Araújo 576 (UFP).31. Gurania acuminata Cogn. (Fig. B) Colombia, Venezuela, Guyanas, Ecuador, Peru, Bolivia, Brazil (Acre, Amazonas, Rondônia, Pernambuco, Bahia) (Kearns 1998).Selected Voucher—Macacos, Melo et al. 61 (UFP).32. Gurania bignoniacea (Poepp. & Endl.) Jeffrey (Fig. 2B)Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil (Amazonas, Pará, Pernambuco, Alagoas, Bahia) (Kearns 1998).Selected Voucher—Piedade, Araújo et al. 381 (UFP).33. Gurania spinulosa (Poepp. & Endl.) Cogn.Venezuela, Ecuador, Bolivia, Brazil (Acre, Rondônia, Pernambuco) (Kearns 1998).Selected Voucher:—Piedade, Araújo et al. 402 (UFP).34. Melothria fluminensis Gardner (Fig. 2D)Honduras, Peru, Bolivia, Brazil (Pernambuco, Bahia, Rio de Janeiro).

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Selected Voucher:—Vespas, 18 June 2007, Ojima 62 (UFP).35. Momordica charantia L. Native to Africa and widely distributed throughout the tropics (Kearns 1998).Selected Vouchers:—Pezinho, Marques et al. 40; Piedade, Albuquerque & Rocha 410 (IPA, ULM).36. Psiguria triphylla (Miq.) JeffreyMexico, Central America, Caribbean, Venezuela, Ecuador, Guyanas, Peru, Brazil (Amazonas) (Kearns 1998). This is the first record in Brazil outside Amazonas. Selected Voucher:—Pezinho, Araújo et al. 446 (UFP); Piedade, Araújo et al. 382 (UFP).37. Psiguria umbrosa (Kunth) Jeffrey (Fig. 1D)Caribbean, Venezuela, Colombia, Brazil (Amazonas, Pará, Pernambuco) (Kearns, 1998).Selected Voucher:—Piedade, Araújo et al. 400 (UFP).

DILLENIACEAE38. Davilla aspera (Aubl.) Benoist (Fig. 4H)Mexico, Central America, Colombia Venezuela, Guyana, Peru Bolivia, Brazil (Pernambuco) (Aymard 1998).Selected Vouchesr:—Pezinho, 11 Araújo et al. 599 (IPA, UFP); Macacos, Araújo et al. 328 (UFP).39. Doliocarpus dentatus (Aubl.) Standl. Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Guyanas, Bolivia, Paraguay, Brazil (from Amazonas to São Paulo) (Aymard 1998).Selected Vouchers:—Zambana, Araújo et al. 483 (IPA, UFP).40. Tetracera breyniana Schltdl.Brazil (Rio Grande do Norte, Ceará, Pernambuco, Bahia, Rio de Janeiro).Selected Vouchers:—Zambana, Albuquerque & Rocha 440 (IPA, ULM).

DIOSCOREACEAE41. Dioscorea marginata Griseb. in Mart.Brazil (Amazonas, Pernambuco, Distrito Federal, Mato-Grosso do Sul).Selected Voucher:—Zambana, Alves-Araújo et al. 625 (UFP), Albuquerque 588 (IPA).

EUPHORBIACEAE42. Dalechampia brasiliensis Lam. (Fig. 4F)Brazil (Pernambuco, Rio de Janeiro). Atlantic forest. (Govaerts et al. 2000).Selected Vouchers:—Piedade, 22 March 2007, Alves-Araújo et al. 201 (IPA, UFP).43. Dalechampia convolvuloides Lam. (Fig. 4E)Brazil (Pernambuco, Sergipe, Rio de Janeiro). Atlantic forest. (Govaerts et al. 2000).Selected Vouchers:—Piedade, Araújo et al. 379 (IPA, UFP); Macacos, Araújo et al. 294 (UFP); Zambana, Araújo et al. 347 (UFP).44. Dalechampia pernambucensis Baill. (Fig. 4G)Brazil (Ceará, Pernambuco) (Govaerts et al. 2000).Selected Voucher:—Pezinho, Alves-Araújo et al. 271 (UFP).45. Tragia lessertiana (Baill.) Müll.Arg.Guyanas, Brazil (Amapá, Maranhão, Pernambuco, Bahia) (Govaerts et al. 2000).Selected Vouchers:—Piedade, 10 January 2008, Alves-Araújo et al. 799 (UFP, IPA).46. Tragia volubilis L. Mexico, Central America, Caribbean, South America (from Ecuador to Argentina), Brazil (widely distributed ), Tropical Africa (Govaerts et al. 2000).Selected Voucher:—Pezinho, Araújo et al. 604 (UFP).

FABACEAE47. Canavalia brasiliensis Mart. ex Benth.United States, Mexico, Central America, Caribbean and South America (widely distributed ) (Aymard et al. 1999).Selected Voucher:—Piedade, Marques 244 (UFP).48. Canavalia dictyota PiperUnited States, Panama, Caribbean, Venezuela, Guyanas, Colombia, Ecuador, Peru, Brazil (Roraima, Amazonas, Pará,

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Ceará, Pernambuco, Bahia) (Aymard et al. 1999).Selected Voucher:—Pezinho, Araújo et al. 609 (UFP).49. Canavalia parviflora Benth. in Mart. Peru and Brazil (Acre, Pernambuco, Bahia, Goiás, Minas Gerais, Rio de Janeiro, São Paulo). Selected Voucher:—Zambana, Albuquerque et al. 495 (UFP). 50. Centrosema brasilianum (L.) Benth.Caribbean, Panama, Venezuela, Colombia, Peru, Guyanas, Bolivia, Brazil (widely distributed ). Savannah, edge of roads and forests boards (Aymard et al. 1999).Selected Vouchers:—Macacos, Marques 218 (IPA, ULM).51. Centrosema plumieri (Turpin ex Pers.) Benth.Mexico, Central America, Caribbean, Venezuela, Ecuador, Guyanas, Peru, Bolivia, Brazil (from Roraima to São Paulo) and introduced in Tropical Africa (Aymard et al. 1999).Selected Voucher:—Piedade, 19 December 2007, Araújo et al. 520 (UFP). 52. Centrosema pubescens Benth.United States, Mexico, Central America, Caribbean, Venezuela, Ecuador, Bolivia, Brazil (from Amazonas to Rio de Janeiro), Argentina.Selected Voucher:—Macacos, Araújo et al. 432 (UFP); Piedade, Melo et al. 136 (UFP). 53. Clitoria falcata Lam.Mexico, Central America, Caribbean, Venezuela, Ecuador, Guyanas, Peru, Brazil (Roraima, Pernambuco, Bahia, Rio de Janeiro), Paraguay (Aymard et al. 1999).Selected Vouchers:—Macacos, Araújo et al. 316 (IPA, UFP); Zambana, Albuquerque 511 (IPA).54. Dioclea violacea Mart. ex Benth.United States, Caribbean, Guyanas, Peru, Brazil (Roraima, Amapá, Pará, Bahia, Mato-Grosso, Mato Grosso do Sul, Espírito Santo, Rio de Janeiro). Selected Voucher:—Zambana, Araújo et al. 448 (UFP). 55. Dioclea virgata (Rich.) Amshoff Mexico, Central America, Caribbean, Venezuela, Colombia, Guyanas, Ecuador, Peru, Bolivia, Brazil (from Roraima to São Paulo), Argentina (Aymard et al. 1999).Selected Voucher:—Macacos, Araújo et al. 484; Zambana, Alves-Araújo et al 548 (UFP).56. Machaerium aculeatum RaddiVenezuela, Guianas, Peru, Bolivia, Brazil, Paraguay, (Paraíba, Pernambuco, Bahia, Goiás, Minas Gerais, Rio de Janeiro), Argentina. Selected Voucher:—Macacos, Marques et al. 214 (IPA). 57. Macroptilium prostratum (Benth.) Urb.Brazil (Pernambuco, Rio Grande do Sul), Paraguay, Argentina.Selected Voucher:—Macacos, Araújo et al. 438 (UFP).58. Mucuna pruriens (L.) DC. (Fig. 4D)United States, Mexico, Central America, Caribbean, Brazil (Pernambuco, Bahia) (Aymard et al. 1999).Selected Vouchers:—Piedade, Araújo et al. 377 (IPA, UFP).59. Phanera outimouta (Aubl.) L.P.QueirozCentral America to eastern South America, Brazil (Pernambuco, Bahia) (Queiroz, 2006).Selected Vouchers:—Piedade, Araújo & Alves-Araújo 513 (IPA, UFP).60. Phanera trichosepala L.P.QueirozBrazil (Pernambuco, Bahia). Humid Atlantic forest. (Queiroz 2006). This is the first collection apart from the type specimen. Selected Voucher:—Piedade, Albuquerque 597 (UFP). 61. Rhynchosia pyramidalis (Lam.) Urb. (= R. phaseoloides (Sw.) DC.)Caribbean, Panama, Venezuela, Colombia, Guyanas, Ecuador, Peru, Bolivia, Brazil (widely distributed ) (Aymard et al.1999).Selected Vouchers:—Macacos, Melo et al. 73 (UFP); Zambana, Araújo et al. 174 (UFP); Pezinho, Araújo et al. 280 (UFP).62. Senegalia tenuifolia (L.) Britton & Rose (= Acacia tenuifolia (L.) Willd.)Venezuela, Caribbean, Peru, Guyanas, Bolivia, Brazil (Roraima, Amazonas, Rondônia, Pará, from Piauí to São Paulo).Selected Voucher:—Pezinho, Araújo et al. 603 (UFP).

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63. Vigna luteola (Jacq.) Benth.United States, Mexico, Central America, Caribbean, South America (from Venezuela to Uruguay), Brazil (Roraima, Acre, Pará, Pernambuco, Bahia) (Aymard et al. 1999).Selected Vouchers:—Piedade, Marques et al. 247 (IPA, ULM); Pezinho, Alves-Araújo et al. 567 (UFP). 64. Vigna vexillata (L.) A.Rich.Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil (Amazonas, Pará, Pernambuco, Bahia) (Aymard et al. 1999).Selected Voucher:—Pezinho, Alves-Araújo et al. 581 (UFP).

LOGANIACEAE65. Strychnos parviflora Spruce ex Benth.Bolivia, Peru, Brazil (from Pará to São Paulo), Paraguay. Selected Voucher:—Piedade, Marques & Albuquerque 16 (IPA).

MALPIGHIACEAE66. Heteropterys nordestina Amorim (Fig. 2J)Brazil (from Rio Grande to Norte ao Espírito Santo) Restingas and humid forests (Amorim 2003).Selected Vouchers:—Piedade, Araújo et al. 389 (IPA, UFP, ULM), Alves-Araújo & Albuquerque 323 (IPA, UFP). 67. Niedenzuella acutifolia (Cav.) W.R.AndersonSouth America (from Guyanas to south Brazil) (Anderson 2006).Selected Voucher:—Piedade, 19, Araújo & Alves-Araújo 533-B (UFP).68. Stigmaphyllon blanchetii W.R.AndersonBrazil (from Paraíba to Espírito Santo) Atlantic forest. Selected Vouchers:—Zambana, Melo et al. 04 (IPA, UFP, ULM); Pezinho, Araújo et al. 447 (UFP)..69. Stigmaphyllon salzmannii Juss. (Fig. 2I)Brazil (from Pernambuco until Rio de Janeiro), Atlantic forest.Selected Vouchers:—Macacos, Melo et al. 59 (UFP, ULM), Alves-Araújo & Araújo 727 (UFP); Pezinho, Alves-Araújo et al. 242 (UFP)..

MENISPERMACEAE70. Cissampelos andromorpha DC.From Central America to southern Brazil (widely distributed ). Selected Vouchers:—Pezinho, Alves-Araújo et al. 275 (IPA, UFP); Macacos, Araújo ET al. 414 (UFP); Zambana, Alves-Araújo 895 (UFP). 71. Cissampelos glaberrima A.St.-Hil.Brazil (Maranhão, Ceará, Pernambuco, Goiás, Mato-Grosso, Minas Gerais, Rio de Janeiro, São Paulo).Selected Vouchers:—Pezinho, Alves-Araújo et al. 274 (IPA, UFP).

PASSIFLORACEAE72. Passiflora alata Curtis (Fig. 3C)Mexico, Central America, Caribbean, Ecuador, Peru, Brazil, (from Bahia to Rio Grande do Sul). Atlantic forest, seasonal forest and restinga (Nunes & Queiroz 2006).Selected Voucher:—Zambana, Alves-Araújo et al. 687 (UFP).73. Passiflora cincinnata MastersSouth America (widely distributed ) (Nunes & Queiroz 2006).Selected Voucher:—Zambana, Albuquerque 457 (IPA).74. Passiflora contracta VittaBrazil (Pernambuco, Bahia, Espírito Santo). Distributed along the litoral: Atlantic forest, seasonal Forest and restingas (Vitta & Bernaci 2004).Selected Vouchers:—Piedade, Araújo et al. 388 (UFP, ULM); Macacos, Araújo et al. 310 (UFP); Zambana, Araújo et al. 576 (UFP).75. Passiflora edmundoi SaccoBrazil (Piauí, Pernambuco, Bahia, Goiás, Minas Gerais, Rio de Janeiro). Occours mainly in campo rupestre and caatinga areas, ocasionally in the seasonal or atlantic forest (Nunes & Queiroz 2006).

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Selected Voucher:—Piedade, Silva s.n. (IPA-61585). 76. Passiflora edulis SimsUnited States, Mexico, Central America, Caribbean, Venezuela, Ecuador, Brazil (widely distributed ), Paraguay, Argentina. Selected Vouchers:—Piedade, Araújo et al. 577 (IPA, UFP).77. Passiflora galbana MastersBrazil (from Paraíba to Rio de Janeiro). Found in carrasco áreas, seasonal areas, campo rupestre, Atlantic forest, restinga and cerrado (Nunes & Queiroz 2006).Selected Voucher:—Pezinho, Araújo et al. 478 (UFP), Albuquerque 431 (IPA).78. Passiflora misera KunthCentral America, Caribbean, South America (widely distributed ). Campo rupestre, Atlantic forest, rarely in the caatinga and seasonal forests (Nunes & Queiroz 2006).Selected Voucher:—Zambana, Araújo et al. 348 (UFP).79. Passiflora suberosa L.United States, Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil (from Pernambuco to Rio Grande do Sul), Paraguay, Argentina (Killip 1938).Selected Voucher:—Macacos, Irapuan 21 (IPA).80. Passiflora watsoniana MastersBrazil (Pernambuco, Bahia, São Paulo). Atlantic and seasonal forests, campo rupestre and near restingas (Nunes & Queiroz 2006).Selected Voucher:— Macacos, Ojima 12 (UFP); Pezinho, Araújo et al. 591 (UFP); Piedade, Albuquerque 256 (IPA); Zambana, Melo et al. 125 (UFP).

POLYGALACEAE81. Securidaca diversifolia (L.) BlakeUnited States, Mexico, Central America, Caribbean, Ecuador, Bolivia, Brazil (Amazonas, Pará, Pernambuco, Alagoas, Bahia, Distrito Federal).Selected Voucher:—Macacos, Alves-Araújo & Araújo 728 (UFP); Zambana, Albuquerque 586 (IPA).

POLYGONACEAE82. Coccoloba ochreolata Wedd. (Fig. 2N)Bolivia, Brazil (Pará, Pernambuco, Alagoas, Bahia, Mato-Grosso).Selected Voucher:—Piedade, Alves-Araújo et al. 204 (IPA).

RANUNCULACEAE83. Clematis dioica L.United States, Mexico, Central America, tropical South America (widely distributed ) (Marcondes-Ferrreira 2003).Selected Vouchers:—Piedade, Alves-Araújo et al. 486 (IPA, UFP, ULM); Pezinho, Alves-Araújo et al. 604 (UFP).

RHAMNACEAE84. Gouania virgata Reissek (Fig. 3B)Venezuela, Guyanas, Bolivia, Brazil (from Pará to São Paulo) (Lima & Giulietti 2005).Selected Voucher:—Piedade, Araújo et al. 398 (UFP); Pezinho, Alves-Araújo 588 (UFP)

RUBIACEAE85. Sabicea grisea Cham. & Schltdl.Brazil (Amazonas, Pernambuco, Bahia). Selected Voucher:—Zambana, Araújo et al. 563 (UFP).

SAPINDACEAE86. Serjania salzmanniana Schltdl.Brazil (from Paraíba to Rio de Janeiro).Selected Voucher:—Zambana, Araújo 554 (UFP); Piedade, Araújo & Alves-Araújo 523 (UFP); Macacos, Araújo et al. 490 (UFP).

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87. Serjania subimpunctata Radlk. (Fig. 3 E, F) Brazil (Pernambuco, Sergipe, Bahia).Selected Voucher:—Piedade, Araújo et al. 588 (UFP); Macacos, Araújo & Alves-Araújo 499 (UFP). 88. Paullinia pinnata L. United States, Mexico, Central America, Caribbean, Ecuador, Peru, Bolivia, Brazil (Amazonas, Amapá, Pará, Maranhão, Pernambuco, Sergipe, Bahia, Goiás, Mato-Grosso, Paraná), Argentina. Selected Voucher:—Pezinho, Araújo et al. 442 (UFP).89. Paullinia racemosa Wawra (Fig. 2E)Brazil (from Pará to São Paulo). Atlantic forest, restinga, carrasco (Somner 2001).Selected Voucher:—Macacos, 13 Araújo et al. 489 (UFP).90. Paullinia trigonia Vell. (Fig. 2F)Brazil (from Pará to Rio Grande do Sul) (Somner 2001). Selected Voucher:—Zambana, Alves-Araújo et al. 659 (IPA).

SMILACACEAE91. Smilax syphilitica Griseb. in Mart. (Fig. 3G)Venezuela, Guyanas, Bolivia, Brazil (Roraima, Pernambuco, Espírito Santo). Selected Voucher:—Pezinho, Araújo et al. 613 (UFP); Piedade, Araújo & Alves-Araújo 525 (UFP); Macacos, Araújo et al. 481 (UFP); Zambana, Araújo et al. 560 (UFP).

VITACEAE92. Cissus erosa Rich.Mexico, Central America, South America until the southeast of Brazil (widely distributed ), Paraguay. Campos, cerrados, restingas and Forest boards (Lombardi 2002). Selected Voucher:—Macacos, Araújo et al. 618 (UFP); Zambana, Alves-Araújo et al. 637 (UFP).93. Cissus verticillata (L.) Nicolson & C.E.JarvisFrom Florida and north Mexico, Caribbean, South America, until the north of Argentina, Brazil (widely distributed ). Found mainly in forest boards, restingas, rarely in cerrados (Lombardi 2002).Selected Voucher:—Piedade, Araújo et al. 437 (UFP).

Acknowledgements

To CNPq/Brazil and BMBF/Germany as partners of the Project “Sustainability of remnants of the Atlantic rainforest in Pernambuco and its implications for conservation and local development” within this study is included. We are also thankful to Alex Monro for his numerous constructive comments that improved this manuscript.

References

Acevedo-Rodriguez, P. (2005) Vines and climbing plants of Puerto Rico and the Virgin Islands. Contributions from de United States National Herbarium 51: 1–483.

Alves-Araújo, A., Araújo, D., Marques, J., Melo, A., Maciel, J.R., Uirapuã, J., Pontes, T., Lucena, M.F.A., Bocage, A.L. & Alves, M. (2008) Diversity of Angiosperms in Atlantic Forest in the State of Pernambuco, Northeastern Brazil. Bioremediation, Biodiversity and Bioavailability 2(1): 14–26.

Amorim, A.M. (2003) The anomalous-stemmed species of Heteropterys subsect. Aptychia (Malpighiaceae). Brittonia55: 127–144.

Anderson, W.R. (2006) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168–204.APG III (2009). An update of the Angiosperms Phylogeny Group classification for the orders and families of flowering

plants: APG III. Botanical Journal of the Linnean Society 161(2): 105–121.Aymard, G.A. (1998) Dilleniaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (Eds), Flora of the

Venezuelan Guayana 4. Missouri Botanical Garden Press, St. Louis, pp. 671–685.Aymard, G.A., Cuello, N.L., Berry, P.E., Rudd, V.E., Cowan, R.S., Frantz, P.R., Maxwell, R.H., Stirton, C.H.,

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Poppendieck, H.-H., Lima, H.C., Fortunato, R.H., Stergios, B., Enrich, N.X., Neil, D.A., Pennington, R.T. & Gil, C. (1999) Fabaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (Eds), Flora of the Venezuelan Guayana 5. Missouri Botanical Garden Press, St. Louis, pp. 231–433.

Brummit, R.K. & Powell, C.E. (1992) Authors of Plant Names, a list of authors of scientific names of plants, with recommended standard forms of their names, including abbreviations. Royal Botanic Gardens, Kew.

Campanello, P.I., Garibaldi, J.F., Gatti, M.G. & Gillermo, G. (2007) Lianas in a subtropical Altantic Forest: Host preference and tree growth. Forest Ecology and Management 242: 250–259.

Chamisso, L.K.A. & Schlechtendal, D.F.L. (1829) De plantis in expeditione speculatoria Romanzoffiana observatis, Rubiaceae. Linnaea 4: 47–202.

Choisy, J.D. (1838) De Convolvulaceis, dissertation secunda. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève 8: 43–83.

CPRH (2003) Diagnóstico socioambiental do litoral norte de Pernambuco. Companhia Pernambucana do Meio Ambiente, Recife, 214 pp.

Cronquist, A. (1981) An integrated system of classification of flowering plants. Columbia University Press, New York, 1262 pp.

Darwin, C. (1867) On the movements and habits of climbing plants. Botanical Journal of the Linnean Society 9: 1–118.Forero, E. (1983) Connaraceae. Flora Neotropica Monograph 36: 1–207. Gentry, A.H. (1985) An ecotaxonomic survey of Panamanian lianas. In: D’Arcy, W.G. & Correa, M. (Eds), História

Natural de Panama. Missouri Botanical Garden Press, St. Louis, pp. 29–42.Gentry, A.H. (1991) The distribution and evolution of climbing plants. In: Putz, F.E. & Mooney, H.A. (Eds), The Biology

of vines. Cambridge University Press, Cambridge, pp. 3–49.Gentry, A.H. (1997) Bignoniaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (Eds), Flora of the

Venezuelan Guayana 3. Missouri Botanical Garden Press, St. Louis, pp. 403–491.Gerwing, J.J., Schnitzer, S.A., Burnham, R.J., Bongers, F., Chave, J., Dewalt, S.J., Ewango, C.E.N., Foster, R., Kenfack,

D., Martinez-Ramos, M., Parren, M., Parthasaranthy, N., Pérez-Salicrup, D.R., Putz, F.E., & Thomas, D.W. (2006) A standard protocol for lianas censuses. Biotropica 38: 256–261.

Govaerts, R., Frodin, D.G. & Radicliffe-Smith, A. (2000) World checklist and bibliography of Euphorbiaceae (and Pandaceae). The Royal Botanic Gardens, Kew, 1662 pp.

Hegarty, E.E. (1991) Vine-host interactions. In: Putz, F.E. & Mooney, H.A. The Biology of Vines. University Press, Cambridge, pp. 3–53.

Hegarty, E.E. & Cabalé, G. (1991 Distribution and abundance in forest comunities. In: Putz, F.E. & Mooney, H.A. The Biology of Vines. University Press, Cambridge, pp. 313–335.

Henderson, A., Galeano, G., Bernal, R. (1995) Field guide to the palms of the Américas. Princeton University Press, New Jersey, 363 pp.

Hora, R.C. & Soares, J.J. (2004) Estrutura fitossociológica da comunidade de lianas em uma floresta estacional semidecidual na Fazenda Canchim, São Carlos, SP. Revista Brasileira de Botânica 25(3): 323–329.

Jeffrey, C. (1978) Further notes on Cucurbitaceae: IV some New-World taxa. Kew Bulletin 33(2): 347–380.Kearns, D.M. (1998) Cucurbitaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (eds.), Flora of the

Venezuelan Guayana 4. Missouri Botanical Garden Press, St. Louis, pp. 431–461.Killip, L.P. (1938) The American species of Passifloraceae. Field Museum of Natural History, Botanical Series 49: 1–

613.Kinoshita, L.S. (2005) Apocynaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (Eds) Flora

Fanerogâmica do Estado de São Paulo 4. FAPESP, São Paulo, pp. 35–91.Klein, V.L. (2000) Estudo taxonômico de Cayaponia Silva Manso (Cucurbitaceae) no Brazil. Unpublished thesis.

Universidade de São Paulo, São Paulo, 289 pp.Lima, R.B. & Giulietti, A.M. (2005) Rhamnaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (Eds), Flora

Fanerogâmica do Estado de São Paulo 4. FAPESP, São Paulo, pp. 331–341.Lima, H.C., Lima, M.P.M., Vaz, A.M.S.F. & Pessoa, S.V.A. (1997) Trepadeiras da Reserva Ecológica de Macaé de Cima.

In: Lima, H.C. & Guedes-Bruni, R.R. (Eds), Serra de Macaé de Cima: Diversidade Florística e Conservação em Mata Atlântica. Jardim Botânico do Rio de Janeiro, Rio de Janeiro, pp. 75–87.

Lombardi, J.A. & Lara, A.C.M. (2003) Hippocrateaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (Eds), Flora Fanerogâmica do Estado de São Paulo 3. FAPESP, São Paulo, pp. 109–122.

Lombardi, J.A. (2002) Vitaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (eds.), Flora Fanerogâmica do Estado de São Paulo 2. FAPESP, São Paulo, pp. 365–374.

Marcondes-Ferreira, W. (2003) Ranunculaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (Eds), Flora Fanerogâmica do Estado de São Paulo 3. FAPESP, São Paulo, pp. 279–283.

Masters, M.T. (1875) Aristolochiaceae In: Martius, C.F.P & Endlicher, S. (Eds) Flora Brasiliensis 4(2): 100.Menega, M.W. & Hedin J.P. (1999) Hippocrateaceae. In: Steyermark, J.A., Berry, P.E., Yatskievych, K.& Holst, B.K.

(Eds), Flora of the Venezuelan Guayana 5. Missouri Botanical Garden Press, St. Louis, pp. 594–617.

Phytotaxa 8 © 2010 Magnolia Press • 23CLIMBING PLANTS OF LOWLAND ATLANTIC FOREST, IGARASSU

Page 24: Climbing plants of a fragmented area of lowland Atlantic Forest, Igarassu, Pernambuco (northeastern Brazil)

MMA (2000) Avaliação e ações prioritárias para a conservação da biodiversidade da Mata Atlântica e Campos Sulinos. Ministério do Meio Ambiente, Brasília, 40 pp.

Nunes, T.S. & Queiroz L.P. (2006) Flora da Bahia: Passifloraceae. Sitientibus Série Ciências Biológicas 6(3): 194–226.Odegaard, F. (2000) The relative importance of trees versus lianas as hosts for phytophagus beetles (Coleoptera) in

tropical; forests. Journal of Biogeography 27: 283–296.Pereira, J.F. (2005) Asclepiadaceae. In: Wanderley, M.G.L., Shepherd, G.J. & Giulietti, A.M. (Eds), Flora Fanerogâmica

do Estado de São Paulo 4. FAPESP, São Paulo, pp. 93–156.Profice, S.R. (1997) Two new species of Mendoncia (Acanthaceae) from Bahia, Brazil. Brittonia 49: 67–70.Putz, F.E. (1984) The natural history of lianas on Barro Colorado Island, Panama. Ecological Society of America 65:

1713–1724.Queiroz, L.P. (2006) New species and new combinations in Phanera Lour. (Caesalpinoideae: Cercideae) from de

caatinga biome. Neodiversity 6: 1–10. Reddy, M.S. & Parthasarathy, N. (2003) Liana diversity and distribution in four tropical dry evergreen forests on the

Coromandel Coast of south India. Biodiversity and Conservation 12: 1609–1627. Rezende, A.A & Ranga, N.T. (2005) Lianas da Estação Ecológica do Noroeste Paulista, São José do Rio Preto/Mirassol,

SP, Brazil. Acta Botânica Brazilica 19(2): 273–279.Schnitzer, S.A. & Bongers, F. (2002) The ecology of lianas and their role in forests. Trends in Ecology & Evolution

17(5): 223–230. Schumann, K. (1895) Bignoniaceae. In: Engler, H.G.A.; Prantl. K. A. (Eds) Die natürlichen pflanzenfamilien. 4(3b).

Eugelmann, Leipzig, pp. 189–306.Silva, H.C.H. (2004) Efeito de borda na fisionomia e estrutura da vegetação em fragmentos de floresta Atlântica de

tamanhos distintos em Igarassu - Pernambuco. Unpublished thesis. PPGB/UFRPE, 91 pp.Simão-Bianchini, R. (1993) Ipomoea L. (Convolvulaceae) no sudeste do Brazil. Umpublished tesis. Universidade de São

Paulo, 476 pp.Solórzano, S. & Ibarra-Manríquez, G. & Oyama, K. (2002) Liana diversity and reproductive attributes in two tropical

forests in Mexico. Biodiversity and Conservation 11: 197–212. Somner, G.V. (2001) Paullinia L. (Sapindaceae): Morfologia, Taxonomia e Revisão de Paullinia sect. Phygoptilon.

Umpublished tesis. Universidade de São Paulo, 275 pp.Souza, C.V. & Lorenzi, H. (2008) Botânica Sistemática. Instituto Plantarum, Nova Odessa, 704 pp.Tibiriçá, Y.J.A., Coelho, L.F.M & Moura, L.C. (2006) Florística de lianas em um fragmento de floresta estacional

semidecidual, Parque Estadual de Vassununga, Santa Rita do Passa Quatro, SP, Brazil. Acta Botanica Brazilica20(2): 339–346.

Udulutsch, R.G., Assis, M.A. & Picchi, D. (2004) Florística de trepadeiras numa floresta estacional semidecídua, Rio Claro - Araras, Estado de São Paulo, Brazil. Revista Brasileira de Botânica 27(1): 125–134.

Vitta, F.A. & Bernaci, L.C. (2004) A new species of Passiflora in section Tetrastylis (Passifloraceae) and two overlooked species of Passiflora from Brazil. Brittonia 56 (1): 89–95.

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