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A new species of Mandevilla (Apocynaceae) from limestone outcrops in Central Brazil with a short note on the occurrence of geophytes in rauvolfioids and apocynoids

ABSTRACT

Mandevilla calcicola, endemic to limestone formations in Goiás state, Brazil, is described. A key and photos for all Mandevilla species reported in Goiás and Distrito Federal, Brazil, are included. This new species is assessed as Critically Endangered (CR). A short note on the occurrence of geophytes in rauvolfioids and apocynoids (focused on Mandevilla) and its taxonomic value is included.

Keywords:
apocynoids; geophytes; Mesechiteae; tubers; tuberous roots

Introduction

Brazil is floristically the most diverse country in South America, with ca. 42,850 plant species recorded (Flora e Funga do Brasil 2020Flora e Funga do Brasil. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/. 23 Jan. 2023.
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). Providing an accurate estimate of the total number of species in the country is a complex task due to the large amount of unidentified material as well as the taxonomic changes proposed every year. As a result of the exploration of remote areas, many new species and genera were described during the second half of the 20th century. At the same time, however, many species have been relegated to synonymy (Brazil Flora Group 2022Brazil Flora Group. 2022. Brazilian Flora 2020. Leveraging the power of a collaborative scientific network. Taxon 71: 178-198.). Phylogenetic studies provide critical results, which could directly affect the species number in a specific group. The state of Goiás in midwestern Brazil, has a remarkable diversity of habitats dominated by Cerrado with its distinct phytophysiognomic classifications (Ribeiro & Walter 2008Ribeiro JF, Walter BMT. 2008. As principais fitofisionomias do bioma Cerrado. In: Sano SM, Almeida SP, Ribeiro JF (eds.). Cerrado: Ecologia e flora. Brasília, Embrapa. p. 151-212.; Santos et al. 2011Santos TRR dos, Pinto JRR, Lenza E, Mews HA. 2011. The tree-shrub vegetation in rocky outcrop cerrado areas in Goiás State, Brazil. Revista Brasileira de Botânica 35: 281-294.; Soares et al. 2015Soares MP, Reys P, Pifano DS, de Sá JL, da Silva P, Santos TM, Silva FG. 2015. Relationships between edaphic factors and vegetation in savannas on the Brazilian midwest region. Rodriguezia 62: 123-137.). Cerrado occurs predominantly on latosol and quartz sand soils as well on dystrophic and mesotrophic soils (Marimon Junior & Haridasan 2005Marimon Junior BH, Haridasan M. 2005. Comparação da vegetação arbórea e características edáficas de um cerradão e cerrado sensu stricto em áreas adjacentes sobre solo distrófico no leste de Mato Grosso, Brasil. Acta Botanica Brasilica 19: 913-926.). Human activities, such as growing field crops (e.g., sugar cane, corn, soy) and cattle ranching, highly threaten this biome. Considerable knowledge has been obtained on Cerrado vegetation in the central region of Goiás, including the Federal District (e.g., Felfili & Fagg 2007Felfili JM, Fagg CW. 2007. Floristic composition and structure of the “cerrado” sensu stricto on rocky soils in northern Goiás and southern Tocantins, Goiás. Brazilian Journal of Botany 30: 375-385.; Santos et al. 2011Santos TRR dos, Pinto JRR, Lenza E, Mews HA. 2011. The tree-shrub vegetation in rocky outcrop cerrado areas in Goiás State, Brazil. Revista Brasileira de Botânica 35: 281-294.; Lenza et al. 2011Lenza E, Pinto JRR, Pinto AS, Maracahipes L, Bruziguessi EL. 2011. Comparação da vegetação arbustivo-arbórea de uma área de cerrado rupestre na Chapada dos Veadeiros, Goiás e áreas de cerrado sentido restrito do Bioma Cerrado. Revista Brasileira de Botânica 34: 247-259.), but many regions in southeastern and southern Goiás remain unexplored.

Apocynaceae comprises 96 genera and ca. 980 species in Brazil. The family is well represented in Goiás (including the Distrito Federal) (Flora e Funga do Brasil 2020Flora e Funga do Brasil. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/. 23 Jan. 2023.
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). With 43 genera and 143 species recorded, it is the ninth richest Brazilian state in Apocynaceae, with the most diverse genera being Mandevilla Lindl. (20 species), Aspidosperma Mart. & Zucc. (15 species), Oxypetalum R. Br. (13 species), Ruehssia H. Karst. (nine species), and Odontadenia Benth. (seven species) (Flora e Funga do Brasil 2020Flora e Funga do Brasil. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/. 23 Jan. 2023.
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). Mandevilla (Mesechiteae) is one of the largest genera of Apocynaceae in the New World, with an estimated 200 species (Morales & Morais 2020Morales JF, Morais IL 2020. Studies in the Neotropical Apocynaceae LV: A new Mandevilla from Bahia, Brazil, with notes on the diversity of the genus. Systematic Botany 45: 183-189.; Morales & Kollmann 2020aMorales JF, Kollmann L. 2020a. Increasing the known floristic diversity of Brazilian inselbergs: two new species of Mandevilla (Apocynaceae) from Espírito Santo. Acta Botanica Brasilica 34: 107-116.; bMorales JF, Kollmann L. 2020b. War, types, and Mandevilla (Apocynaceae): A new species and the rediscovery of a rare species from Brazil. Darwiniana, nueva serie 8: 449-459.; Morales et al. 2022Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093.), distributed from southern USA and the West Indies to northern Argentina, except in Chile (Morales 2013Morales JF. 2013. Estudios en las Apocynaceae Neotropicales XLIX: Sinopsis de las Apocynaceae (Apocynoideae, Rauvolfioideae) de Chile. Darwiniana, nueva serie 1: 39-45.; Alvarado-Cárdenas & Morales 2014Alvarado-Cárdenas LO, Morales JF. 2014. El género Mandevilla (Apocynaceae: Apocynoideae, Mesechiteae) en México. Botanical Sciences 92: 59-79.). The genus exhibits a broad range of growth forms, enabling it to thrive in a variety of habitats, from coastal dunes (Restingas) and white-sand formations in the Amazon basin to the Andean forest and campos rupestres in Brazil, as well as dry forest, swamp areas, and savannas (Morales 2007Morales JF. 2007. Estudios en las Apocynaceae Neotropicales XXX: Tres nuevas especies andinas de Mandevilla (Apocynoideae, Mesechiteae). Journal of the Botanical Research Institute of Texas 1: 853-857.; 2009Morales JF. 2009. Estudios en las Apocynaceae Neotropicales XXXVIII: Tres nuevas especies de Mandevilla (Apocynoideae, Mesechiteae) para Colombia y Venezuela. Journal of Botanical Research Institute of Texas 3: 565-571.; 2011Morales JF. 2011. Estudios en las Apocynaceae Neotropicales XLII: sinopsis del género Mandevilla (Apocynoideae: Mesechiteae) en Colombia. Journal of the Botanical Research Institute of Texas 5: 521-543.). White-sand savannas are rich in endemic plants (Adeney et al. 2016Adeney JM, Christensen NL, Vicentini A, Cohn-Haft M. 2016. White‐sand Ecosystems in Amazonia. Biotropica 48: 7-23.).

Mandevilla is highly diverse in Brazil. Woodson (1933)Woodson RE Jr. 1933. Studies in the Apocynaceae IV. The American genera of Echitoideae. Annals of the Missouri Botanical Garden 20: 605-790. recorded 36 species for the country, of which 27 were endemic. Since then, the number of species has increased substantially. Sales (1993)Sales MF. 1993. Estudos taxonômicos de Mandevilla Lindley subgênero Mandevilla (Apocynaceae) no Brasil. - Campinas. PhD Thesis, Universidade Estadual de Campinas, Brazil. recognized 40 species of Mandevilla subgenus Mandevilla (sensu Woodson 1933Woodson RE Jr. 1933. Studies in the Apocynaceae IV. The American genera of Echitoideae. Annals of the Missouri Botanical Garden 20: 605-790. ). Fifteen new species have been published for Brazil in the last 15 years, almost all of them endemic (Morales 2005aMorales JF. 2005a. Estudios en las Apocynaceae neotropicales XVIII: Dos nuevas especies de Mandevilla (Apocynoideae, Mesechiteae) para Brasil. Darwiniana, nueva serie 43: 84-89. ; bMorales JF. 2005b. Estudios en las Apocynaceae neotropicales XI: Una nueva especie de Mandevilla (Apocynoideae: Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay. Sida 21: 1549-1555.; Sales et al. 2006Sales MF, Kinoshita LS, Simões AO. 2006. Eight new species of Mandevilla Lindley (Apocynaceae, Apocynoideae) from Brazil. Novon 16: 112-128.; Souza-Silva et al. 2010Souza-Silva RF, Rapini A, Morales JF. 2010. Mandevilla catimbauensis (Apocynaceae), a new species from the semi-arid region, Pernambuco, Brazil. Edinburgh Journal of Botany 67: 1-5. ; Morales 2018aMorales JF. 2018a. Studies in Neotropical Apocynaceae LIV: A synopsis of Asketanthera. Candollea 73: 7-17.; bMorales JF. 2018b. New species and combinations of Apocynaceae, Bignoniaceae, Clethraceae, and Cunoniaceae from the Neotropics. Anales del Jardín Botánico de Madrid 75: e071.; Coelho et al. 2020Coelho CA, da Silva Pereira MR, Amorin BS. 2020. Preliminary Angiosperm Checklist in an Area South of the Madeira River, Manicoré, Amazonas, Brazil. Acta Brasiliensis 4: 29.; Morales & Kollmann 2020aMorales JF, Kollmann L. 2020a. Increasing the known floristic diversity of Brazilian inselbergs: two new species of Mandevilla (Apocynaceae) from Espírito Santo. Acta Botanica Brasilica 34: 107-116.; bMorales JF, Kollmann L. 2020b. War, types, and Mandevilla (Apocynaceae): A new species and the rediscovery of a rare species from Brazil. Darwiniana, nueva serie 8: 449-459.; Morales & Morais 2020Morales JF, Morais IL 2020. Studies in the Neotropical Apocynaceae LV: A new Mandevilla from Bahia, Brazil, with notes on the diversity of the genus. Systematic Botany 45: 183-189.; Morales et al. 2022Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093.). The Flora and Fungi of Brazil recognized 71 species of Mandevilla (Flora e Funga do Brasil 2020Flora e Funga do Brasil. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/. 23 Jan. 2023.
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), 44 of them endemic. However, two species must be excluded as endemic: Mandevilla alexicaca (Mart. ex Stadelm.) J.F. Morales & A. Fuentes is present in Peru and Bolivia and M. illustris (Vell.) Woodson has been recorded in Peru, Bolivia, and Paraguay (Ezcurra et al. 1992Ezcurra C, Endress ME, Leeuwenberg AJM. 1992. Apocynaceae. In: Spichiger R, Ramella L (eds.). Flora del Paraguay. Missouri, Conservatoire et Jardin Botaniques de Genève, Suiza, y Missouri Botanical Garden, U.S.A . p. 1-121.; Fuentes & Morales 2014Fuentes AF, Morales JF. 2014. Apocynaceae. In: Jørgensen PM, Nee MH, Beck S (eds.). Catalogo de las Plantas Vasculares de Bolivia. Missouri, Missouri Botanical Garden Press. vol. 127, p. 234-255.). The current species number for all of Brazil could be around 78 species considering several species described recently (Coelho et al. 2020Coelho CA, da Silva Pereira MR, Amorin BS. 2020. Preliminary Angiosperm Checklist in an Area South of the Madeira River, Manicoré, Amazonas, Brazil. Acta Brasiliensis 4: 29.; Morales et al. 2022Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093.). Taking into consideration the large number of unexplored Inselbergs in Espírito Santo as well as Mandevilla's high diversity and narrow endemism in these formations, this number is likely an under-estimate.

In preparation for a revision on Mandevilla reported for Goiás and Distrito Federal, a new species endemic to limestone formations in Ponte de Pedra, Paraúna, Goiás state, was found and is described here. A note regarding geophytic species in traditional Apocynaceae (focused on Mandevilla) is provided.

Material and methods

This paper is part of the monograph of Mandevilla. Specimens (including types) from the following herbaria were examined: A, AAU, ALCB, AS, ASE, B, BHCB, BM, BOTU, BR, C, CAS, CAY, CEN, CEPEC, CESJ, CGE, CHOCO, CIIDIR, CIMI, CM, COAH, COL, CPUN, CR, CVRD, CUCV, CUZ, DUKE, E, EAP, ECON, ENCB, ESA, ESAL, F, FI, FI-W, FLAS, FLOR, FTG, FUEL, G, G-DC, GA, GFJP, GH, GUA, HAL, HAO, HAS, HB, HBG, HRB, HRCB, HTO, HUA, HUEFS, HUFU, HUQ, IAN, IBGE, ICN, INPA, JAR, JAUM, JPB, K, L, LAGU, LD, LIL, LP, LPB, LV, LZ, M, MA, MBM, MBML, MCNS, MEDEL, MEXU, MICH, MIN, MG, MO, MOL, MVFA, MVJB, NY, O, P, P-HB, P-JU, P-LA, PAMG, PEL, PEUFR, PMA, PR, Q, QCA, QCNE, R, RB, S, SI, SPF, SPFR, SP, TRIN, TULV, U, UC, UB, ULM, UNAH, UPCB, UPS, US, USF, USJ, USM, USZ, UTEP, UVAL, W, WAG, WIS, WU, VALLE, VEN, VIC, Z and ZT. Acronyms cited follow Thiers (2023)Thiers B. 2023 [continuously updated]. Index herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s virtual herbarium. http://sweetgum.nybg.org/science/ih/. 1 Jan. 2023.
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, continuously updated). Description of the new species was based on herbarium specimens and fresh material. The species concept of Templeton (1989)Templeton AR. 1989. The meaning of species and speciation: A genetic perspective. In: Otte D, Endler JA (eds.). Speciation and its consequences. Sunderland, Sinauer Associates, Inc. p. 3-27. is followed for the description of the new species, considering the phenotypic variability and habitat restrictions. Typification of species (including homotypic and heterotypic synonyms) reported for Goiás and Distrito Federal is provided in Morales (2022)Morales JF. 2022. Miscellaneous typifications and new synonyms toward a revision of Mandevilla (Apocynaceae). Novon 30: 128-137.. Morphological terminology follows Radford et al. (1974)Radford AE, Dickison WE, Massey JR, Bell CR. 1974. Vascular plant systematics. New York, Harper & Row.. Pertinent morphological characters are illustrated in Morales (2005c)Morales JF. 2005c. Estudios en las Apocynaceae Neotropicales XIX: La familia Apocynaceae s. str. (Apocynoideae y Rauvolfioideae) de Costa Rica. Darwiniana, nueva serie 43: 90-191., and definition of parts of the corolla follow Morales and Fuentes (2004)Morales JF, Fuentes A. 2004. Estudios en las Apocynaceae Neotropicales VIII: Nuevas especies de Mandevilla para Peru y Bolivia, con notas sobre la morfología floral en corolas infundibuliformes. Candollea 59: 167-174.. Measurements given in the key are based on an unpublished monograph of Mandevilla (Morales, unpub. data) and these can differ with respect to those given in previous papers.

The conservation status was assessed using the IUCN Red List Categories and Criteria, version 3.1 (IUCN 2014IUCN - International Union for Conservation of Nature. 2014. The IUCN Red List of Threatened Species. Version 2014.3. http://www.iucnredlist.org. 23 Jan. 2023.
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). Calculating the extent of occurrence (EOO) or the area of occupancy (AOO) was not made because at least three localities are needed. Population size is poorly understood in neotropical apocynoids (Morales 2018aMorales JF. 2018a. Studies in Neotropical Apocynaceae LIV: A synopsis of Asketanthera. Candollea 73: 7-17.; Morales & Cornejo 2019Morales JF, Cornejo X. 2019. Two New Species of Prestonia (Apocynaceae) from Colombia, Ecuador, and Peru. Systematic Botany 44: 197-202. ), because evaluating this feature is not possible at this time.

Taxonomic treatment

Mandevilla calcicola J. F. Morales, M.E. Endress & I. L. Morais, sp. nov.

Type:-BRAZIL. Paraúna, Goiás: estrada GO-50, c. 30 km após a cidade de Motividiu, estrada para Paraúna, Ponte de Pedra, margens do Ribeirão Corrente, afloramentos de rocha calcárea, 6/I/2020 (fl.), I.L. de Morais, J.F. Morales 5850 (Holotype JAR!; isotype: HUFU!).

Figs. 1-3, 4C.

Figure 1
Mandevilla calcicola. A. flowering branch. B. detail of adaxial base of leaf blade and petiole, showing the arrangement of colleters. C. adaxial view of two sepals, showing colleters at the base. D. corolla tube partially open, showing anthers. E. anther, dorsal view. F. style-head. G. nectaries and ovary (Fabio de Barros 2196, HST; drawn by J F Morales).

Figure 2
Mandevilla calcicola (Ponte de Pedra, Paraúna, Goiás, I.L. de Morais & J.F. Morales 5850, JAR). A. Tuberous roots. B. stem with inflorescence. C. detail of leaf base. D. habitat.

Figure 3
Distribution map of Mandevilla calcicole.

Geophytic woody vine or scandent shrub with tuberous roots and milky latex; branchlets cylindrical to subcylindrical, glabrous to glabrescent; interpetiolar colleters inconspicuous, up to 0.3 mm long. Leaves opposite, sessile to subsessile, petioles (when present) up to 1.1 mm long; leaf blades 8.8-11.6 × 5.3-7.2 cm, obovate, obovate-elliptic to elliptic, acute to retuse apically, conspicuously cordate basally, with numerous colleters clustered at base of the midrib adaxially, sometimes with a few colleters spread along the secondary and tertiary veins, not bullate, membranaceous, glabrous on both surfaces, not revolute, secondary veins slightly impressed adaxially, conspicuously impressed abaxially, tertiary veins slightly impressed adaxially, inconspicuous abaxially. Inflorescences longer than subtending leaves, terminal to subterminal, glabrous, 10-13-flowered, peduncle 40-52 mm long, pedicels 10-13 mm long, bracts 1-1.7 × 0.5-0.8 mm, linear-ovate, scarious; sepals 6-7.5 × 1.5-1.7 mm, similar in length, narrowly ovate, acuminate at the apex, the apex slightly reflexed, subfoliaceous, glabrous on the abaxial surface, with numerous colleters irregularly disposed at the base adaxially; corolla infundibuliform, pink, the throat yellow within, glabrous, lower part 20-25 × 2.3-3 mm long, upper part 31-33 mm long, campanulate, 24-26 mm diam., apex of the floral bud acuminate; lobes 38-41 × 27-30 mm, obovate, extended; stamens inserted at the base of the upper part of the corolla tube, anthers 7.5-8.5 mm long, dorsally glabrous, base auriculate, with auricles rounded; style-head 1.9-2.2 mm long, the main body ca. 1.5 mm long, apical appendages ca. 0.2 mm long; ovary 2-2.5 mm long, glabrous; nectaries two, entire to subentire. Follicles unknown.

Distribution and habitat: Endemic to Goiás, where it grows on limestone cliffs in riparian forest at 675-725 m.

Phenology: Flowering in January and February.

Etymology: The name refers to the limestone cliff habitat where it grows.

Conservation status: Mandevilla calcicola is only known from limestone cliffs in a small area within a single locality and on a specific substrate (limestone). Therefore, the extent of occurrence (EOO) and the area of occupancy (AOO) were not calculated. The type locality is an unprotected area surrounded by extensive soy, sugarcane, and corn monoculture. The forest along the river is well preserved but confined along the river margins and under tremendous pressure due to monocultures around.

We assessed this species as Critically Endangered (CR) based on criteria B1ab (iii, iv) + B2ab (iii, iv), considering the number of localities and the severely fragmented and continuing decline of its habitat. Further fieldwork is necessary to discover if there are additional populations along the river or nearby localities with limestone formations.

Observations: Of the species of Mandevilla in Brazil, M. calcicola shows some similarity to M. splendens, which is endemic to inselbergs in Minas Gerais and Rio de Janeiro states, in its subsessile leaves, leaf blade obovate, obovate-elliptic to elliptic, and pink corolla, but can be distinguished by its glabrous leaf blade (vs. tomentulose), sepals 6-7.5 mm long (vs. 9-14 mm), the upper part of the corolla tube campanulate (vs. broadly conic) and corolla lobes 38-41 mm long (vs. 25-34 mm). Mandevilla calcicola is the second species described as endemic to Goiás (following M. myriophyllum (Taub.) Woodson).

Paratypes: BRAZIL. Paraúna, Goiás: estrada GO-411, ca. 90 km após a cidade de Paraúna, localidade denominada Ponte de Pedra, margens do Ribeirão Corrente, 25/I/1991 (fl), Fabio de Barros 2196 (CR, HST [HST005584], SP ).

Mandevilla grows in a variety of habitats, from dry forest to tropical wet forest, premontane and montane forest, white-sand formations in the Guyana shield and Amazon basin, inselbergs, and Restinga in Brazil. However, this is the first record of an edaphic endemic species of Mandevilla specialized on limestone and few Apocynaceae are reported as endemic to this substrate (e.g., Allamanda (Souza-Silva & Rapini 2009Souza-Silva RF, Rapini A. 2009. Allamanda calcicola (Apocynaceae), an Overlooked New Species from Limestone Outcrops in the States of Minas Gerais and Bahia, Brazil. Kew Bulletin 64: 171-174.); Ceropegia (Kidyoo 2021Kidyoo M. 2021. Two New Species of Ceropegia (Apocynaceae, Asclepiadoideae) on Limestone Hills in Northern Thailand. Tropical Natural History 21: 1-11.); Hoya (Rodda & Juhonewe 2012Rodda M, Juhonewe NS. 2012. Hoya vangviengiensis (Apocynaceae, Asclepiadoideae), a new species from limestone formations of Vang Vieng, Lao PDR. Webbia 67: 23-27.); Ruehssia (as Marsdenia) Espírito Santo et al. (2018)Espírito Santo FS, Bitencourt C, Ribeiro PL, Rapini A. 2018. Two new species of Marsdenia (Apocynaceae) from limestone outcrops in Brazil. Willdenowia 48: 109-116.; Wrightia (Middleton & Santisuk 2001Middleton DJ, Santisuk T. 2001. A new species of Wrightia (Apocynaceae: Apocynoideae) from Thailand. Thai Forest Bulletin (Botany) 29: 1-10. ; Middleton 2007Middleton DJ. 2007. A new species of Wrightia (Apocynaceae: Apocynoideae) from Thailand. Thai Forest Bulletin (Botany) 35: 80-85.)). Mandevilla calcicola was known from a single specimen collected 29 years ago, but recent fieldwork has revealed that it is locally common at the type locality, but always restricted to limestone cliffs. Ponte de Pedra is highly deforested and the remnant forests are restricted to strips along the Correntes river surrounded by cane and soy fields.

Note on the occurrence of geophytes in rauvolfioids and apocynoids: Geophytes - plants that regrow from buds on underground storage organs, such as rhizomes, tubers, xylopods, bulbs, and corms - are widespread among the land plants (Tribble et al. 2021Tribble CM, Martínez-Gómez J, Howard CC et al. 2021. Get the shovel: Morphological and evolutionary complexities of belowground organs in geophytes. American Journal of Botany 108: 372-387.). This adaptive feature allows them to survive during periods of environmental stress such as drought or fire (Howard et al. 2019Howard DD, Landis JB, Beaulieu JM, Cellinese N. 2019. Geophytism in monocots leads to higher rates of diversification. New Phyologist 225: 1023-1032.). In Apocynaceae, geophytism is well-documented in some of the derived lineages like Periplocoideae and in Fockeeae, Ceropegieae and diverse Asclepiadeae of Asclepiadoideae (Verhoeven et al. 2003Verhoeven RL, Liede S, Endress ME. 2003. The tribal position of Fockea and Cibirhiza (Apocynaceae-Asclepiadoideae): Evidence from pollinium structure and cpDNA sequence data. Grana 42: 70-81.; Meve & Liede 2004Meve U, Liede S. 2004. Generic delimitations in tuberous Periplocoideae (Apocynaceae) from Africa and Madagascar. Annals of Botany 93: 407-414.; Bruyns et al. 2015Bruyns PV, Klak C, Hanáček P. 2015. Recent radiation of Brachystelma and Ceropegia (Apocynaceae) across the Old World against a background of climatic change. Molecular Phylogenetics and Evolution 90: 49-66.; Brito & Bruyns 2016Brito SJ, Bruyns P. 2016. Three New Species of Brachystelma from Tamil Nadu, India. Haseltonia 22: 48-54.; Endress et al. 2019Endress ME, Meve U, Middleton DJ, Liede-Schumann S. 2019. Apocynaceae. In: Kadereit JW, Bittrich V (eds.). Flowering plants. Eudicots Apiales and Gentianales (except Rubiaceae), K. Kubitzki (ed.), Families and genera of vascular plants. Cham, Springer. vol. 15, p. 207-411.). In the basal nodes of the family (rauvolfioids and apocynoids), this condition has been less well studied, but in a handful of genera in geophytic species have been reported. In rauvolfioids, most species of the temperate to subtropical genus Amsonia die back in winter and resprout from rhizomes in spring (Woodson 1928Woodson RE Jr. 1928. Studies in the Apocynaceae. III. A monograph of the genus Amsonia. Annals of the Missouri Botanical Garden 15: 379-434.). Few geophytic species have been known in neotropical rauvolfioids, such as Rauvolfia anomala Rapini & I. Koch and R. weddeliana Müll. Arg. However, several questions need to be addressed, such as how many genera have geophytic species, why sympatric species of the same genus or different genera in habitats with strong drought seasons do not have underground storage organs or if the presence of geophytes species is restricted to specific clades within a genus.

Among apocynoids, Apocynum and Cycladenia, which occur mainly in colder regions or higher elevations, are rhizomatous and dormant in the winter (Woodson 1930Woodson RE Jr. 1930. Studies in the Apocynaceae. I. A Critical Study of the Apocynoideae (with special reference to the genus Apocynum). Annals of the Missouri Botanical Garden 17: 212.; Sipes & Tepedino 1996Sipes SD, Tepedino VJ, 1996. Cycladenia humilis var. jonesii (Apocynaceae). In: Maschinski J, Hammond HD, Holter L (eds.). Southwestern rare and endangered plants. Proceedings of the Second Conference Darby, PA, USA, DIANE Publishing. p. 158.). Most genera of apocynoids with geophytic species, however, occur in habitats where the stress factor is a long-lasting dry season or extreme drought conditions. Most of these has been reported from Brazil, grow in Cerrados, campos rupestres and inselbergs, and have tubers, tuberous roots, or a xylopod. Except for Rhodocalyx, most of these have been reported in the genus Mandevilla (e.g., Kinoshita et al. 2005Kinoshita LS, Simões AO, Koch I, Sales MF, Rio MCS, Marcondes-Ferreira W. 2005. Apocynaceae. In: Wanderley MGL, Shepherd GJ, Melhem TS, Martins SE, Kirizawa M, Giulietti AM (eds.). Flora Fanerogâmica do Estado de São Paulo. São Paulo, Instituto de Botânica. vol. 4, p. 35-92.; Sales et al. 2006Sales MF, Kinoshita LS, Simões AO. 2006. Eight new species of Mandevilla Lindley (Apocynaceae, Apocynoideae) from Brazil. Novon 16: 112-128.; Monteiro et al. 2017Monteiro FK, de Morais-Rodrigues E, do Nascimento VM, de Melo JIM. 2017. Primer registro de Mandevilla dardanoi (Apocynaceae) para el estado de Paraíba, Brasil. Revista Mexicana de Biodiversidad 88: 755-758.; Morales et al. 2022Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093.), but specimens (and subsequently descriptions) rarely include or describe these underground parts. The first author has cultivated or studied in situ different species of Echiteae (e.g., Echites, Macropharynx, Prestonia, Temnadenia), Odontadenieae (e.g., Odontadenia, Pentalinon, Secondatia, Stipecoma), and Rhabdadeniaeae (Rhabdadenia) without ever observing the presence of geophytic species. A number of Mandevilla species growing on inselbergs have these structures, where geophytism is a common condition (Porembski 2007Porembski S. 2007. Tropical inselbergs: Habitat types, adaptive strategies and diversity patterns. Revista Brasileira de Botânica 30: 579-586.), as do species from Cerrados or campos rupestres (Appezato-da-Glória & Estelita 2000Appezato-da-Glória B, Estelita MEM. 2000. The development anatomy of the subterranean system in Mandevilla illustris (Vell.) Woodson and M. velutina (Mart. ex. Stadelm.) Woodson (Apocynaceae). Revista Brasileira de Botânica 23: 27-35.), but specific studies concerning the characterization of geophyte species in the genus are absent. The few available studies focus on anatomical examinations of a few species (e.g., M. illustris (Vell.) Woodson and M. pohliana (Stadelm.) A.H. Gentry [Appezato-da-Gloria & Estelita 2000]; M. tenuifolia [Costa 2002Costa ATL. 2002. Adaptações Morfo-anatômicas da folha e do sistema subterrâneo de Mandevilla tenuifolia (Mikan) Woodson (Apocynaceae). - Ceará. MSc Thesis, Universidade Federal do Ceará, Brazil.]; M. atroviolacea (Stadelm.) Woodson, [Lopes 2007Lopes KLB. 2007. Morfoanatomia dos órgãos vegetativos de Mandevilla atroviolacea (Stadelm.) Woodson (Apocynaceae, Apocynoideae) em um afloramento rochoso no parque estadual da Serra do Brigadeiro. PhD Thesis, Universidade Federal de Viçosa, Brazil.; Lopes-Mattos et al. 2013Lopes-Mattos KLB, Azevedo AA, Soares AA, Meira RMSA. 2013. Underground system of Mandevilla atroviolacea (Stadelm.) Woodson (Apocynaceae, Apocynoideae) from the Brazilian high-altitude grassland. South African Journal of Botany 87: 27-33.]).

Mandevilla calcicola has tuberous roots, a common feature of Mandevilla species growing on rocks in South America. Following the clades names proposed by Morales et al. (2022)Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093., almost all the species studied by the first author with tuberous roots, tubers, or xylopods belong to the Mandevilla clade, except for a group of species from the Exothostemon clade, restricted to granitic or quarzitic outcrops from the Guyana shield in Colombia and Venezuela (e.g., M. lancifolia Woodson). However, the presence of this condition in the genus has not been studied in a phylogenetic context, which could provide helpful characters or synapomorphies to characterize clades.

A key to Mandevilla in Goiás and Distrito Federal, Brazil is provided here. A total of 20 species are included, with only M. myriophyllum and M. calcicola being endemic to Goiás (Fig. 4). Mandevilla antennacea (= M. rugellosa (Rich) Allorge fide Fuentes and Morales (2014)Fuentes AF, Morales JF. 2014. Apocynaceae. In: Jørgensen PM, Nee MH, Beck S (eds.). Catalogo de las Plantas Vasculares de Bolivia. Missouri, Missouri Botanical Garden Press. vol. 127, p. 234-255.) is reported for Distrito Federal in Flora of Brazil. Five vouchers were cited: Ferreira 11588 (NY) [Mato Grosso], Teixeira et al. 510 (F, INPA, K, MG, MO, NY, RB, US) [Rondônia], Hatschbach 62580 (ESA, MT, NY, UEC) [Mato Grosso], and Ducke 7636 (RB) [Amazonas]. The specimen Ducke 7636 is misidentified: its correct identification is M. scabra. In Brazil, Mandevilla rugellosa has been recorded only in Acre, Mato Grosso, and Rondônia states. Working on a new revision of Mandevilla, the first author has never seen a specimen from Goiás or Distrito Federal. Therefore, this species is not included in the key.

Figure 4
Mandevilla in Goiás and Distrito Federal. A. M. abortiva (Brazil, Bahia, 2019, J F Morales). B. M. alexicaca (Brazil, Goiás, 2018, J F Morales). C. M. calcicola (Brazil, Goiás, 2019, J F Morales). D. M. clandestina (Brazil, Goiás, 2018, I L Morais). E. M. coccinea (Brazil, Rio Grande do Sul, 2017, J F Morales). F. M. emarginata (Brazil, Goiás, 2015, J F Morales). G. M. hirsuta (Brazil, Goiás, 2019, J F Morales). H. M. illustris (Brazil, Goiás, 2018, I L Morais). I. M. longiflora (Brazil, Goiás, 2019, J F Morales). J. M. martii (Brazil, Goiás, 2018, J. F Morales). K. M. myriophyllum (Brazil, Goiás, 2017, J F Morales). L. M. novocapitalis (Brazil, Goiás, 2017, J F Morales). M. M. petraea (showing verticillate leaves) (Brazil, Paraná, 2015, J F Morales). N. M. pohliana (Brazil, Goiás, 2018, J F Morales). O. M. scabra (Brazil, Goiás, 2019, J F Morales). P. M. spigeliiflora (Brazil, Goiás, 2019, I L Morais). Q. M. tenuifolia (Brazil, Goiás, 2018, J F Morales). R. M. velame (Brazil, Goiás, 2015, J F Morales). S. M. virescens (Brazil, Paraná, 2016, J F Morales). T. M. widgrenii (Brazil, Goiás, 2019, J F Morales).

Key to species of Mandevilla in Goiás and Distrito Federal, Brazil

1. Leaf blades with colleters spread along midvein adaxially; corolla tube gibbous at the base . . . . . . . . .. . . . . . . . . . . . . . . 2

2. Floral bracts 11-21 × 1.5-8 mm, foliaceous; sepals 7-12(-15) × 1-3 mm . . . . . . . . .. . . . . . . . . . . . . . . M. hirsuta

2. Floral bracts 1-5(-8) × 0.5-1.5(-2) mm, scarious; sepals 1.7-3.5 × 1-2.7 mm . . . . . . . . .. . . . . . . . . . . . . . . 3

3. Leaf blades with tertiary veins not impressed abaxially; erect to suberect herbs or herbaceous vines . . . . . . . . .. . . . . . . . . . . . . . . M. abortiva

3. Leaf blades with tertiary veins impressed abaxially; herbaceous to woody vines . . . . . . . . .. . . . . . . . . . . . . . . 4

4. Corolla lobes white; corolla mouth 5-6 mm diam. . . . . . . . . .. . . . . . . . . . . . . . . M. clandestina

4. Corolla lobes yellow; corolla mouth 15-24(-27) mm diam. . . . . . . . . .. . . . . . . . . . . . . . . M. scabra

1. Leaf blades with colleters grouped at the base of midvein adaxially; corolla tube straight at the base . . . . . . . . .. . . . . . . . . . . . . . . 5

5. Corolla tube 8-18.7 cm long; leaf blades conspicuously white-pubescent abaxially . . . . . . . . .. . . . . . . . . . . . . . . 6

6. Leaf blades white-lanuginose on both surfaces . . . . . . . . .. . . . . . . . . . . . . . . M. velame

6. Leaf blades with only the abaxial surface white-lanuginose, adaxial surface green . . . . . . . . .. . . . . . . . . . . . . . . 7

7. Leaves usually verticillate, rarely opposite at some nodes, blades 0.1-1 cm wide . . . . . . . . .. . . . . . . . . . . . . . . M. petraea

7. Leaves usually opposite, rarely verticillate at some nodes, blades more than 1.4 cm wide . . . . . . . . .. . . . . . . . . . . . . . . 8

8. Inflorescence usually 5-12-flowered . . . . . . . . .. . . . . . . . . . . . . . . M. martii

8. Inflorescence 1-3(4)-flowered . . . . . . . . .. . . . . . . . . . . . . . . 9

9. Lower part of the corolla tube 10.5-15.4 cm long, mouth 15-23 mm diam. . . . . . . . . .. . . . . . . . . . . . . . . M. longiflora

9. Lower part of the corolla tube 5.8-7.8 cm long, mouth 6-11 mm diam. . . . . . . . . .. . . . . . . . . . . . . . . M. virescens

5. Corolla tube less than 6 cm long; leaf blades glabrous, glabrate abaxially, or if pubescent, then this never white . . . . . . . . .. . . . . . . . . . . . . . . 10

10. Corolla red to orange . . . . . . . . .. . . . . . . . . . . . . . . 11

11. Corolla hypocrateriform, mouth 2.5-3.1 mm diam.; corolla lobes obovate to narrowly obovate . . . . . . . . .. . . . . . . . . . . . . . . M. coccinea

11. Corolla narrowly infundibuliform, mouth 3-6 mm diam.; corolla lobes narrowly elliptic to narrowly obovate-elliptic . . . . . . . . .. . . . . . . . . . . . . . . M. spigeliiflora

10. Corolla pink, lilac, pinkish-white to green or greenish-white . . . . . . . . .. . . . . . . . . . . . . . . 12

12. Corolla hypocrateriform . . . . . . . . .. . . . . . . . . . . . . . . 13

13. Sepals 13-21(-24) mm long, as long as the corolla tube; corolla lobes green to greenish-white . . . . . . . . .. . . . . . . . . . . . . . . M. emarginata

13. Sepals 1.5-10 mm long, much shorter than the corolla tube; corolla lobes pink, lilac to pinkish white. . . . . . . . .. . . . . . . . . . . . . . . 14

14. Corolla lobes 17-31 × 11-23 mm; follicles 19-21 cm long . . . . . . . . .. . . . . . . . . . . . . . . M. novocapitalis

14. Corolla lobes 6-9 × 4-6 mm; follicles 3.5-6.2 cm long . . . . . . . . .. . . . . . . . . . . . . . . 15

15. Leaf blades 5-15 × 0.2-0.7(-1) mm, filiform . . . . . . . . .. . . . . . . . . . . . . . . M. myriophyllum

15. Leaf blades (10-)14-95(-135) × 1.1-90(-160) mm, linear-elliptic, narrowly elliptic, narrowly ovate-elliptic to narrowly obovate . . . . . . . . .. . . . . . . . . . . . . . . M. tenuifolia

12. Corolla infundibuliform . . . . . . . . .. . . . . . . . . . . . . . . 16

16. Woody vines; leaf blade conspicuously cordate basally; limestone cliffs (gallery forest) . . . . . . . . .. . . . . . . . . . . . . . . M. calcicola

16. Erect plants; leaf blade cuneate, obtuse, rounded to slightly cordate basally; Cerrados and swamp areas . . . . . . . . .. . . . . . . . . . . . . . . 17

17. Leaves opposite to verticillate, blades 2-3 mm wide; swamp areas . . . . . . . . .. . . . . . . . . . . . . . . M. widgrenii

17. Leaves opposite, blades 5-95 mm wide; Cerrados . . . . . . . . .. . . . . . . . . . . . . . . 18

18. Upper part of the corolla tube cylindrical to urceolate-cylindrical, 21-48 mm long . . . . . . . . .. . . . . . . . . . . . . . . M. pohliana

18. Upper part of the corolla tube conical to narrowly conical, 10-23 mm long . . . . . . . . .. . . . . . . . . . . . . . . 19

19. Corolla mouth 11-19 mm diam.; upper part of corolla tube broadly conical . . . . . . . . .. . . . . . . . . . . . . . . M. alexicaca

19. Corolla mouth 4-9 mm diam.; upper part of corolla tube narrowly conical . . . . . . . . .. . . . . . . . . . . . . . . M. illustris

Acknowledgments

This study was partially supported by the University of the West Indies (Trinidad and Tobago), and the Herbarium JAR, Universidade Estadual de Goiás (Brazil).

References

  • Adeney JM, Christensen NL, Vicentini A, Cohn-Haft M. 2016. White‐sand Ecosystems in Amazonia. Biotropica 48: 7-23.
  • Alvarado-Cárdenas LO, Morales JF. 2014. El género Mandevilla (Apocynaceae: Apocynoideae, Mesechiteae) en México. Botanical Sciences 92: 59-79.
  • Appezato-da-Glória B, Estelita MEM. 2000. The development anatomy of the subterranean system in Mandevilla illustris (Vell.) Woodson and M. velutina (Mart. ex. Stadelm.) Woodson (Apocynaceae). Revista Brasileira de Botânica 23: 27-35.
  • Brazil Flora Group. 2022. Brazilian Flora 2020. Leveraging the power of a collaborative scientific network. Taxon 71: 178-198.
  • Brito SJ, Bruyns P. 2016. Three New Species of Brachystelma from Tamil Nadu, India. Haseltonia 22: 48-54.
  • Bruyns PV, Klak C, Hanáček P. 2015. Recent radiation of Brachystelma and Ceropegia (Apocynaceae) across the Old World against a background of climatic change. Molecular Phylogenetics and Evolution 90: 49-66.
  • Coelho CA, da Silva Pereira MR, Amorin BS. 2020. Preliminary Angiosperm Checklist in an Area South of the Madeira River, Manicoré, Amazonas, Brazil. Acta Brasiliensis 4: 29.
  • Costa ATL. 2002. Adaptações Morfo-anatômicas da folha e do sistema subterrâneo de Mandevilla tenuifolia (Mikan) Woodson (Apocynaceae). - Ceará. MSc Thesis, Universidade Federal do Ceará, Brazil.
  • Endress ME, Meve U, Middleton DJ, Liede-Schumann S. 2019. Apocynaceae. In: Kadereit JW, Bittrich V (eds.). Flowering plants. Eudicots Apiales and Gentianales (except Rubiaceae), K. Kubitzki (ed.), Families and genera of vascular plants. Cham, Springer. vol. 15, p. 207-411.
  • Espírito Santo FS, Bitencourt C, Ribeiro PL, Rapini A. 2018. Two new species of Marsdenia (Apocynaceae) from limestone outcrops in Brazil. Willdenowia 48: 109-116.
  • Ezcurra C, Endress ME, Leeuwenberg AJM. 1992. Apocynaceae. In: Spichiger R, Ramella L (eds.). Flora del Paraguay. Missouri, Conservatoire et Jardin Botaniques de Genève, Suiza, y Missouri Botanical Garden, U.S.A . p. 1-121.
  • Felfili JM, Fagg CW. 2007. Floristic composition and structure of the “cerrado” sensu stricto on rocky soils in northern Goiás and southern Tocantins, Goiás. Brazilian Journal of Botany 30: 375-385.
  • Flora e Funga do Brasil. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/ 23 Jan. 2023.
    » http://floradobrasil.jbrj.gov.br/
  • Fuentes AF, Morales JF. 2014. Apocynaceae. In: Jørgensen PM, Nee MH, Beck S (eds.). Catalogo de las Plantas Vasculares de Bolivia. Missouri, Missouri Botanical Garden Press. vol. 127, p. 234-255.
  • Howard DD, Landis JB, Beaulieu JM, Cellinese N. 2019. Geophytism in monocots leads to higher rates of diversification. New Phyologist 225: 1023-1032.
  • IUCN - International Union for Conservation of Nature. 2014. The IUCN Red List of Threatened Species. Version 2014.3. http://www.iucnredlist.org 23 Jan. 2023.
    » http://www.iucnredlist.org
  • Kidyoo M. 2021. Two New Species of Ceropegia (Apocynaceae, Asclepiadoideae) on Limestone Hills in Northern Thailand. Tropical Natural History 21: 1-11.
  • Kinoshita LS, Simões AO, Koch I, Sales MF, Rio MCS, Marcondes-Ferreira W. 2005. Apocynaceae. In: Wanderley MGL, Shepherd GJ, Melhem TS, Martins SE, Kirizawa M, Giulietti AM (eds.). Flora Fanerogâmica do Estado de São Paulo. São Paulo, Instituto de Botânica. vol. 4, p. 35-92.
  • Lenza E, Pinto JRR, Pinto AS, Maracahipes L, Bruziguessi EL. 2011. Comparação da vegetação arbustivo-arbórea de uma área de cerrado rupestre na Chapada dos Veadeiros, Goiás e áreas de cerrado sentido restrito do Bioma Cerrado. Revista Brasileira de Botânica 34: 247-259.
  • Lopes KLB. 2007. Morfoanatomia dos órgãos vegetativos de Mandevilla atroviolacea (Stadelm.) Woodson (Apocynaceae, Apocynoideae) em um afloramento rochoso no parque estadual da Serra do Brigadeiro. PhD Thesis, Universidade Federal de Viçosa, Brazil.
  • Lopes-Mattos KLB, Azevedo AA, Soares AA, Meira RMSA. 2013. Underground system of Mandevilla atroviolacea (Stadelm.) Woodson (Apocynaceae, Apocynoideae) from the Brazilian high-altitude grassland. South African Journal of Botany 87: 27-33.
  • Marimon Junior BH, Haridasan M. 2005. Comparação da vegetação arbórea e características edáficas de um cerradão e cerrado sensu stricto em áreas adjacentes sobre solo distrófico no leste de Mato Grosso, Brasil. Acta Botanica Brasilica 19: 913-926.
  • Meve U, Liede S. 2004. Generic delimitations in tuberous Periplocoideae (Apocynaceae) from Africa and Madagascar. Annals of Botany 93: 407-414.
  • Middleton DJ. 2007. A new species of Wrightia (Apocynaceae: Apocynoideae) from Thailand. Thai Forest Bulletin (Botany) 35: 80-85.
  • Middleton DJ, Santisuk T. 2001. A new species of Wrightia (Apocynaceae: Apocynoideae) from Thailand. Thai Forest Bulletin (Botany) 29: 1-10.
  • Monteiro FK, de Morais-Rodrigues E, do Nascimento VM, de Melo JIM. 2017. Primer registro de Mandevilla dardanoi (Apocynaceae) para el estado de Paraíba, Brasil. Revista Mexicana de Biodiversidad 88: 755-758.
  • Morales JF. 2005a. Estudios en las Apocynaceae neotropicales XVIII: Dos nuevas especies de Mandevilla (Apocynoideae, Mesechiteae) para Brasil. Darwiniana, nueva serie 43: 84-89.
  • Morales JF. 2005b. Estudios en las Apocynaceae neotropicales XI: Una nueva especie de Mandevilla (Apocynoideae: Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay. Sida 21: 1549-1555.
  • Morales JF. 2005c. Estudios en las Apocynaceae Neotropicales XIX: La familia Apocynaceae s. str. (Apocynoideae y Rauvolfioideae) de Costa Rica. Darwiniana, nueva serie 43: 90-191.
  • Morales JF. 2007. Estudios en las Apocynaceae Neotropicales XXX: Tres nuevas especies andinas de Mandevilla (Apocynoideae, Mesechiteae). Journal of the Botanical Research Institute of Texas 1: 853-857.
  • Morales JF. 2009. Estudios en las Apocynaceae Neotropicales XXXVIII: Tres nuevas especies de Mandevilla (Apocynoideae, Mesechiteae) para Colombia y Venezuela. Journal of Botanical Research Institute of Texas 3: 565-571.
  • Morales JF. 2011. Estudios en las Apocynaceae Neotropicales XLII: sinopsis del género Mandevilla (Apocynoideae: Mesechiteae) en Colombia. Journal of the Botanical Research Institute of Texas 5: 521-543.
  • Morales JF. 2013. Estudios en las Apocynaceae Neotropicales XLIX: Sinopsis de las Apocynaceae (Apocynoideae, Rauvolfioideae) de Chile. Darwiniana, nueva serie 1: 39-45.
  • Morales JF. 2018a. Studies in Neotropical Apocynaceae LIV: A synopsis of Asketanthera Candollea 73: 7-17.
  • Morales JF. 2018b. New species and combinations of Apocynaceae, Bignoniaceae, Clethraceae, and Cunoniaceae from the Neotropics. Anales del Jardín Botánico de Madrid 75: e071.
  • Morales JF, Cornejo X. 2019. Two New Species of Prestonia (Apocynaceae) from Colombia, Ecuador, and Peru. Systematic Botany 44: 197-202.
  • Morales JF. 2022. Miscellaneous typifications and new synonyms toward a revision of Mandevilla (Apocynaceae). Novon 30: 128-137.
  • Morales JF, Fuentes A. 2004. Estudios en las Apocynaceae Neotropicales VIII: Nuevas especies de Mandevilla para Peru y Bolivia, con notas sobre la morfología floral en corolas infundibuliformes. Candollea 59: 167-174.
  • Morales JF, Kollmann L. 2020a. Increasing the known floristic diversity of Brazilian inselbergs: two new species of Mandevilla (Apocynaceae) from Espírito Santo. Acta Botanica Brasilica 34: 107-116.
  • Morales JF, Kollmann L. 2020b. War, types, and Mandevilla (Apocynaceae): A new species and the rediscovery of a rare species from Brazil. Darwiniana, nueva serie 8: 449-459.
  • Morales JF, Morais IL 2020. Studies in the Neotropical Apocynaceae LV: A new Mandevilla from Bahia, Brazil, with notes on the diversity of the genus. Systematic Botany 45: 183-189.
  • Morales JF, Fontana AP, Kollmann L, Fraga CN. 2022. Inselbergs again: Four new species of Mandevilla (Apocynaceae) from Brazil. Systematic Botany 47: 1080-1093.
  • Porembski S. 2007. Tropical inselbergs: Habitat types, adaptive strategies and diversity patterns. Revista Brasileira de Botânica 30: 579-586.
  • Radford AE, Dickison WE, Massey JR, Bell CR. 1974. Vascular plant systematics. New York, Harper & Row.
  • Ribeiro JF, Walter BMT. 2008. As principais fitofisionomias do bioma Cerrado. In: Sano SM, Almeida SP, Ribeiro JF (eds.). Cerrado: Ecologia e flora. Brasília, Embrapa. p. 151-212.
  • Rodda M, Juhonewe NS. 2012. Hoya vangviengiensis (Apocynaceae, Asclepiadoideae), a new species from limestone formations of Vang Vieng, Lao PDR. Webbia 67: 23-27.
  • Sales MF. 1993. Estudos taxonômicos de Mandevilla Lindley subgênero Mandevilla (Apocynaceae) no Brasil. - Campinas. PhD Thesis, Universidade Estadual de Campinas, Brazil.
  • Sales MF, Kinoshita LS, Simões AO. 2006. Eight new species of Mandevilla Lindley (Apocynaceae, Apocynoideae) from Brazil. Novon 16: 112-128.
  • Santos TRR dos, Pinto JRR, Lenza E, Mews HA. 2011. The tree-shrub vegetation in rocky outcrop cerrado areas in Goiás State, Brazil. Revista Brasileira de Botânica 35: 281-294.
  • Sipes SD, Tepedino VJ, 1996. Cycladenia humilis var. jonesii (Apocynaceae). In: Maschinski J, Hammond HD, Holter L (eds.). Southwestern rare and endangered plants. Proceedings of the Second Conference Darby, PA, USA, DIANE Publishing. p. 158.
  • Soares MP, Reys P, Pifano DS, de Sá JL, da Silva P, Santos TM, Silva FG. 2015. Relationships between edaphic factors and vegetation in savannas on the Brazilian midwest region. Rodriguezia 62: 123-137.
  • Souza-Silva RF, Rapini A. 2009. Allamanda calcicola (Apocynaceae), an Overlooked New Species from Limestone Outcrops in the States of Minas Gerais and Bahia, Brazil. Kew Bulletin 64: 171-174.
  • Souza-Silva RF, Rapini A, Morales JF. 2010. Mandevilla catimbauensis (Apocynaceae), a new species from the semi-arid region, Pernambuco, Brazil. Edinburgh Journal of Botany 67: 1-5.
  • Templeton AR. 1989. The meaning of species and speciation: A genetic perspective. In: Otte D, Endler JA (eds.). Speciation and its consequences. Sunderland, Sinauer Associates, Inc. p. 3-27.
  • Thiers B. 2023 [continuously updated]. Index herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s virtual herbarium. http://sweetgum.nybg.org/science/ih/ 1 Jan. 2023.
    » http://sweetgum.nybg.org/science/ih/
  • Tribble CM, Martínez-Gómez J, Howard CC et al 2021. Get the shovel: Morphological and evolutionary complexities of belowground organs in geophytes. American Journal of Botany 108: 372-387.
  • Verhoeven RL, Liede S, Endress ME. 2003. The tribal position of Fockea and Cibirhiza (Apocynaceae-Asclepiadoideae): Evidence from pollinium structure and cpDNA sequence data. Grana 42: 70-81.
  • Woodson RE Jr. 1928. Studies in the Apocynaceae. III. A monograph of the genus Amsonia Annals of the Missouri Botanical Garden 15: 379-434.
  • Woodson RE Jr. 1930. Studies in the Apocynaceae. I. A Critical Study of the Apocynoideae (with special reference to the genus Apocynum). Annals of the Missouri Botanical Garden 17: 212.
  • Woodson RE Jr. 1933. Studies in the Apocynaceae IV. The American genera of Echitoideae. Annals of the Missouri Botanical Garden 20: 605-790.

Publication Dates

  • Publication in this collection
    27 Oct 2023
  • Date of issue
    2023

History

  • Received
    20 May 2023
  • Accepted
    08 Aug 2023
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