Menu
Brasil
home Sumário navigate_before Anterior Atual Seguinte navigate_next
home Sumário
Original Article • Acta Bot. Bras. 39 • 2025 • https://doi.org/10.1590/1677-941X-ABB-2024-0075 linkcopy

Open-access A new species of Ipomoea (Convolvulaceae) for the Brazilian Flora discovered in an anthropic habitat

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    Abstract

    Ipomoea is one of the most diverse genera of Angiosperms, with 700 to 800 species. Several of them are widespread ruderal, often highly polymorphic, making species delimitation a challenge. When collecting Brazilian Convolvulaceae, one specimen from an anthropic Cerrado area in Mato Grosso do Sul state, caught our attention. Morphological, palynological, and anatomical analyses were performed to compare this unidentified species to its closest relatives. Ipomoea cryptocarpa Belo & Buril, sp. nov., the new species described here, is similar to I. bahiensis, a widespread ruderal plant, due to the shape of the leaves and the presence of an appendage on the sepals. However, they can be distinguished by the shape of the sepals appendage, fruit length, pollen morphology, indument on the leaf blade, petiole, peduncle, and anatomical characters, such as the contour of epidermis, distribution, and types of stomata, the number of laticiferous canals in the petiole, and types of collenchyma observed in the stem. We provide a diagnosis, besides morphological, anatomical, and palynological descriptions, illustrations, and notes on the distribution of the new species. Its informal conservation status is presented as Data Deficient (DD) since only one population is known so far.

    Keywords:
    Brazilian flora; leaf anatomy; morning glory; palynology; species boundaries

    Introduction

    Convolvulaceae Juss. comprises 60 genera and approximately 1,900 species distributed worldwide but most diverse in tropical areas, with few representatives in temperate zones (Staples & Brummitt, 2007; WCSP, 2024). It includes the important crop sweet-potato (Ipomoea batatas L.), and other species with pharmacological and ornamental uses (Meira et al., 2012; Wood et al., 2020; Magalhães et al., 2022). Around 430 species are recorded in Brazil (Flora e Funga do Brasil, 2024), of which some were described in the last decade (e.g. Pastore & Simão-Bianchini, 2016; Wood et al., 2016; Belo et al., 2023a; Santos & Buril, 2024).

    Ipomoea L. is the most diverse genus of Convolvulaceae, and it is among the most diverse of angiosperms, distributed mainly in the Tropics, comprising between 700 and 800 species, depending on the circumscription (Eserman et al., 2020; Wood et al., 2020). It is characterized by the following features: an entire style, a stigma usually with two globose lobes, echinate and pantoporate pollen grains, and a capsular fruit (Stefanović et al., 2003; Wood et al., 2020). It was the focus of a recent global monograph and molecular phylogenetic inference that led to a proposal to recognize it in a broader sense, including all other echinate-pollen genera within the tribe Ipomoeeae Hall. f., such as Argyreia Lour., Astripomoea A.Meeuse, and Rivea Choisy (Muñoz-Rodríguez et al., 2023). As a result of these studies, several new species were combined (Wood et al., 2020; WCSP, 2024).

    In Brazil, 160 species of Ipomoea are currently known, 40% of which are endemic to the country (Santos et al., 2021; Simão-Bianchini et al., 2024). The greatest diversity is found in the Cerrado (96 spp.) and Caatinga (58 spp.) domains, in areas of open vegetation with high light incidence (Santos et al., 2020a; Simão-Bianchini et al., 2024). Various species of Ipomoea are ruderal, being abundant in areas with anthropic influence - such as I. asarifolia Roem. & Schult., I. bahiensis Willd. ex Roem. & Schult., I. cairica (L.) Sweet, I. nil (L.) Roth, I. tricolor Cav., and I. triloba L. Some are rare, and others are considered as threatened, such as I. cavalcantei D.F.Austin, I. lanifolia D.Santos & Buril, I. maranhensis D.Santos & Buril, and I. pantanalensis J.R.I.Wood & C.Urbanetz (Wood et al., 2016; Santos et al., 2020b; Wood et al., 2020; Santos et al., 2021).

    Its species are often polymorphic, and species delimitation can be a challenge due to morphological characters overlapping in closely related species (Miller et al., 1999; Stefanović et al., 2003; Muñoz-Rodríguez et al., 2019; Wood et al., 2020). Integrative approaches are being used in Convolvulaceae to enlighten species recognition, especially when morphology itself is too variable or difficult to interpret. Thus, recent studies focused on resolving species boundaries are laying on morphometrics - e.g. in Daustinia Buril & A.R.Simões (Alencar et al., 2020; 2024), and Jacquemontia Choisy (Belo et al., 2024a); molecular - e.g. in Distimake Raf. (Pisuttimarn et al., 2023), and Ipomoea (Muñoz-Rodríguez et al., 2019); palynological - e.g. in Evolvulus L., Ipomoea, Jacquemontia, Distimake ( Vital et al., 2008; Vasconcelos et al., 2015; Belo et al., 2024b); and anatomical characters - e.g. in Argyreia (Traiperm et al., 2017), Camonea Raf. (Santos et al., 2024), Daustinia (Alencar et al., 2022), Ipomoea (Santos et al., 2020a; b), and Jacquemontia (Belo et al., 2023b; c).

    During field expeditions in Brazilian territory, we collected a morphotype of Ipomoea in the Brazilian Cerrado with unusual sepals and fruits, growing in an anthropic area. In the first analysis, we identified it as belonging to the morphological variation of I. bahiensis. However, after in-depth and careful analyses, remarkable characters were found, excluding this specimen from the morphological range reported to I. bahiensis. The present study provides analyses through morphological, anatomical, and palynological studies at the population level to test the hypothesis that there are morphological reasons to differentiate these two biological entities. Therefore, here we describe this morphotype as a new species.

    Material and methods

    Macromorphological analyses

    Field expeditions were conducted in the Cerrado, Amazon, Pantanal, and Caatinga domains, in the states of Bahia, Mato Grosso do Sul, Pará, Pernambuco, and Piauí, Brazil, between May 2023 and April 2024. Specimens of morphologically close species were analyzed in the following herbaria: ALCB, ASE, CGMS, CEN, EAC, EAN, HUEFS, IAN, INPA, IPA, K, MAC, MBM, MG, NY, P, PEUFR, RB, SP, SPF, UEC, and UFP (Thiers, 2024). We also carefully examined the type material of each possibly related species available at JSTOR (https://plants.jstor.org/). The taxonomic description and terminology follow Harris & Harris (2001) and Wood et al. (2020). The phenological period was defined based on the collected specimens. A preliminary conservation status assessment was performed according to the IUCN Red List Categories and Criteria (IUCN, 2012; 2024). The map of occurrence was produced using QGIS Software 3.22 (QGIS.ORG, 2023) (Fig. 1). The geographic coordinates used to prepare the distribution map of the populations used in the micromorphological analyses and of the specimens of I. bahiensis can be found in the supplementary material 1. A list of examined specimens that were used for the taxonomic treatment of I. bahiensis can be found in supplementary material 2. Dehydrated samples of leaves near the leaf margins and the median portions of petioles and peduncles were prepared for scanning electron microscopy (SEM) analysis by fixing them to aluminum supports (stubs) using double-sided adhesive tape. The samples were then photomicrographed using a Hitachi SEM, model TM4000 Plus. The images were processed using CorelDRAW® 2021 software.

    thumbnail of Figure 1.
    Figure 1.
    Populations of Ipomoea bahiensis Willd. ex Roem. & Schult. (white) and Ipomoea cryptocarpa Belo & Buril, sp. nov. (red) analyzed in this study. Specimens of Ipomoea bahiensis Willd. ex Roem. & Schult. (green) examined in herbaria.

    Anatomical analyses

    We proceeded with a comparative anatomical analysis using leaves, sepals, and stems of the new species and its closest relative, I. bahiensis (Table 1). Three leaves from the third shoot node, two sepals, and stem samples were obtained from the specimens collected in the field and fixed in FAA 50 (formaldehyde, acetic acid, and 50% ethanol) for 48 hours (Johansen, 1940) and then stored in 70% ethanol. Freehand sections were made in the median region of the leaf blade, petiole, and stem, cleared by 50% sodium hypochlorite treatment, and stained with Safranin-astra blue (Bukatsch, 1972). Slides were prepared according to the protocols of Kraus & Arduin (1997), analyzed under a Leica DM500 microscope, and subsequently deposited in the Integrative Systematics Laboratory at Universidade Federal Rural de Pernambuco, Brazil. The images were processed using CorelDRAW® 2021 software.

    Palynological analysis

    To describe pollen morphology, at least two unopened flowers were collected from each specimen per population, close to anthesis, to obtain a significant sample of pollen material (Table 1). The pollen grains were acetolyzed according to Erdtman (1960) with the modifications mentioned in Melhem et al. (2003). Diameter measurements were taken on 25 pollen grains over one week (Salgado-Labouriau et al., 1965). Other measurements (exine thickness and apertures) were taken from ten pollen grains. The slides obtained were incorporated into the pollen collection of the Laboratory of Plant Morphology and Palynology - LaMPali/UNESP.

    Statistical analyses were performed to obtain the mean (χ) and standard deviation (Sₓ) of diameters, apertures, spines, the distance between spines, and exine layers of pollen grains (Zar, 2010). The pollen morphology and terminology follow Punt et al. (2007) and Halbritter et al. (2018). The number of apertures was calculated based on Hoen & Punt (1989), and to calculate the number of spines, we followed what was proposed by Hanks & Fryxell (1979).

    Thumbnail
    Table 1.
    Specimens used in micromorphological analyses.

    Results

    Anatomy

    In the parademic section and the frontal view, the leaf blade epidermis of Ipomoea sp. nov. comprises cells with sinuous anticline walls on the adaxial and abaxial surfaces, with tector trichomes, and sessile peltate glandular trichomes. The leaf is amphihypostomatic, with anisocytic and paracytic stomata (Fig. 2A-B). In cross-section, the petiole has a concave-convex contour with uniseriate epidermis and absent trichomes (Fig. 2E). Adjacent to the epidermis is found the angular collenchyma with 2-3 layers of cells, arranged continuously along the petiole, followed by the fundamental parenchyma, composed of visually isodiametric cells circular (Fig. 2E). The vascular system is bicollateral comprising three bundles, the main one being the arch, located in the central region of the petiole, and two smaller ones positioned on the adaxial surface (Fig. 2E). Laticiferous canals are concentrated in the region above the main vascular bundle of the petiole (Fig. 2E). The secondary stem has a circular shape and uniseriate epidermis (Fig. 2G). The cortical region of the stem is adjacent to the epidermis and composed of 3 layers of lamellar collenchyma, followed by 2-3 layers of parenchyma. Internally to the cortical region, sclerenchyma fibers were observed surrounding the phloem. In the vascular system in a single growth ring, the xylem forms a continuous ring with xylem elements. The central region of the stem is occupied by a medulla composed of fundamental parenchyma with rounded and isodiametric cells, the largest ones in the central region and the smallest close to the xylem, in addition to the phloem (2-3 layers), and idioblasts containing drusen and laticiferous canals in the central region (Fig. 2I).

    thumbnail of Figure 2.
    Figure 2.
    Paradermic and transverse sections of the leaves and stems of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult. A. Adaxial epidermis with sinuous cell walls, anisocytic and paracytic stomata in I. cryptocarpa: glandular trichome (gl tr), stomata (st). B. Abaxial epidermis with sinuous cell walls and paracytic stomata in I. cryptocarpa: glandular trichome (gl tr), stomata (st). C. Adaxial epidermis with straight to slightly curved walls and paracytic stomata in I. bahiensis: tector trichome (tec tr), stomata (st). D. Abaxial epidermis with straight to curved walls and paracytic stomata in I. bahiensis: glandular trichome (gl tr), tector trichome (tec tr). E. Petiole concave-convex with absent trichomes in I. cryptocarpa. F. Petiole concavo-convex with tector trichomes in I. bahiensis: tector trichome (tr). G. Stem of I. cryptocarpa with few laticiferous canals in the medulla region and xylem. H. Stem of I. bahiensis with many laticiferous canals in the medulla region and xylem. I. Details of the stem of I. cryptocarpa: epidermis (ep), collenchyma (col), laticiferous canals (lc), sclerenchyma (scl), phloem (ph), xylem (xy), vessel elements (ve). J. Details of the stem of I. bahiensis: collenchyma (col), laticiferous canals (lc), sclerenchyma (scl), phloem (ph), xylem (xy). Scale bars: A-D = 100 μm; E-H = 200 μm; I-J = 100 μm. Photographs: Deibson Belo.

    In the paradermal section, the sepals of Ipomoea sp. nov. present straight anticlinal walls on both epidermal surfaces, with the presence of druses on the adaxial surface (Fig. 3C). The stomatal distribution is hypostomatic, with predominantly anomocytic stomata, with anisocytic ones being rare, and occasionally forming stomatal groupings in pairs (Fig. 3D). In cross-section, the sepals have a straight adaxial surface and a prominent abaxial surface, with sharp ends. The epidermis is uniseriate, with glandular trichomes emerging from the surface on both surfaces (Fig. 3F). The mesophyll is homogeneous (Fig. 3F), composed of compact parenchyma cells, without noticeable intercellular spaces, numerous laticiferous canals are present (Fig. 3H), distributed in the central region and along the mesophyll.

    In paradermal section, the epidermal cells of the leaf blade of I. bahiensis exhibit straight to curved anticlinal walls on both surfaces (adaxial and abaxial) (Figs. 2C-D). The stomatal distribution is amphihypostomatic, and paracytic stomata commonly occur on both surfaces of the leaf blade (Figs. 2C-D). In the transverse section, the mesophyll is dorsiventral, with palisade parenchyma arranged in 2 to 3 layers and spongy parenchyma in 3 to 4 layers. The main midrib displays a biconvex outline, angular collenchyma, and a vascular system consisting of a single central bicolateral bundle in an arch shape. The petiole is concave-convex, with an undulated epidermis and a shallow, closed adaxial concavity (Figs. 2F).

    thumbnail of Figure 3.
    Figure 3.
    Comparative anatomy of the sepals of Ipomoea bahiensis Willd. ex Roem. & Schult. and Ipomoea cryptocarpa Belo & Buril. A. Adaxial surface of Ipomoea bahiensis shows no druses and stomatal clusters. B. Abaxial surface of Ipomoea bahiensis. C. Adaxial surface of Ipomoea cryptocarpa showing druses (dr). D. Abaxial surface of Ipomoea cryptocarpa showing stomatal clusters in pairs (st). E. Mesophyll of Ipomoea bahiensis showing aerenchyma below the epidermis (aer). F. Mesophyll of Ipomoea cryptocarpa showing compact homogeneous parenchyma. G. Detail of the abaxial region showing few laticiferous canals near the epidermis of Ipomoea bahiensis. H. Laticiferous canals in the central region of the mesophyll of Ipomoea cryptocarpa.

    Palynology

    The pollen grains of Ipomoea sp. nov. are monads, apolar, large size (x̄ = 90.8 × 90.4 μm) (Fig. 4A), spheroidal, pantoporate, narrow pores (Fig. 4B), ca. 111 circular pores (x̄ = 2.8 μm) annulus absent; exine tectate, echinate, and perforate. Bulbous spines (x̄ = 12.2 μm) (Fig. 4C), ca. 250, rounded apex, wide and polygonal base. Nexine is thicker than sexine (exine total x̄ = 9.6 μm; sexine x̄ = 3.9 μm; nexine x̄ = 5.7 μm).

    In I. bahiensis, the pollen grains are monads, apolar, very large (x̄ = 111.7 × 110.9 μm) (Fig. 4E), spheroidal, pantoporate, large pores (Fig. 4F), ca. 96 circular pores (x̄ = 9.9 μm) annulus absent; exine tectate, echinate, perforate. Coniform spines (x̄ = 20.3 μm) (Fig. 4G), ca. 165, acute apex, wide and rounded base. Sexine is thicker than nexine (exine total x̄ = 10.3 μm; sexine x̄ = 6.3 μm; nexine x̄ = 4.1 μm).

    Here we present a taxonomic treatment of the new species and a complete description of I. bahiensis, clarifying its morphological boundaries, based on populations studied from its entire distribution range.

    thumbnail of Figure 4.
    Figure 4.
    Comparison of the pollen grains of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult. in light micrographs. A. General view of I. cryptocarpa pollen grains. B. Aperture details of I. cryptocarpa pollen grains. C. Spines details of I. cryptocarpa pollen grains. D. Base of spines details of I. cryptocarpa pollen grains. E. General view of I. bahiensis pollen grains. F. Aperture details of I. bahiensis pollen grains. G. Spines details of I. bahiensis pollen grains. H. Base of spines details of I. bahiensis pollen grains. Scale bars: 50 μm. Photographs: Eduardo Gasparino.

    Taxonomic Treatment

    Ipomoea cryptocarpa Belo & Buril, sp. nov. (Figs. 5, 6, 7, 8, 9)

    Type: BRAZIL. Mato Grosso do Sul, Anastácio, entrada da cidade, sentido Campo Grande-Anastácio; 20°29’52” S, 55°47’15” W; 171 m; 2 Sep. 2023 (fl., fr.), E. Barbier & D. Belo 13 (Holotype: PEUFR!; Isotype: CGMS!).

    Diagnosis: Ipomoea cryptocarpa is similar to I. bahiensis due to its hastate leaves, with the base hastate to sagittate, and sepals with an appendage at the apex. However, I. cryptocarpa bears a glabrous petiole (vs. pubescent in I. bahiensis), inflorescence 1-3-flowered (vs. 4-20-flowered), corolla ≤ 30 mm long (vs. > 30 mm long), sepals with a dorsal globose appendage (vs. sepals with a dorsal tooth-like appendage), accrescent, enclosing the capsules (vs. not enclosing the capsules), and seeds lanate (vs. pubescent).

    thumbnail of Figure 5.
    Figure 5.
    Comparison of the macromorphological structures of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult. A. Flower of I. cryptocarpa. B. Flower of I. bahiensis. C. Fruits of I. cryptocarpa. D. Fruits of I. bahiensis. E. Seed of I. cryptocarpa. F. Seeds of I. bahiensis. Photographs: Eder Barbier (A-E); Diego Gonzaga (F).

    thumbnail of Figure 6.
    Figure 6.
    Ipomoea cryptocarpa Belo & Buril, sp. nov. A. Habit. B. Striated stem. C. Trichomes on the adaxial leaf surface. D. Bifid trichomes. E. Striated petiole and glabrous. F. Inflorescence. G. Floral bud. H. Flower. I. Bracteole. J. Sepals with dorsal globose appendage. K. Corolla inner surface. L. Stamen. M. Gynoecium. N. Fruit immature. Illustration of the holotype by Regina Carvalho.

    thumbnail of Figure 7.
    Figure 7.
    Comparison of the morphological structures of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult. A. Adaxial surface glabrous in I. cryptocarpa. B. Adaxial surface pubescent in I. bahiensis. C. Petiole glabrous in I. cryptocarpa. D. Petiole pubescent in I. bahiensis. E. Peduncle glabrous in I. cryptocarpa. F. Peduncle pubescent in I. bahiensis. Photographs: Deibson Belo.

    thumbnail of Figure 8.
    Figure 8.
    Comparison of the morphological structures of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult. view by scanning electron microscopy. A. Adaxial surface with sessile peltate glandular trichomes (gl tr) in I. cryptocarpa. B. Adaxial surface pubescent with tector trichomes (tec tr) and peltate glandular trichomes (gl tr) in I. bahiensis. C. Petiole with sessile peltate glandular trichomes (gl tr) in I. cryptocarpa. D. Petiole pubescent with tector trichomes (tec tr) and peltate glandular trichomes (gl tr) in I. bahiensis. E. Peduncle with sessile peltate glandular trichomes (gl tr) in I. cryptocarpa. F. Peduncle pubescent with tector trichomes (tec tr) and peltate glandular trichomes (gl tr) in I. bahiensis. Photographs: Hianna Fagundes.

    thumbnail of Figure 9.
    Figure 9.
    Comparison of the morphological structures of Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis. A. Sepals with dorsal globose appendage in I. cryptocarpa. B. Sepals with dorsal tooth-like appendage in I. bahiensis. C. Bracteoles (brac) lanceolate with the apex acute in I. cryptocarpa. D. Bracteoles (brac) lanceolate with the apex acuminate to caudate in I. bahiensis. Photographs: Deibson Belo.

    Description: Climbing plants, herbaceous; twining, branches glabrous, striated; internodes 35.6-141.2 mm long. Leaf blades 21.6-45.6 × 7.3-14.9 mm, hastate to sagittate, margin entire, ciliate, the base hastate to sagittate, the apex caudate, rare mucronate to acute, adaxial surface glabrous to glabrescent with tector trichomes simple or bifid, and sessile peltate glandular trichomes, abaxial surface glabrescent with tector trichomes simple, and sessile peltate glandular trichomes viewed under scanning electron microscopy, venation peninervea, cladodromous type, with eight to ten pairs of secondary veins; petiole 8.8-31.2 mm long, glabrous to glabrescent, sessile peltate glandular trichomes viewed under scanning electron microscopy, striated with protuberances. Inflorescence cymose 1-3-flowers; peduncle 3.3-7.2 mm long, glabrous, striated viewed under stereomicroscope, and sessile peltate glandular trichomes viewed under scanning electron microscopy; bracteoles 3.7-6.6 mm long, lanceolate, the base rounded, the apex acute, glabrous; pedicels 4.5-10.5 mm long, glabrous, striated. Sepals 5, unequal, fleshy, the 2 outer ones 5.4-7 × 5.5-5.8 mm, ovate, the base rounded to subtruncate, the apex retuse with subtly dorsal globose appendage between 1-1.2 mm long, glabrous, the intermediate one 7-7.6 × 5.2-5.6 mm, rotund, the base rounded to truncate, the apex retuse with dorsal globose appendage ca. 1.2 mm long, glabrous, the 2 inner ones 7.1-7.5 × 5.2-5.8 mm, rotund, the base rounded, the apex retuse with subtly dorsal globose appendage between 1-1.2 mm long, glabrous. Corolla 24.1-28.6 mm long, funnelform, pink, glabrous. Filaments 5, emerging from the base of the corolla, 8.1-15.7 mm long, glabrous; anthers usually narrowly oblong 3-3.5 mm long, glabrous. Ovary 1.1-1.3 × 1.8-1.9 mm, globose; style entire, 6-6.2 mm long, stigmatic lobes 2, 0.5-0.6 mm long, oval. Capsules 4-valvar, 8-10 mm long, globose, enclosed by accrescent sepals. Seeds 4.7-5.6 mm long, lanate, golden.

    Etymology: The specific epithet is derived from the Greek crypto-, covered, hidden, and -carpos, fruit; referring to the fruit being hidden by the sepals until maturity, a diagnostic character of the species.

    Phenology: Ipomoea cryptocarpa was found with flowers and young fruits until early September. Senescent flowers and fruits were also found in loco.

    Distribution and habitat: This species is known only from Anastácio municipality (state of Mato Grosso do Sul), in the Brazilian Cerrado. It was found near a livestock grazing area in a peri-urban transition zone. Ipomoea cryptocarpa is potentially a native plant adapted to anthropic areas. It was observed climbing a species of Poaceae, even though only a few individuals were found in the area.

    Preliminary IUCN conservation assessment: Assessing the conservation status of I. cryptocarpa, we considered it as Data Deficient (DD) since it is known only from the type locality. The species was discovered in an anthropic environment, an urban and peri-urban transition zone of the Brazilian Cerrado, near pastures used for domestic animal farming. These areas experience constant human interference, whether from the upkeep of urban roads or the maintenance of crop and livestock management. The Brazilian Cerrado is one of Brazil’s biodiversity hotspots and the second-largest phytogeographic domain (Klink & Machado, 2005; Hofmann et al., 2021). The number of vascular plants in this phytogeographic domain exceeds 13,900 known species, of which around 33% are endemic (Flora e Funga do Brasil, 2024). A large part of the Cerrado was transformed into pasture and crop areas (Klink & Machado, 2005), and the destruction of this domain continues, mainly due to human action (Machado et al., 2004). Many biological entities that are still undescribed and inhabit the Cerrado may be threatened, due to the countless impactful activities that directly or indirectly affect this domain (Fernandes & Pessôa, 2011).

    Additional specimens examined (Paratypes): Brazil. Mato Grosso do Sul: Anastácio, 20°29’52”S, 55°47’15”W, 2 Sep. 2023 (fl,, fr.), E. Barbier & D. Belo 14 (PEUFR); ibid., 2 Sep. 2023 (fl,), E. Barbier & D. Belo 15 (UFP).

    Taxonomic notes: The sagittate and hastate leaves found in the new species are also found in other species of the genus in areas of the Brazilian Cerrado or close to the type locality of I. cryptocarpa, such as I. acanthocarpa (Choisy) Hochst. ex Schweinf. & Asch., I. aequiloba J.R.I.Wood & Scotland, I. aquatica Forssk., I. bahiensis, I. cryptica J.R.I.Wood & Scotland, I. maurandioides Meisn., I. mucronatoproducta J.R.I.Wood & Scotland, I. paludicola J.R.I.Wood & Scotland, and I. squamosa Choisy (Wood et al., 2020; Delgado-Junior et al., 2023). Among them, only I. cryptocarpa and I. bahiensis share the sepals with appendages and possibly are closely related. However, the absence of indument in the branches, in addition to the shape of the sepal appendage, and the capsule enclosed by the sepal when maturing, are relevant macromorphological characters to differentiate them (Figs. 7, 8, 9, Table 2).

    Thumbnail
    Table 2.
    Morphological comparison between Ipomoea cryptocarpa Belo & Buril, sp. nov. and Ipomoea bahiensis Willd. ex Roem. & Schult.

    Ipomoea bahiensis Willd. ex Roem. & Schult., Syst. Veg. 4: 769. 1819.

    Type: Brazil. T. Hoffmannsegg s.n. (holotype B-W 03753-010).

    Ipomoea salzmannii Choisy, Mém. Soc. Phys. Genève 8(1): 59. 1838. Type. Brazil. Bahia, Salzmann s.n. (lectotype M0184904, designated by Wood et al., 2020:670).

    Ipomoea salzmannii var. uniflora Choisy in A.P. de Candolle, Prodr. 9: 379. 1845. Type. BRAZIL. Minas Gerais, Salgado, Martius s.n. (holotype M0184905). Ipomoea bahiensis var. uniflora (Choisy) Meisn. in Martius et al., Fl. Brasil. 7: 269. 1869.

    Ipomoea bahiensis var. sagittifolia Meisn. in Martius et al., Fl. Brasil. 7: 269. 1869. Type. Brazil. Rio São Francisco, Gardner 1359 (lectotype K000944834, designated by Wood et al., 2020:670).

    Quamoclit rochai Hoehne, Anexos Mem. Inst. Butantan, Bot. 1, fasc. 6: 79. 1922. Type. Brazil. Ceará, D. Rocha 16 (holotype SP).

    Description: Climbing plants, herbaceous; twining, branches glabrescent to pubescent, smooth to striated; internodes 10-82 mm long. Leaf blades 12.5-89.18 × 5.79-95.56 mm, cordate, sagittate to hastate, margin entire, the base cordate, subcordate to hastate, the apex acuminate, apiculate with mucro, aristulate with mucro, mucronate to acute, adaxial surface pubescent, sericeous to tomentose, tector trichomes simple, and peltate glandular trichomes, abaxial surface pubescent, sericeous to tomentose, tector trichomes simple, and peltate glandular trichomes, venation peninervea, camptodromous type, with seven to nine pairs of secondary veins; petiole 3.36-63.45 mm long, pubescent, tomentose to hirsute, tector trichomes simple, and peltate glandular trichomes. Inflorescence cymose 4-20-flowers; peduncle 9.97-130.50 mm long, pubescent, tomentose to hirsute; bracteoles 1.73-6.63 mm long, linear to ensiform, the base truncated, the apex acuminate to caudate, glabrescent; pedicels 2-12.56 mm long, glabrous, smooth. Sepals 5, unequal, the 2 outer ones 3.46-7.22 × 2.07-4.1 mm, oval, ovate, obovate, oblong to elliptic, the base subtruncate, cuneate to aequilateral, the apex acute with mucro, subtly tooth-like appendage between 1.3-2.7 mm long, glabrous to glabrescent, with tector trichomes in the margin, the intermediate one 4.17-7.45 × 3.27-6.46 mm, rotund, the base subtruncate, the apex obtuse with mucro, subtly tooth-like appendage between 1.5-2.1 mm long, glabrous to glabrescent, with tector trichomes in the margin, the 2 inner ones 4.79-8.18 × 4.34-8.19 mm, rotund to orbicular, the base cordate to subcordate, rare subtruncate, the apex retuse to obtuse with mucro, with subtly tooth-like appendage between 1.3-2.5 mm long, glabrous. Corolla 30.14-54 mm long, funnelform, pink, glabrous. Filaments 5, emerging from the base of the corolla, 5.93-21.44 mm long, tector trichomes in the base, anthers elliptic 3.1-5.04 mm long, glabrous. Ovary 1-1.98 × 0.8-1.46 mm, ovoid; style entire, 7.36-17.82 mm long, stigmatic lobes 2, 0.89-1.21 mm long, oval, glabrescent. Capsules 4-valvar, 11-20 mm long, globose; seeds 4.5-5.7 mm long, pubescent, gray to black.

    Distribution and habitat: Ipomoea bahiensis occurs predominantly in Caatinga areas of northeastern Brazil, mainly on forest edges and disturbed environments. There are also records in other areas of Brazil, such as the Eastern Amazon, Cerrado, Atlantic Forest, and Pantanal (Flora e Funga do Brasil, 2024). It does not occur in the southern region of Brazil. After analyzing specimens in K, we also observed records of I. bahiensis in eastern Bolivia (Wood et al., 2015; 2020).

    Specimens examined: BOLIVIA - Santa Cruz. Santa Cruz. Nuflo de Chavez, 14 Mar 2019, J.R.I. Wood et al. 28973 (K barcode 000544433!). Chiquitos, 18 Apr 2013, J.R.I. Wood et al. 27877 (K barcode 001755498!). BRAZIL - Alagoas: Marechal Deodoro, 12 Sep 2008, R.P. Lyra-Lemos et al. 11512 (MAC barcode 034411!). Murici, margens da Estação Ecológica de Murici, 15 Nov 2023, D. Belo and E. Barbier 638 (PEUFR). Bahia: Andaraí, 17 Apr 2024, D. Belo et al. 654 (PEUFR). Juazeiro, 21 May 2023, D. Belo and E. Barbier 621 (PEUFR). Morro do Chapéu, 18 Apr 2024, D. Belo et al. 656 (PEUFR). Ceará: Juazeiro do Norte, 27 May 2023, D. Belo and E. Barbier 630 (PEUFR). Missão Velha, 28 May 2023, D. Belo and E. Barbier 631 (PEUFR). Goiás: Cavalcante, 14 Apr 2001, G. Pereira-Silva et al. 4913 (CEN barcode 00042107!, MAC 12046!, SP 370075!). Maranhão: Carolina, 24 May 1950, J.M. Pires et al. 2032 (IAN 58462!). Mato Grosso: Cáceres, 21 Jul 1987, W.L. Werneck 64 (CEN barcode 00011380). Novo Mundo Parque Estadual do Rio Cristalino, 9°26’28”S, 55°48’56”W, 1 Jun 2007, D. Sasaki et al. 1694 (INPA 222983!). Mato Grosso do Sul: Aquidauana, 12 Sep 2004, A. Pott 11798 (CGMS 52986!, PEUFR 53365!). Bonito, 4 Dec 2010, A. Quinet et al. 2354 (RB, SP). Pará: Belém, 13 May 1997, A. Nitta 17036 (IAN 167912!). Bragança, E.S. Galvão-Júnior 17 (MG barcode 160172!). Rurópolis, 17 May 2024, D. Belo and E. Barbier 25 (PEUFR). Paraíba: Cajazeiras, 3 Aug 2023, D. Belo and E. Barbier 636 (PEUFR). Pernambuco: Buíque, Parque Nacional do Catimbau, 29 July 2023, D. Belo and E. Barbier 633 (PEUFR). Pesqueira, 30 July 2023, D. Belo and E. Barbier 634 (PEUFR). Triunfo, 28 July 2023, D. Belo and E. Barbier 632 (PEUFR). Piauí: São Raimundo Nonato, Parque Nacional Serra da Capivara, 25 May 2023, D. Belo and E. Barbier 628 (PEUFR). Tocantins: Pedro Afonso, 28 Apr 1999, K.G. Kissmann s.n. (MBM barcode 248689!).

    Discussion

    Ipomoea cryptocarpa is, to date, known only from its type locality, an anthropic area within the Cerrado domain, in the state of Mato Grosso do Sul. Although we are currently conducting a review of I. bahiensis, its most closely related species, we have not detected any specimens that fit its concept in the herbarium collections. However, there may be other populations that have not been collected due to the morphological similarity to a widespread, very common species.

    Several authors have been considering I. bahiensis as a morphologically hypervariable species (Simão-Bianchini, 1998; Wood et al., 2020; Delgado-Junior et al., 2023). Thus, proposing new species related to it can be tricky. Starting from the hypothesis that the current concept of I. bahiensis is hiding a greater diversity, the ongoing thesis project of the first author aims to evaluate this variability through replicable methods, using different sources of information. After analyzing more than 20 populations and approximately 400 samples, our preliminary results corroborate this hypothesis so far. We can observe discontinuities between morphotypes (Belo & Buril, unpublished data). The description of I. cryptocarpa is the first result of our analysis since many characters support the idea that it does not belong to the typical I. bahiensis.

    Fifty-two species of Ipomoea are found in Mato Grosso do Sul state, but only two of them, I. cardenasiana O'Donell and I. pantanalensis J.R.I.Wood & Urbanetz, are endemic to the state (Wood et al., 2020; Simão-Bianchini et al., 2024). As in I. cryptocarpa, only four other species of Ipomoea that occur in Mato Grosso do Sul have fruits enclosed by accrescent sepals: I. cardenasiana, I. revoluta J.R.I. Wood & Scotland, I. rubens Choisy, and I. setifera Poir. (see Wood et al., 2020; Simão-Bianchini et al., 2024).

    However, I. cryptocarpa has a glabrous stem (vs. glabrescent in I. cardenasiana, pubescent in I. revoluta, tomentose in I. rubens, in hirsute in I. setifera), leaves entire, hastate, glabrescent in both surfaces (vs. 3-lobed, glabrous in surface adaxial, pubescent in surface abaxial in I. cardenasiana, digitate, 5-7 free leaflets, glabrous in surface adaxial, tomentose in surface abaxial in I. revoluta, ovate-deltoid, pubescent in surface adaxial, tomentose in surface abaxial in I. rubens, ovate-deltoid, glabrous in both surfaces in I. setifera), the base hastate to sagittate (vs. cordate in I. cardenasiana and I. rubens, tapered in I. rubens, cordate to hastate in I. setifera), the apex mucronate to acute (vs. acuminate and mucronate in I. cardenasiana, mucronate in I. revoluta, acute in I. rubens, obtuse to mucronate in I. setifera).

    Comparing the anatomical characters, the angular-type collenchyma and collateral vascular system in the petiole are shared by I. cryptocarpa and I. bahiensis. However, the sinuous walls on the adaxial and abaxial surfaces in I. cryptocarpa differentiate these two species (vs. straight to curved walls on the adaxial and abaxial surfaces in all populations of I. bahiensis in this study) (Figs. 2A-D). The distribution of stomata on the leaf blade is different in the two species. In I. cryptocarpa the leaves are amphihypostomatic with anisocytic and paracytic stomata (Figs. 2A-B), while in I. bahiensis it is amphistomatic with only paracytic stomata (Figs. 2C-D). The unique presence of paracytic stomata in all populations of I. bahiensis was also observed by EV Santos (unpubl. res.) when comparing four species of Ipomoea. In I. cryptocarpa, the laticiferous canals in the petiole occur in greater quantity and more frequently in the region above the main vascular bundle. We observed a frequency of nine laticiferous canals in the analyzed individuals of I. cryptocarpa (Fig. 2E), a characteristic not observed in the I. bahiensis populations from this study (Fig. 2F). When we compare the structure of the stem, both species share sclerenchyma fibers surrounding the phloem; however lamellar collenchyma with 3-layers occurs in I. cryptocarpa (Figs. 2G, 2I) (vs. angular 2-3-layers in all populations of I. bahiensis). Leaf anatomical characters used to support delimitations such as the type of epidermal walls, mesophyll, and shape of the petiole and midrib, in addition to the stem, have also been considered relevant in other studies of Convolvulaceae (Alencar et al., 2022; Belo et al., 2023b; c), as in other families - e.g. in Lamiaceae (Rashid & Parnell, 2017), and Oxalidaceae (Richetti et al., 2023).

    When comparing the anatomy of outer sepals, we also observed structural differences between the two species. The adaxial epidermis of I. cryptocarpa presents druses (Fig. 3C), predominantly anomocytic stomata, and stomatal groupings in pairs (Fig. 3D), contrasting with I. bahiensis, which presents paracytic stomata and does not present druses or stomatal groupings in the epidermis ((Figs. 3A-(B). In cross-section, the adaxial surface of I. cryptocarpa is rectilinear, with a prominent abaxial surface with absent lobes, while I. bahiensis has a curvilinear adaxial surface and three conspicuous lobes. In addition, I. bahiensis presents aerenchyma in its mesophyll (Fig. 3F), while in I. cryptocarpa the mesophyll is homogeneous and compact (Fig. 3E). Besides that laticiferous canals in I. cryptocarpa sepals are numerous and are predominantly found in the central region along the mesophyll (Fig. 3H), while in I. bahiensis, these canals are restricted to the proximity of the epidermis and in smaller numbers (Fig. 3G).

    The pollen grains of I. cryptocarpa fit into the characteristic pattern of the genus: echinate and pantoporate pollen grains (Sengupta, 1972; Wood et al., 2020). They are large (Fig. 4A), small pores (Fig. 4B), with bulbous spines (Fig. 4C), rounded apex, wide and polygonal base (Fig. 4D), and nexine thicker than sexine. When we compare Ipomoea cryptocarpa with I. bahiensis, we observe a variation in pollen and aperture size, the shape of the spines, and the thickness of the exine layers. The pollen results found in I. bahiensis in the present study, corroborate the description of I. bahiensis provided by Vital et al. (2008), and Vasconcelos et al. (2015), when studying other populations of this variable species.

    Bulbous spines on pollen grains have been previously described in other species of the genus. Sengupta (1972) cites spines as broad and bulbous at the base for the species I. clausa Rudolph ex Ledeb & Adlerstam, I. indica(Burm.) Merr., gradually tapering towards the apex with a blunt tip and constricted neck, characteristics very similar to the spines observed for I. cryptocarpa. Vasconcelos et al. (2015) observed pollen grains with bulbous spines, similar to those described here, for I. hederifolia L. and I. incarnata Choisy. Tiwari et al. (2023) proposed a Type III - wide bulbous base, progressively tapering toward the apex with a blunt tip for the species I. purpurea (L.) Roth and I. triloba. Bulbous spines were also found by Romeiro et al. (2023) for the species I. cavalcantei D.F.Austin, I. decora Meisn., I. goyazensis Gardner, I. hederifolia, and I. marabaensis D.F.Austin & Secco.

    Sengupta (1972) cited this spine morphology as spines broad at the base and gradually tapering towards the apex with more or less rounded tips. Coniform spines such as those observed in I. bahiensis were described for I. procumbens Mart. ex Choisy, I. rosea Choisy, I. rupestris Sim.-Bianch. & Pirani, I. subincana (Choisy) Meisn., and I. maurandioides Meisn. (Vasconcelos et al., 2015). Tiwari et al. (2023) described spines wide at the base, progressively tapering toward the apex with a blunt tip (Type II) for the species I. aitonii Lindl., I. carnea Jacq., I. eriocarpa R.Br., I. hederifolia, and I. nil; Romeiro et al. (2023) observed coniform spines in I. asplundii O'Donell, I. carajasensis D.F.Austin, I. procumbens, and I. setifera Poir. pollen grains. Wood et al. (2020) indicate that there is continuous variation in pollen morphology with little or no variation that correlates with the phylogeny of Ipomoea. However, despite pollen morphology being homogeneous within the genus, our studies indicate that population studies have the potential to differentiate morphologically related specimens.

    Species considered ruderal may be hiding a greater diversity of species, and more detailed studies are needed to understand the morphological patterns and species boundaries. Greater collection efforts are needed for verification of the conservation status of I. cryptocarpa since current information is still considered data deficient. Many weedy species are studied only locally, which can lead to the common practice of putting all “similar ones” within its taxonomic concept. This can bias the diversity knowledge and hamper conservation policies and bioprospecting. We also emphasize that it is necessary to advance in phylogenetic and population genetic studies to better elucidate the diversity in Ipomoea.

    Acknowledgments

    DPB thanks the Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco (FACEPE-IBPG-1957-2.03/22) for the PhD scholarship. To the Laboratório Avançado de Microscopia e Imagem, Departamento de Zoologia/UFPE, process FACEPE-APQ-0522/2.04/19 for providing the SEM images. We also thank Dr. Edinalva Santos for their assistance with anatomical analyses, and Dr. Eder Barbier for the partnership in the botanical expeditions. This research is part of the PhD thesis of the first author at the Programa de Pós-Graduação em Biodiversidade (PPGBio/UFRPE).

    References

    • Alencar J, Nascimento ALB, Duarte-Neto P, Buril MT. 2020. Disentangling leaf morphology in Daustinia montana (Convolvulaceae). Systematic Botany 45: 921-930. doi: 10.1600/036364420X16033962925132.
      » https://doi.org/10.1600/036364420X16033962925132
    • Alencar J, Alves JV, Arruda E, Buril MT. 2022. Population-level analysis of leaf anatomy of Daustinia montana (Moric.) Buril & A.R. Simões: A polymorphic species with tangled taxonomic history. Brazilian Journal of Botany 45: 1279-1295. doi: 10.1007/s40415-022-00836-x
      » https://doi.org/10.1007/s40415-022-00836-x
    • Alencar J, Maciel JR, Buril MT. 2024. Morphologically hypervariable species hinder our knowledge of biodiversity: Daustinia montana (Convolvulaceae) as a case study. Botanical Journal of the Linnean Society 204: 86-101. doi: 10.1093/botlinnean/boad040.
      » https://doi.org/10.1093/botlinnean/boad040
    • Belo DP, Buril MT, Arruda E, Louzada RB. 2023a. A new Jacquemontia Choisy (Convolvulaceae) species from the Brazilian Amazon forest. Acta Amazonica 53: 302-309. doi: 10.1590/1809-4392202300341.
      » https://doi.org/10.1590/1809-4392202300341
    • Belo DP, Buril MT, Arruda E, Louzada RB. 2023b. Disentangling the identity of two Jacquemontia Choisy (Convolvulaceae Juss.) species using an integrative approach. Brazilian Journal of Botany 46: 85-101. doi: 10.1007/s40415-023-00872-1.
      » https://doi.org/10.1007/s40415-023-00872-1
    • Belo DP, Buril MT, Santos EAV, Arruda E, Louzada RB. 2023c. Leaf and stem micromorphology of Jacquemontia evolvuloides (Moric.) Meisn. (Convolvulaceae) populations: New insights for taxonomic classification using light and scanning electron microscopy. Microscopy Research and Technique 86: 1177-1196. doi: 10.1002/jemt.24391.
      » https://doi.org/10.1002/jemt.24391
    • Belo DP, Louzada RB, Buril MT. 2024a. Alpha-taxonomy underestimates the diversity in Jacquemontia (Convolvulaceae): A new hypothesis for Jacquemontia evolvuloides complex based on morphological, anatomical, and morphometric analyses. Plant Biosystems 158: 613-631. doi: 10.1080/11263504.2024.2347853.
      » https://doi.org/10.1080/11263504.2024.2347853
    • Belo DP, Santos EAV, Gasparino EC. 2024b. Ipomoea buriliae (Convolvulaceae), a new species of morning glory discovered in the Borborema Plateau, northeastern Brazil. Rheedea 34: 488-505. doi: 10.22244/rheedea.2024.34.05.10.
      » https://doi.org/10.22244/rheedea.2024.34.05.10
    • Bukatsch F. 1972. Bemerkungen zur doppelfarbung. Microkosmos 61: 1-255.
    • Delgado-Junior GC, Costa SL, Staples G, Buril MT. 2023. Flora of Pernambuco, Brazil: Ipomoea (Convolvulaceae). Rodriguésia 74: e01152020. doi: 10.1590/2175-7860202374009.
      » https://doi.org/10.1590/2175-7860202374009
    • Erdtman G. 1960. The acetolysis method. A revised description. Svensk Botanisk Tidskrift 54: 561-564.
    • Eserman L, Sosef MSM, Simão-Bianchini R et al 2020. Proposal to change the conserved type of Ipomoea, nom. cons. (Convolvulaceae). Taxon 69: 1369-1371. doi: 10.1002/tax.12400.
      » https://doi.org/10.1002/tax.12400
    • Fernandes PA, Pessôa VLS. 2011. O Cerrado e Suas Atividades Impactantes: Uma Leitura Sobre o Garimpo, a Mineração e a Agricultura Mecanizada. Observatorium: Revista Eletrônica de Geografia 3: 19-3.
    • Flora e Funga do Brasil. 2024. Jardim Botânico do Rio de Janeiro. https://floradobrasil.jbrj.gov.br/FB93 01 Nov. 2024.
      » https://floradobrasil.jbrj.gov.br/FB93
    • Halbritter H, Ulrich S, Grímsson F et al 2018. Illustrated Pollen Terminology. 2nd. edn. Cham, Springer International Publishing.
    • Hanks S, Fryxell PA. 1979. Palynological studies of Gaya and Herissantia (Malvaceae). American Journal of Botany 66: 494-501. doi: 10.1002/j.1537-2197.1979.tb06251.x.
      » https://doi.org/10.1002/j.1537-2197.1979.tb06251.x
    • Harris JG, Harris MW. 2001. Plant identification terminology: An illustrated glossary. 2nd. edn. Utah, Spring Lake Publishing.
    • Hoen PP, Punt W. 1989. Pollen morphology of the tribe Dorstenieae (Moraceae). Review of Palaeobotany and Palynology 57: 187-220.
    • Hofmann GS, Cardoso MF, Alves RJV et al 2021. The Brazilian Cerrado is becoming hotter and drier. Global Change Biology 27: 4060-4073. doi: 10.1111/gcb.15712.
      » https://doi.org/10.1111/gcb.15712
    • IUCN - International Union for Conservation of Nature. 2012. Red list categories and criteria. Version 3.1. 2nd edition. Prepared by the IUCN Species Survival Commission, Gland and Cambridge. https://www.iucnredlist.org/ 28 Oct. 2024.
      » https://www.iucnredlist.org/
    • IUCN - International Union for Conservation of Nature. 2024. Guidelines for Using the IUCN Red List Categories and Criteria. Version 16. Prepared by the Standards and Petitions Committee. https://www.iucnredlist.org/resources/redlistguidelines 28 Oct. 2024.
      » https://www.iucnredlist.org/resources/redlistguidelines
    • Johansen DA. 1940. Plant microtechnique. New York, McGraw-Hill Book Company.
    • Klink CA, Machado RB. 2005. Conservation of the Brazilian Cerrado. Conservation Biology 19: 707-713. doi: 10.1111/j.1523-1739.2005.00702.x.
      » https://doi.org/10.1111/j.1523-1739.2005.00702.x
    • Kraus JE, Arduin M. 1997. Manual básico de métodos em morfologia vegetal. Rio de Janeiro, Edur.
    • Machado RB, Ramos Neto MB, Pereira P et al 2004. Estimativas de perda da área do Cerrado brasileiro. Relatório técnico não publicado. Brasília, Conservação Internacional. https://cmbbc.cpac.embrapa.br/RelatDesmatamCerrado%20CIBrasil%20JUL2004.pdf 26 Feb. 2024.
      » https://cmbbc.cpac.embrapa.br/RelatDesmatamCerrado%20CIBrasil%20JUL2004.pdf
    • Magalhães HF, Queiroz RT, Cruz DD, Lucena RFP. 2022. Ipomoea bahiensis Willd. ex Roem. & Schult. Convolvulaceae. In: Lucena, RFP, Cruz, DD (eds.). Ethnobotany of the Mountain Regions of Brazil. Cham, Springer International Publishing . p. 455-459.
    • Meira M, Silva EP, David JM, David JP. 2012. Review of the genus Ipomoea: Traditional uses, chemistry and biological activities. Revista Brasileira de Farmacognosia 22: 682-713. doi: 10.1590/S0102-695X2012005000025.
      » https://doi.org/10.1590/S0102-695X2012005000025
    • Melhem TS, Cruz-Barros MAV, Corrêa AS, Makino-Watanabe H, Silvestre-Capelato MSF, Gonçalves-Esteves VL. 2003. Variabilidade polínica em plantas de Campos do Jordão, São Paulo, Brazil. Boletim do Instituto de Botânica de São Paulo 16: 10-104.
    • Miller RE, Rausher MD, Manos PS. 1999. Phylogenetic systematics of Ipomoea (Convolvulaceae) based on ITS and waxy sequences. Systematic Botany 24: 209-227. doi: 10.2307/2419549.
      » https://doi.org/10.2307/2419549
    • Muñoz-Rodríguez P, Carruthers T, Wood JRI et al 2019. A taxonomic monograph of Ipomoea integrated across phylogenetic scales. Nature Plants 5: 1136-1144. doi: 10.1038/s41477-019-0535-4.
      » https://doi.org/10.1038/s41477-019-0535-4
    • Muñoz-Rodríguez P, Wood JRI, Wells T, Carruthers T, Sumadijaya A, Scotland RW. 2023. The challenges of classifying big genera such as Ipomoea Taxon 72: 1201-1215. doi: 10.1002/tax.12887
      » https://doi.org/10.1002/tax.12887
    • Pastore M, Simão-Bianchini R. 2016. Jacquemontia aequisepala (Convolvulaceae), a new species from Brazil. Kew Bulletin 71:26. doi: 10.1007/s12225-016-9640-y.
      » https://doi.org/10.1007/s12225-016-9640-y
    • Pisuttimarn P, Simões ARG, Petrongari FS et al 2023. Distimake vitifolius (Convolvulaceae): Reclassification of a widespread species in view of phylogenetics and convergent pollen evolution. Botanical Journal of the Linnean Society 202: 363-388. doi: 10.1093/botlinnean/boac077.
      » https://doi.org/10.1093/botlinnean/boac077
    • Punt W, Hoen PP, Blackmore S, Nilsson S, Le Thomas A. 2007. Glossary of pollen and spore terminology. Review of Palaeobotany and Palynology 143: 1-81. doi: 10.1016/j.revpalbo.2006.06.008.
      » https://doi.org/10.1016/j.revpalbo.2006.06.008
    • QGIS.ORG. 2023. QGIS geographic information system. QGIS association. https://www.qgis.org 08 Nov. 2024.
      » https://www.qgis.org
    • Rashid MH, Parnell J. 2017. Petiolar anatomical characters and its taxonomic significance in some species of Premna L. (Lamiaceae). Pleione 11: 405-419. doi: 10.26679/pleione.11.2.2017.405-419.
      » https://doi.org/10.26679/pleione.11.2.2017.405-419
    • Richetti, E., Lima, DF., Fiaschi, P., Lusa, MG. 2023. Macro- and micromorphology reveal four entities in the highly variable Oxalis polymorpha Mart. ex Zucc. (Oxalidaceae). Botany 101: 43-60. doi: 10.1139/cjb-2022-0073.
      » https://doi.org/10.1139/cjb-2022-0073
    • Romeiro LA, Silva EF, Vasconcelos LV, Lopes KS, Carreira LMM, Guimarães JTF. 2023. Pollen morphology of Convolvulaceae from Southeastern Amazonian Cangas and its relevance for interaction networks and paleoenvironmental studies. Plants 12: 2256. doi: 10.3390/plants12122256.
      » https://doi.org/10.3390/plants12122256
    • Salgado-Labouriau ML, Vanzolini PE, Melhem TS. 1965. Variation of polar axes and equatorial diameters in pollen grains of two species of Cassia Grana 6: 98-105. doi: 10.1080/00173136509429142.
      » https://doi.org/10.1080/00173136509429142
    • Santos D, Alencar J, Loiola MIB, Buril MT. 2020a. Ipomoea bonsai (Convolvulaceae), a magnificent new species from the Caatinga domain, Brazil. Systematic Botany 45: 652-657. doi: 10.1600/036364420X15935295449907.
      » https://doi.org/10.1600/036364420X15935295449907
    • Santos D, Buril MT. 2024. A new species of Evolvulus (Cresseae-Convolvulaceae) from the Brazilian savanna. Brittonia 76: 253-258. doi: 10.1007/s12228-023-09767-7.
      » https://doi.org/10.1007/s12228-023-09767-7
    • Santos EAV, Leite AV, Arruda ECP. 2024. Anatomical features of ecological importance and taxonomic value revealed by scanning electron microscopy and light microscopy in Camonea umbellata (L.) A.R. Simões & Staples (Convolvulaceae). Microscopy Research and Technique 87: 1889-1903. doi: 10.1002/jemt.24554.
      » https://doi.org/10.1002/jemt.24554
    • Santos D, Saraiva RVC, Ferraz TM, Arruda ECP, Buril MT. 2020b. A threatened new species of Ipomoea (Convolvulaceae) from the Brazilian Cerrado revealed by morpho-anatomical analysis. PhytoKeys 151: 93-106. doi: 10.3897/phytokeys.151.49833.
      » https://doi.org/10.3897/phytokeys.151.49833
    • Santos D, Souza EB, Buril MT. 2021. Ipomoea lanifolia sp. nov. (Convolvulaceae), a new species endemic to the Ibiapaba plateau in northeastern Brazil. Rodriguésia 72: e00782020. doi: 10.1590/2175-7860202172076.
      » https://doi.org/10.1590/2175-7860202172076
    • Sengupta S. 1972. On the pollen morphology of Convolvulaceae with special reference to taxonomy. Review of Palaeobotany and Palynology 13: 157-212. doi: 10.1016/0034-6667(72)90030-9.
      » https://doi.org/10.1016/0034-6667(72)90030-9
    • Simão-Bianchini R. 1998. Ipomoea L. (Convolvulaceae) no Sudeste do Brasil. PhD Thesis, Universidade de São Paulo, Brasil.
    • Simão-Bianchini R, Ferreira PPA, Vasconcelos LV. 2024. Ipomoea in Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. https://floradobrasil.jbrj.gov.br/FB7021 08 Nov. 2024.
      » https://floradobrasil.jbrj.gov.br/FB7021
    • Staples GW, Brummitt RK. 2007. Convolvulaceae. In: Heywood VH, Brummitt RK, Culham A, Seberg O (eds.). Flowering Plant Families of the World. London, Royal Botanic Gardens, Kew. p. 108-110.
    • Stefanović S, Austin DF, Olmstead RG. 2003. Classification of Convolvulaceae: A Phylogenetic Approach. Systematic Botany 28: 791-806. doi: 10.1043/02-45.1.
      » https://doi.org/10.1043/02-45.1
    • Thiers B. 2024. Index herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s virtual herbarium, New York. https://sweetgum.nybg.org/ih/ 26 Jan. 2024.
      » https://sweetgum.nybg.org/ih/
    • Tiwari P, Rawat R, Negi S, Rawat DS. 2023. Pollen morphology of Ipomoea L. from Garhwal Himalaya, India. Palynology 48. doi: 10.1080/01916122.2023.2264357.
      » https://doi.org/10.1080/01916122.2023.2264357
    • Traiperm P, Chow J, Nopun P, Staples G, Swangpol SC. 2017. Identification among morphologically similar Argyreia (Convolvulaceae) based on leaf anatomy and phenetic analyses. Botanical Studies 58: 25. doi: 10.1186/s40529-017-0178-6.
      » https://doi.org/10.1186/s40529-017-0178-6
    • Vasconcelos LV, Saba MD, Junqueira MER, Simão-Bianchini R. 2015. Morfologia polínica de espécies das tribos Ipomoeeae Hallier f. e Merremieae D.F. Austin (Convolvulaceae) ocorrentes numa região de ecótono do município de Caetité, BA, Brasil. Hoehnea 42: 253-264. doi: 10.1590/2236-8906-37/2014.
      » https://doi.org/10.1590/2236-8906-37/2014
    • Vital MTAB, Santos FAR, Alves M. 2008. Diversidade palinológica das Convolvulaceae do Parque Nacional do Catimbau, Buíque, PE, Brasil. Acta Botanica Brasilica 22: 1163-1171. doi: 10.1590/S0102-33062008000400027.
      » https://doi.org/10.1590/S0102-33062008000400027
    • WCSP - World checklist of selected plant families. 2024. Facilitated by the Royal Botanic Gardens, Kew. https://wcsp.science.kew.org/ 26 Feb. 2024.
      » https://wcsp.science.kew.org/
    • Wood JRI, Carine MA, Harris DJ, Wilkin P, Williams BRM, Scotland RW. 2015. Ipomoea (Convolvulaceae) in Bolivia. Kew Bulletin 70: 31. doi: 10.1007/s12225-015-9592-7
      » https://doi.org/10.1007/s12225-015-9592-7
    • Wood JRI, Urbanetz C, Scotland RW. 2016. Ipomoea pantanalensis, a new species of Ipomoea L. (Convolvulaceae) from the Pantanal, Brazil. Kew Bulletin 71: 6. doi: 10.1007/S12225-016-9619-8.
      » https://doi.org/10.1007/S12225-016-9619-8
    • Wood JRI, Muñoz-Rodríguez P, Williams BRM, Scotland RW. 2020. A foundation monograph of Ipomoea (Convolvulaceae) in the New World. PhytoKeys 143: 1-823. doi: 10.3897/phytokeys.143.32821.
      » https://doi.org/10.3897/phytokeys.143.32821
    • Zar JH. 2010. Biostatistical analysis. New Jersey, Prentice-Hall.
    • Data availability:
      All data supporting the findings of this study are included in the article.

    Supplementary Materials

    The following online material is available for this article:

    S1 -

    S2 -

    Edited by

    • Associate Editor:
      Leandro Giacomin
    • Editor-in-Chief:
      Thais Elias Almeida

    Data availability

    All data supporting the findings of this study are included in the article.

    Publication Dates

    • Publication in this collection
      22 Sept 2025
    • Date of issue
      2025

    History

    • Received
      14 Mar 2024
    • Accepted
      27 May 2025
    Creative Common - by 4.0
    This is an open-access article distributed under the terms of the Creative Commons Attribution License
    vertical_align_top show_chart
    location_on
    Sociedade Botânica do Brasil SCLN 307 - Bloco B - Sala 218 - Ed. Constrol Center Asa Norte CEP: 70746-520 Brasília/DF. - Alta Floresta - MT - Brazil
    E-mail: acta@botanica.org.br
    rss_feed Acompanhe os números deste periódico no seu leitor de RSS
    Reportar erro