Abstract
A new species of Borreria from savanna vegetation found within the Caatinga in northwestern Ceará, Brazil, is described and illustrated. Borreria savannicola has an affinity with B. scabiosoides, sharing with it the habitat preference and similar morphological characteristics such as corolla infundibuliform, stigma bilobed and shape of seeds. Nevertheless, the new species differs mainly in its erect habit, linear leaves, terminal glomerule with up to 10 involucral bracts, and obovoid capsule. Images of anatomical characters, electro microphotography of flower, fruit, seed, and pollen, together with photographs of the habitat, distribution map and conservation status are provided.
References
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<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Carmo, J.A.M., Reginato, M., Florentín, J.E., Nuñez-Florentin, M., Salas, R.M. & Simões, A.O. (2021) One more piece to the puzzle: <em>Diadorimia</em>, a new monotypic genus in the Spermacoceae (Rubiaceae), endemic to the campo rupestre of Minas Gerais, southeastern Brazil. <em>Taxon </em>71 (2): 396–419. https://doi.org/10.1002/tax.12643</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Dessein, S. (2003) Systematic studies in the Spermacoceae (Rubiaceae). PhD Thesis, Leuven, Belgium.</span></span></span></p>
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<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Groeninckx, I., Dessein, S., Ochoterena, H., Persson, C., Motley, T.J., Kårehed, J., Bremer, J., Huysmans, S. & Smets, E. (2009) Phylogeny of the herbaceous tribe Spermacoceae (Rubiaceae) based on plastid DNA data. <em>Annals of the Missouri Botanical Garden</em> 96: 109–132. https://doi.org/10.3417/2006201</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">IUCN (2022) The IUCN Red List of Threatened Species. Version 2021–3. [https://www.iucnredlist.org] </span></span></span></p>
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<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Kårehed, J., Groeninckx, I., Dessein, S., Motley, T.J. & Bremer, J. (2008) The phylogenetic utility of chloroplast and nuclear DNA markers and the phylogeny of the Rubiaceae tribe Spermacoceae. <em>Molecular Phylogenetics and Evolution</em> 49: 843–866. https://doi.org/10.1016/j.ympev.2008.09.025</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Meyer, G. (1818) <em>Borreria </em>Meyer. <em>Primitiae Florae Essequeboensis</em> 79–83, pl. 1.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M. (2016) Estudios biosistemáticos en especies sudamericanas de <em>Borreria</em> subsecc. <em>Borreria</em> y filogenia de las especies del complejo <em>Spermacoce–Borreria</em> (Rubiaceae). Tesis Doctoral. Facultad de Ciencias Exactas y Naturales y Agrimensura. Universidad Nacional del Nordeste. Corrientes, Argentina, 375 pp.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M. & Cabral, E.L. (2013) <em>Borreria krapocarmeniana</em>, a new cryptic species recovered through taxonomic analyses of <em>Borreria scabiosoides</em> and <em>Borreria linoides </em>(Spermacoceae, Rubiaceae). <em>Systematic Botany</em> 38: 769–781. https://doi.org/10.1600/036364413X670368</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M., Sobrado, S.V., Janssens, S., Dessein, S. & Cabral, E.L. (2018) The monotypic Brazilian genus <em>Diacrodon</em> is a synonym of <em>Borreria</em> (Spermacoceae, Rubiaceae): morphological and molecular evidences. <em>Anais da Academia Brasileira de Ciências </em>90 (2): 1397–1415. https://doi.org/10.1590/0001-3765201820170314</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M., Sobrado, S.V., Cabral, E.L., Salas, R.M., Souza, E.B., Florentín, J.E., Nicora–Chequin, R., Fader, A.A.C., Nepomuceno, F.A.A., Carmo, J.A.M. & Nuñez-Florentin, M. (2020) <em>Borreria</em> <em>in</em> Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Disponível em: http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB20690 (Acesssed 30 November 2021)</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nepomuceno, F.A.A., Souza, E.B., Nepomuceno, I.V., Miguel, L.M., Cabral, E.L. & Loiola, M.I.B. (2018) The genus <em>Borreria </em>(Spermacoceae, Rubiaceae) in the state of Ceará, Brazil. <em>Rodriguésia</em> 69: 715–731. https://doi.org/10.1590/2175-7860201869232</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nepomuceno, I.V., Souza, E.B., Zappi, D.C., Moreira, M.C., Nepomuceno, F.A.A. & Moro, M.F. (2021) Savannas of the Brazilian semiarid region: what do we learn from floristics. <em>Acta Botanica Brasilica</em> 35 (3): 361–380. https://doi.org/10.1590/0102-33062020abb0259</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nuñez–Florentin, M., Salas, R.M., Janssens, S., Dessein, S. & Cardoso, D. (2021) Molecular–based phylogenetic placement and revision of <em>Micrasepalum</em> (Spermacoceae–Rubiaceae). <em>Taxon</em> 70 (6): 1300–1316. https://doi.org/10.1002/tax.12593</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Punt, W., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007) Glossary of pollen and spore terminology. <em>Review of Palaeobotany and Palynology</em> 143: 1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">QGIS Development Team. (2022) QGIS Geographic Information System. Open Source Geospatial Foundation Project.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Rôças, G., Barros, C.F. & Scarano, F.R. (1997) Leaf anatomy plasticity of <em>Alchornea triplinervia</em> (Euphorbiaceae) under distinct light regimes in a Brazilian Montane Atlantic Rain Forest. <em>Trees</em> 11 (8): 469–473. https://doi.org/10.1007/PL00009688</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Salas, R.M., Viana, P.L., Cabral, E.L., Dessien, S. & Janssens, S. (2015) <em>Carajasia </em>(Rubiaceae), a new and endangered genus from Carajás mountain range, Pará, Brazil. <em>Phytotaxa</em> 206: 14–29. https://doi.org/10.11646/phytotaxa.206.1.4</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sobrado, S.V. (2016) Estudios biosistemáticos en especies de <em>Borreria</em> (Spermacoceae–Rubiaceae) con énfasis en <em>Borreria</em> subsecc. <em>Latifoliae</em>. Tesis Doctoral. Facultad de Ciencias Exactas y Naturales y Agrimensura. Universidad Nacional del Nordeste. Corrientes, Argentina, 283 pp.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Souza, E.B., Miguel, L.M., Cabral, E.L., Nepomuceno, F.A.A. & Loiola, M.I.B. (2016) <em>Borreria apodiensis</em> (Rubiaceae: Spermacoceae), a new species from Ceará and Rio Grande do Norte, Brazil. <em>Acta Botanica Brasilica</em> 30 (2): 283–289. https://doi.org/10.1590/0102-33062016abb0014</span></span></span></p>
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<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zini, A.S., Martins, S., Toderke, M.L. & Temponi, L.G. (2016) Anatomia foliar de Rubiaceae ocorrentes em fragmento florestal urbano de Mata Atlântica, PR, Brasil. <em>Hoehnea</em> 43 (2): 173–182. https://doi.org/10.1590/2236-8906-59/2015</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Bacigalupo, N.M. & Cabral, E.L. (1996) Infrageneric classification of <em>Borreria </em>(Rubiaceae–Spermacoceae) on the basis of American species. <em>Opera Botanica Belgica</em> 7: 297–308.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Barros, I.O. & Soares, A.A. (2013) Adaptações anatômicas em folhas de marmeleiro e velame da caatinga brasileira. <em>Revista Ciência Agronômica</em> 44: 192–198. https://doi.org/10.1590/S1806-66902013000100024</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Cabral, E.L., Miguel, L.M. & Salas, R.M. (2011) Dos especies nuevas de <em>Borreria </em>(Rubiaceae), sinopsis y clave de las especies para Bahia, Brasil. <em>Acta Botanica Brasilica</em> 25: 255–276. https://doi.org/10.1590/S0102-33062011000200002</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Carmo, J.A.M., Reginato, M., Florentín, J.E., Nuñez-Florentin, M., Salas, R.M. & Simões, A.O. (2021) One more piece to the puzzle: <em>Diadorimia</em>, a new monotypic genus in the Spermacoceae (Rubiaceae), endemic to the campo rupestre of Minas Gerais, southeastern Brazil. <em>Taxon </em>71 (2): 396–419. https://doi.org/10.1002/tax.12643</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Dessein, S. (2003) Systematic studies in the Spermacoceae (Rubiaceae). PhD Thesis, Leuven, Belgium.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Dickson, W.C. (2000) <em>Integrative Plant Anatomy.</em> Academic Press, Florida, 492 pp.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Erdtman, G. (1966) <em>Pollen Morphology and Plant Taxonomy: Angiosperms. An introduction to Palynology–I.</em> New York and London, Hafner Publishing Company.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Florentín, J.E., Salas, R.M., Miguel, L.M. & Cabral, E.L. (2016) Taxonomía de <em>Spermacoce eryngioides</em> (Rubiaceae) y transferencia de <em>Borreria</em> secc. <em>Pseudodiodia</em> a <em>Spermacoce</em>. <em>Boletín de la Sociedad Argentina de Botánica</em> 51 (3): 551–564. https://doi.org/10.31055/1851.2372.v51.n3.15393</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Florentín, J.E., Cabaña Fader, A.A., Salas, R.M., Janssens, S. & Cabral, E.L. (2017) Morphological and molecular data confirm the transfer of homostylous species in the typically distylous genus <em>Galianthe</em> (Rubiaceae), and description of the new species <em>Galianthe</em> <em>vasquezii</em> from Peru and Colombia. <em>PeerJ.</em> 5: e4012 [1–23]. https://doi.org/10.7717/peerj.4012</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Groeninckx, I., Dessein, S., Ochoterena, H., Persson, C., Motley, T.J., Kårehed, J., Bremer, J., Huysmans, S. & Smets, E. (2009) Phylogeny of the herbaceous tribe Spermacoceae (Rubiaceae) based on plastid DNA data. <em>Annals of the Missouri Botanical Garden</em> 96: 109–132. https://doi.org/10.3417/2006201</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">IUCN (2022) The IUCN Red List of Threatened Species. Version 2021–3. [https://www.iucnredlist.org] </span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Kraus, J.E. & Arduin, M. (1997) <em>Manual básico de métodos em morfologia vegetal.</em> Edur, Seropédica, 198 pp.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Kårehed, J., Groeninckx, I., Dessein, S., Motley, T.J. & Bremer, J. (2008) The phylogenetic utility of chloroplast and nuclear DNA markers and the phylogeny of the Rubiaceae tribe Spermacoceae. <em>Molecular Phylogenetics and Evolution</em> 49: 843–866. https://doi.org/10.1016/j.ympev.2008.09.025</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Meyer, G. (1818) <em>Borreria </em>Meyer. <em>Primitiae Florae Essequeboensis</em> 79–83, pl. 1.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M. (2016) Estudios biosistemáticos en especies sudamericanas de <em>Borreria</em> subsecc. <em>Borreria</em> y filogenia de las especies del complejo <em>Spermacoce–Borreria</em> (Rubiaceae). Tesis Doctoral. Facultad de Ciencias Exactas y Naturales y Agrimensura. Universidad Nacional del Nordeste. Corrientes, Argentina, 375 pp.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M. & Cabral, E.L. (2013) <em>Borreria krapocarmeniana</em>, a new cryptic species recovered through taxonomic analyses of <em>Borreria scabiosoides</em> and <em>Borreria linoides </em>(Spermacoceae, Rubiaceae). <em>Systematic Botany</em> 38: 769–781. https://doi.org/10.1600/036364413X670368</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M., Sobrado, S.V., Janssens, S., Dessein, S. & Cabral, E.L. (2018) The monotypic Brazilian genus <em>Diacrodon</em> is a synonym of <em>Borreria</em> (Spermacoceae, Rubiaceae): morphological and molecular evidences. <em>Anais da Academia Brasileira de Ciências </em>90 (2): 1397–1415. https://doi.org/10.1590/0001-3765201820170314</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miguel, L.M., Sobrado, S.V., Cabral, E.L., Salas, R.M., Souza, E.B., Florentín, J.E., Nicora–Chequin, R., Fader, A.A.C., Nepomuceno, F.A.A., Carmo, J.A.M. & Nuñez-Florentin, M. (2020) <em>Borreria</em> <em>in</em> Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Disponível em: http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB20690 (Acesssed 30 November 2021)</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nepomuceno, F.A.A., Souza, E.B., Nepomuceno, I.V., Miguel, L.M., Cabral, E.L. & Loiola, M.I.B. (2018) The genus <em>Borreria </em>(Spermacoceae, Rubiaceae) in the state of Ceará, Brazil. <em>Rodriguésia</em> 69: 715–731. https://doi.org/10.1590/2175-7860201869232</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nepomuceno, I.V., Souza, E.B., Zappi, D.C., Moreira, M.C., Nepomuceno, F.A.A. & Moro, M.F. (2021) Savannas of the Brazilian semiarid region: what do we learn from floristics. <em>Acta Botanica Brasilica</em> 35 (3): 361–380. https://doi.org/10.1590/0102-33062020abb0259</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Nuñez–Florentin, M., Salas, R.M., Janssens, S., Dessein, S. & Cardoso, D. (2021) Molecular–based phylogenetic placement and revision of <em>Micrasepalum</em> (Spermacoceae–Rubiaceae). <em>Taxon</em> 70 (6): 1300–1316. https://doi.org/10.1002/tax.12593</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Punt, W., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007) Glossary of pollen and spore terminology. <em>Review of Palaeobotany and Palynology</em> 143: 1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">QGIS Development Team. (2022) QGIS Geographic Information System. Open Source Geospatial Foundation Project.</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Rôças, G., Barros, C.F. & Scarano, F.R. (1997) Leaf anatomy plasticity of <em>Alchornea triplinervia</em> (Euphorbiaceae) under distinct light regimes in a Brazilian Montane Atlantic Rain Forest. <em>Trees</em> 11 (8): 469–473. https://doi.org/10.1007/PL00009688</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Salas, R.M., Viana, P.L., Cabral, E.L., Dessien, S. & Janssens, S. (2015) <em>Carajasia </em>(Rubiaceae), a new and endangered genus from Carajás mountain range, Pará, Brazil. <em>Phytotaxa</em> 206: 14–29. https://doi.org/10.11646/phytotaxa.206.1.4</span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sobrado, S.V. (2016) Estudios biosistemáticos en especies de <em>Borreria</em> (Spermacoceae–Rubiaceae) con énfasis en <em>Borreria</em> subsecc. <em>Latifoliae</em>. Tesis Doctoral. Facultad de Ciencias Exactas y Naturales y Agrimensura. Universidad Nacional del Nordeste. Corrientes, Argentina, 283 pp.</span></span></span></p>
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