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Inventory • Biota Neotrop. 25 (4) • 2025 • https://doi.org/10.1590/1676-0611-BN-2025-1765 linkcopy

Open-access Climbers of an insular landscape in upper Paraná River of the Southeast Brazil: floristic composition and identification keys

Trepadeiras de uma ilha fluvial no alto Rio Paraná no Sudeste do Brasil: composição florística e chaves de identificação

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

Abstract

Climbing species in the Neotropics are diverse and polyphyletic, contributing approximately 10,000 species to angiosperms and occurring across a wide range of forests and savannas. They play essential roles in these ecosystems by interacting with pollinators and influencing the growth dynamics of host plants. Despite their ecological importance, little is known about the richness and composition of climbing plants in fluvial island environments. In this study, we aimed to: (i) inventory the climbing plants species of Ilha Solteira, São Paulo, Southeast Brazil; and (ii) provide identification keys for the species recorded in the area. We conducted unsystematic walks, with bimonthly collections in 2018 and monthly collections in 2019, mainly along edges and trails. We deposited the voucher specimens in the Ilha Solteira Herbarium (HISA) and identified the plants using standard taxonomic methodology. We recorded 39 species of climbing plants, belonging to 33 genera and 18 families, with Fabaceae, Apocynaceae, and Convolvulaceae being the richest families. Most species were woody (59%), and zoochorous dispersal was the predominant syndrome. We also developed an identification key for the families of climbing plants of Ilha Solteira, which can be further expanded into seven family-specific keys: Apocynaceae, Bignoniaceae, Convolvulaceae, Fabaceae, Malpighiaceae, Passifloraceae, Sapindaceae, and Vitaceae. As the first inventory of climbing plants in a fluvial island ecosystem, our findings are expected to support further ecological and botanical research, as well as contribute to practical measures for biodiversity conservation and management in the region.

Keywords
Atlantic Forest; Seasonal Semideciduous Forest; liana; vines

Resumo

As espécies de trepadeiras nos Neotrópicos são diversas e polifiléticas, contribuem com aproximadamente 10.000 espécies de angiospermas e estão presentes em vários tipos de florestas e savanas. Elas desempenham papéis cruciais nestes ecossistemas, estabelecendo interações com polinizadores e influenciando o crescimento de plantas hospedeiras. Considerando a lacuna de informações sobre a riqueza e composição desta forma de vida em ambientes insulares fluviais, este estudo teve como objetivo: i) inventariar as espécies trepadeiras da Ilha Solteira no Rio Paraná, que inspirou o nome do municipio de Ilha Solteira, no estado de São Paulo, Brasil; e ii) fornecer chaves de identificação para as espécies encontradas na área. Por meio de caminhadas assistemáticas, as espécies de trepadeiras foram inventariadas em coletas bimestrais, em 2018, e mensais em 2019, principalmente ao longo das bordas e trilhas. Os espécimes foram depositados no Herbário Ilha Solteira (HISA) e identificados usando literatura especializada. Foram identificadas 39 espécies de trepadeiras, distribuídas em 33 gêneros e 18 famílias, em que se destaca a riqueza de espécies de Fabaceae, Apocynaceae e Convolvulaceae. A maioria das espécies era lenhosa (59%), e a síndrome de dispersão zoocórica foi a mais comum. Foi organizada uma chave de identificação para as famílias das trepadeiras da Ilha Solteira, a qual leva, eventualmente, a outras sete chaves de famílias, especificamente: Apocynaceae, Bignoniaceae, Convolvulaceae, Fabaceae, Malpighiaceae, Passifloraceae, Sapindaceae e Vitaceae. Dado o ineditismo deste inventário para ecossistemas insulares fluviais, tem-se a expectativa de que estes resultados subsidiarão experimentos científicos em ecologia e botânica, além de auxiliarem medidas práticas de conservação e manejo da biodiversidade na região.

Palavras-chave
Floresta Atlântica; Floresta Estacional Semidecidual; liana; vinhas

Introduction

The richness of climbing species in the Neotropics, as well as their polyphyletic origin (Gianoli 2004, 2015; Burnham & Romero-Saltos 2015), is widely recognized and its contribution to the total percentage of angiosperm species is around 10 thousand species (Polisel 2017), distributed across humid, seasonal and dry tropical forests, and savannas (Schnitzer & Bongers 2002; Gentry 1991).

The effects of the presence of climbing species in forest ecosystems are also unquestionable (Schnitzer 2018), either as key resources for pollinators and herbivores (Schnitzer et al. 2020; Hilje et al. 2017) or as interferers in the development of phorophytes, in which positive correlations are expected between their functional traits (Werden et al. 2018; Sfair et al. 2010) and the diversity of tree species (Sfair & Martins 2011). Or even, in canopy height (Meyer et al. 2019), infestation levels, age, height and diameter of the phorophyte (Reis et al. 2020, Visser et al. 2017, Zulqarnain et al. 2016). Its richness and abundance of climbing species are influenced by the temperature and latitudinal gradient (Lobos-Catalán & Jiménez-Castillo 2019), altitude (Mohandass et al. 2017), precipitation seasonality (Parolari et al. 2019) and water retention in the soil (Manzané-Pizón et al. 2018).

Particularly, the nature and intensity of disturbances are related to the speed of propagation of lianas, which would modulate the composition and structure of forest communities, as well as the direction of secondary succession. (Melis et al. 2020; Hogan et al. 2017; Ledo & Schnitzer 2014). On the one hand, it is possible to draw inferences on the effects of forest fragmentation in the presence and abundance of hyperabundant climbing species in intensely disturbed sites (Ledo & Schnitzer 2014), on the other hand, it is necessary to consider those species of climbers that remain relatively rare or in population decline in these same areas and are therefore undersampled in floristic inventories and possibly, maintaining underestimations of the total richness of climber species. In any case, these gaps reveal research potential in ecology and conservation with this group of plants. (Schnitzer et al. 2020; Vargas et al. 2020; Schnitzer 2018). It is worth noting that in semideciduous seasonal forests of the Atlantic Forest, which have a long history of intensive disturbances, climbing plants have increased in abundance (Püttker et al. 2020; Haddad et al. 2015; Farah et al. 2014), due to the morphological and ecophysiological traits that allow them to maintain their leaves during drought and the way their root system exploit soil water (Chen et al. 2015; De Guzman et al. 2017; De Deurwaerder et al. 2018; Schnitzer 2018).

In the state of São Paulo, Southeastern Brazil, several floristic surveys highlighted the high richness of climbing species in seasonal forests, including fragmented forests (Vargas et al. 2020; Polisel 2017; Rezende et al. 2007; Rezende & Ranga 2005). Although these works have emphasized the importance of climbers in terms of richness, the flora of insular environments in Brazil remains little known (Meira-Neto et al. 1998), and no inventories of climbing plants have yet been conducted in insular environments in Brazil (Polisel 2017). This gap implies limited knowledge of the specific composition of climbing species in these ecosystems, as well as the environmental and biological factors that effectively determine their richness and structure.

Referring to the theory of island biogeography and its predictions about the effects of habitat fragmentation (MacArthur & Wilson 1963; 1967), it would be expected that the composition, richness and diversity of plants in river island systems would be influenced, a priori, by the total area of the island, its shape, distances from the banks, river flow dynamics (directly linked to temporal changes in shape, topography and island pedology). Landscape characteristics would also exert an influence on island colonization patterns, modulating gene flow based on the likelihood of seed and diaspore arrival, given in part by the movement of dispersal agents and pollinators (Nascimento et al. 2012; Haila 2002; Brown & Kodric-Brown 1977). However, these communities’ floristic composition, structure, and dynamics also respond to disturbance variables directly and indirectly associated with fragmentation and intensive human intervention (Bueno & Peres, 2019; Zambrano et al. 2020).

Identification keys are essential tools for recognizing local flora. However, they typically rely on reproductive traits (Edwards & Morse 1995), which limit their usefulness for identifying plants without flowers or fruits (Freitas & Oliveira 2002). Therefore, developing identification keys based exclusively on vegetative characters is particularly valuable for research and technical studies that require accurate plant identification within a limited time frame. Moreover, keys based on vegetative traits enable plant collection throughout the year.

In this context, we aimed to inventory the climbing plants of a fluvial island in the Paraná River, in the municipality of Ilha Solteira, Southeast Brazil, classify their climbing mechanisms and dispersal syndromes, and provide an identification key based on vegetative traits for all recorded species.

Material and Methods

1. Study area

Ilha Solteira, a river island on the Paraná River, belongs to the municipality of the same name in São Paulo State, near the border with Mato Grosso do Sul State, where the extremes are located geographic coordinates: 20º23’21”S e 51º21’54”W; 20º24’21”S e 51º22’37”W (Figure 1). The Ilha Solteira gave its name to the municipality of which it is part and is considered historical-cultural heritage by Law 1054 of August 21, 2003, as well as an Environmental Protection Area, by Law 1125, of February 6, 2004 (http://leismunicipa.is/wphke).

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Figure 1.
Study area, Ilha Solteira, São Paulo, Southeast Brazil (20°23'56.56"S, 51°22'25.24"W). a. General view of the island; b. Location of the island on the Paraná River; c. Border between the states of São Paulo and Mato Grosso do Sul. Source: Google Earth.

The vegetation of the river island Ilha Solteira is classified as Semideciduous Seasonal Forest (Figure 2), a phytophysiognomy of the Atlantic Forest (IBGE, 2012). Originally, the island covered an area of 43 ha, however, due to the impacts associated with the construction of the Ilha Solteira Hydroelectric Powerplant in the 1960s, its area was reduced by approximately 65% and it now spans less than 15 ha (Wilson Júnior et al. 2017).

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Figure 2.
Overview of Ilha Solteira, São Paulo State, Southeast Brazil, highlighting the vegetation — Seasonal Semideciduous Forest. a. Vegetation during the wet season; b. Vegetation during the dry season. Source: Santos, A.M.D.

The climate of Ilha Solteira is tropical, with an average annual temperature of 23.5°C and higher rainfall during the summer. The total annual rainfall is 1,316 mm. The driest month is August, with an average of 22 mm of rainfall, while January, the month with the highest rainfall, averages 237 mm (CLIMATE-DATA 2021).

2. Sampling

To conduct the floristic inventory, we collected climbing plant species in reproductive or vegetative stages bimonthly, in 2018, and monthly, in 2019. We sampled along the edges and trails of the island using an unsystematic approach.

Classification of climbing mechanisms was based on the criteria of Sperotto et al. (2020): 1) passive climbing plants, containing scrambling, hooks, and adhesive roots; and 2) active climbing plants, containing twining, tendrils, prehensile branches, twining petioles, and twining inflorescences. For growth habit, we include all climbers that germinate on the forest ground and have true secondary growth stems (woody climbers = liana) and those lacking true wood they were identified as non-woody (herbaceous climbers) (Gerwing 2006), and dispersal syndrome (anemochory, zoochory or autochory) we followed what was proposed by Van der Pijl (1982).

We dried the collected material using standard techniques (Fidalgo & Bononi, 1984) and deposited it in the Ilha Solteira Herbarium (HISA). We identified the specimens with the assistance of specialized references. Accepted names were verified according to Flora e Funga do Brasil (2025). We organized the identification key based exclusively on vegetative characters, and the nomenclature of the structures followed Bell and Bryan (2008), in addition to specific references for each family.

Results

A total of 39 climbing plant species were identified, belonging to 33 genera and 18 families (Figure 3), and one species was identified to the genus level (Table 1). The families with the highest number of species were Fabaceae (six species), representing approximately 15% of the species sampled; Apocynaceae and Convolvulaceae (four species each), each accounting for around 10%; and Bignoniaceae, Malpighiaceae, Passifloraceae, and Vitaceae (three species each), each contributing about 7%. Of the species sampled, 23 were woody climbers, accounting for 59% and 16 were herbaceous, comprising 40%.

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Figure 3.
Climber species sampled on Ilha Solteira: a. Cynanchum montevidense (Apocynaceae); b. Aristolochia odoratissima (Aristolochiaceae); c. Psiguria ternata (Cucurbitaceae); d. Dioclea virgata (Fabaceae); e. Amorimia pubiflora (Malpighiaceae); f. Passiflora foetida (Passifloraceae). Source: Sawakuchi, S.Y.
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Table 1.
List of climbing species on Ilha Solteira, Paraná River, São Paulo, Brazil. Habit: Herbaceous (non-woody climbers) (H), Liana (woody climbers) (L); Climbing Mechanism (CM): Scrambling, Hooks, Twining, Tendrils; Dispersal Syndrome (SD): Anemochory (ANE), Zoochory (ZOO), Autochory (AUT).

Among the species listed (Table 1), 22 exhibit a twining climbing mechanism, representing 57% of the species; 14 species have tendrils (35%), and three are scandent species (7%). Therefore, according to the classification of Sperotto et al. (2020) there are 36 active species and only three passive species. Regarding dispersal syndrome, 15 species are zoochorous, accounting for about 36% of the total species sampled; 14 are anemochoric (35%), and 11 are autochoric (28%).

Based on the data obtained, an identification key was developed for the climbing plant families recorded on Ilha Solteira. This key, in turn, is directed to keys for seven specific families: Apocynaceae, Bignoniaceae, Convolvulaceae, Fabaceae, Malpighiaceae, Passifloraceae, Sapindaceae, and Vitaceae. We found that the most developed characteristics for species differentiation were the presence or absence of tendrils, leaf type, venation, and indument type.

Furthermore, some families, such as Vitaceae and Cucurbitaceae, appear more than once in the key due to variations in the primary vegetative characteristics of their representatives, with species with both simple and compound leaves found within the same family.

Key to families of lianas from the River Island

  • 1 Compound leaves ................................................................................ 2

  • 1’Simple leaves ................................................................................ 6

  • 2 Opposite leaves ................................................................................ Bignoniaceae (Chave 2)

  • 2’Alternate leaves ................................................................................ 3

  • 3 Absence of tendril ................................................................................ Fabaceae (Chave 4)

  • 3’ Presence of tendril ................................................................................ 4

  • 4 Circinate tendril ................................................................................ Sapindaceae (Chave 7)

  • 4’Other types of tendrils ................................................................................ 5

  • 5 Axillary tendrils ................................................................................ Cucurbitaceae (Psiguria ternata)

  • 5’ Opposite tendrils ................................................................................ Vitaceae (Chave 8)

  • 6 Opposite leaves or fasciculated ................................................................................ 7

  • 6’Alternate leaves ................................................................................ 12

  • 7 Fasciculated leaves ................................................................................ Nyctaginaceae (Pisonia aculeata)

  • 7’ Opposite leaves ................................................................................ 8

  • 8 Presence of latex ................................................................................ Apocynaceae (Chave 1)

  • 8’Absence of latex ................................................................................ 9

  • 9 Petiole < 1 cm ................................................................................ Combretaceae (Combretum laxum)

  • 9’ Petiole > 1 cm ................................................................................ 10

  • 10 Presence of stipule ................................................................................ Malpighiaceae (Chave 5)

  • 10’Absence of stipule ................................................................................ 11

  • 11 Actinodromous nervation ................................................................................ Asteraceae (Mikania triangularis)

  • 11’ Craspedodromous nervation ................................................................................ Dilleniaceae (Dioliocarpus dentatus)

  • 12 Presence of tendril ................................................................................ 13

  • 12’Absence of tendril ................................................................................ 15

  • 13 Gland in the petiole and/or leaves margin ................................................................................ Passifloraceae (Chave 6)

  • 13’Absence of glands in the petiole and/or leaves margin ................................................................................ 14

  • 14 Actinodromous nervation ................................................................................ Smilacaceae (Smilax brasiliensis)

  • 14’ Nervation other types ................................................................................ Vitaceae (Chave 8)

  • 15 Armed plant ................................................................................ Cannabaceae (Celtis chichape)

  • 15’ Non-armed plant ................................................................................ 16

  • 16 Presence of latex ................................................................................ Convolvulaceae (Chave 3)

  • 16’Absence of latex ................................................................................ 17

  • 17 Presence of stipule ................................................................................ 18

  • 17’Absence of stipule ................................................................................ Cucurbitaceae (Momordica charantia)

  • 18 Lobed leaves ................................................................................ Euphorbiaceae (Dalechampia scandens)

  • 18’ Entire leaves ................................................................................ 19

  • 19 Presence of pseudostipule ................................................................................ Aristolochiaceae (Aristolochia odoratissima)

  • 19’Absence of stipules ................................................................................ Menispermaceae (Cissampelos andromorpha)

KEY 1: Apocynaceae

  • 1 White latex ................................................................................ 2

  • 1’ Colorless latex ................................................................................ 3

  • 2 Base of the lamina cordate ................................................................................ Cynanchum montevidense

  • 2’ Base of the lamina truncate ................................................................................ Funastrum clausum

  • 3 Glabrous leaves ................................................................................ Mesechites mansoana

  • 3’ Leaves with indument ................................................................................ Prestonia tomentosa

KEY 2: Bignoniaceae

  • 1 Trifid tendril, uncinates ................................................................................ Dolichandra unguis-cati

  • 1’Simple tendril ................................................................................ 2

  • 2 Articulated petiole and laciniate lamina ................................................................................ Adenocalymma paulistarum

  • 2’ Non-articulated petiole and oblong to ovate lamina ................................................................................ Tanaecium pyramidatum

KEY 3: Convolvulaceae

  • 1 Stem with solid pith ................................................................................ Ipomoea sidifolia

  • 1’ Stem with fistulous pith ................................................................................ 2

  • 2 Limbus with glabrous adaxial surface ................................................................................ Ipomoea alba

  • 2’ Limbus with incana to pubescent adaxial surface ................................................................................ 3

  • 3 Actinodromous venation ................................................................................ Camonea umbellata

  • 3’ Craspedodromous venation ................................................................................ Ipomoea carnea

KEY 4: Fabaceae

  • 1 Unifoliate leaf ................................................................................ Centrosema sagittatum

  • 1’Trifoliolate leaf ................................................................................ 2

  • 2 Glabrous stem and petiole ................................................................................ Macropsychanthus latifolius

  • 2’ Stem and petiole with indument ................................................................................ 3

  • 3 Parallel venation ................................................................................ Rhynchosia phaseloides

  • 3’ Pinnate venation (other type) ................................................................................ 4

  • 4 Asymmetrical lateral leaflets ................................................................................ Calopogonium caeruleum

  • 4’Symmetrical lateral leaflets ................................................................................ 5

  • 5 Lanceolate stipules with parallel striations and leaves with an acute apex ................................................................................ Clitoria falcata

  • 5’ Inconspicuous stipules and culminated apex ................................................................................ Dioclea virgata

KEY 5: Malpighiaceae

  • 1 Limbus with sericeous adaxial surface ................................................................................ Banisteriopsis muricata

  • 1’ Limbus with other type of indumentum in the adaxial surface ................................................................................ 2

  • 2 Lamina discolor ................................................................................ Amorimia pubiflora

  • 2’ Lamina concolor ................................................................................ Diplopterys sp.

KEY 6: Passifloraceae

  • 1 Entire lamina ................................................................................ Passiflora alata

  • 1’ Lobed leaves ................................................................................ 2

  • 2 Leaves villose ................................................................................ Passiflora capsularis

  • 2’ Leaves pilose ................................................................................ Passiflora foetida

KEY 7: Sapindaceae

  • 1 Transverse section of the stem with 1 larger central cylinder and 3 peripheral cylinders ................................................................................ Serjania marginat

  • 1’ Transverse section of the stem with 1 large central cylinder and 5 peripheral cylinders ................................................................................ Paullinia elegans

KEY 8: Vitaceae

  • 1 Canaliculated and reddish branches ................................................................................ Cissus erosa

  • 1’ Non-canaliculated branches or canaliculated and non-reddis. ................................................................................ 2

  • 2 Limbus membranaceous and blackish when dry ................................................................................ Cissus tinctoria

  • 2’ Chartaceous limbus and not blackish when dry ................................................................................ Cissus verticillata

Discussion

This is the first studies to inventory climbing plants on a river island in Brazil. Furthermore, these are the first collections from the study area recorded in a herbarium, emphasizing the pioneering nature of this study.

Observing the number of families and species of climbing plants in surveys carried out in Semideciduous Seasonal Forest of São Paulo state, we found that this number varies from 33 families and 148 species to 19 families and 65 species (Udulutsch et al. 2004, Rezende & Ranga 2005; Tibiriçá et al. 2006; Udulutsch et al. 2010; Vargas et al. 2013; Ribeiro-Neto et al. 2018; Scudeler et al. 2019). The number of species sampled on the Ilha Solteira (39) represents only 62% of the number of species sampled by Scudeler et al. (2019), which was the study that sampled the smallest number of climbing species.

Difference in sampling methods and fragment size can affect species richness (Santos et al. 2009). However, proximity to water is also a relevant factor for lower species richness, as pointed out in other studies that sampled climbing plants in riparian forest (Funch et al. 1996; Alcalá et al. 2006; Vargas et al. 2013). Liana richness is lower in seasonally flooded forests, such as those inIlha Solteira, than in non-flooded forests because many liana species apparently lack ecophysiological traits, such as hypertrophied lenticels on their stems, through which potentially toxic compounds associated with anaerobiosis are released (Oliveira et al. 2014; Joly 1994).

Anemochory is the dominant form of dispersal in most studies that sampled climbing plants. (Morellato & Leitão-Filho 1996; Vargas et al. 2013; 2018). However, in the present study most species exhibited zoochoric dispersion. This result may possibly be related to the adaptation of the species to proximity to water and to the fact that it is an island. Kubitzki (1971) reported that Doliocarpus dentatus is more frequent in humid environments, occurring in both primary and secondary forests. In Brazil it is found mainly in gallery forests and, less frequently, in Cerrado areas near watercourses. In Ilha Solteira, this species was common, occurring both along the riverbanks and within the island interior.

The families with the highest number of species were Fabaceae, Apocynaceae and Convolvulaceae. This pattern, in which few families account for more than half of the species, was observed by Gentry (1991) and has also been reported in other studies (Morellato & Leitão-Filho, 1998; Hora & Soares, 2002; Rezende & Ranga, 2005; Tibiriçá et al., 2006; Udulutsch et al., 2010; Vargas et al., 2018; Ribeiro-Neto et al., 2018). Although Bignoniaceae is commonly the richest family in low altitude Neotropical Forest, particularly Semideciduous Seasonal Forest (Gentry 1991), it was not among the most species-rich families in the study. Fabaceae, which is generally not the family with the highest species richness in Semideciduous Seasonal Forest areas, emerged as the richest family here, a pattern also observed in surveys carried out in ecotone areas between the Dense Ombrophilous Forest and the Semideciduous Seasonal Forest (Araújo & Alves 2010; Villagra & Romaniuc-Neto 2010; Barros et al., 2009).

Udulutsch et al. (2004) carried out a study on the climbing floristics in a Semideciduous Seasonal Forest in Rio Claro and Araras in the State of São Paulo, and for the climbing mechanisms, they found the predominance of voluble species, with a total of 43% of the species, followed by forms with tendrils (39%) and, to a lesser extent, by nongripping vines, with 18% of the species. Ilha Solteira, twining vines had a predominance of 56.41%. This dominance was also found in Semideciduous Seasonal Forest by Tibiriçá et al. (2006) and Durigon et al. (2009). And in riparian forests Alcalá et al. (2006) and Vargas et al. (2013).

Due to the degradation caused during the construction of the Ilha Solteira Hydroelectric Powerplant, it was expected that herbaceous climbers would be predominant (Lima et al. 1997; Haber 2000), once they are commonly found in clearings, disturbed areas, and regions affected by forest fragmentation. Factors that increase light availability, such as edges and gaps, generally favor the development of herbaceous climbers (Morellato & Leitão-Filho 1996). However, on the river island, despite the highlight incidence at the edges, a greater number of woody species was observed, a pattern also reported in Semideciduous Seasonal Forest (Udulutsch et al. 2004; Vargas et al. 2013). This indicates that, even after prolonged anthropogenic impact, the climbing flora of Ilha Solteira remains diverse. Moreover, this study contributes to expanding knowledge about climbing plants on river islands, an area still poorly documented in Brazil.

This novelty underscores the importance of this study as the first inventory of climbing plants in a fluvial island ecosystem. The findings are expected to support further ecological and botanical research and contribute to practical measures for biodiversity conservation and management in the region.

Data Availability

The datasets generated during and/or analyzed during the current study are available at: https://doi.org/10.48331/scielodata.F8UVKY.

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Edited by

  • Associate Editor
    Alexander Vibrans.

Publication Dates

  • Publication in this collection
    08 Dec 2025
  • Date of issue
    2025

History

  • Received
    06 Mar 2025
  • Accepted
    28 Oct 2025
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