Abstract
A distinctive species, Vanilla denshikoira, is described from the North West Amazon, in Colombia, within the Guiana Shield region. The species has morphological features similar to those of species in the Vanilla planifolia group. It is an important addition to the vanilla crop wild relatives, bringing the total number of species in the V. planifolia group to 21. Vanilla denshikoira is a narrow endemic, known from only a single locality, and highly vulnerable to anthropological disturbance. Under IUCN criteria it is categorized CR. The species has potential value as a non-timber forest product. We recommend a conservation program that includes support for circa situm actions implemented by the local communities.
References
Ames, O. (1934) An addition to the genus Vanilla. Botanical Museum Leaflets 2: 101–103.
Andrews, H.C. (1808) Vanilla planifolia. Botanist’s Repository, for new, and rare plants 8: 552.
Aublet, J.B.C.F. (1775) Histoire des Plantes de la Guiane Françoise. Didot, Paris, 621 pp.
Azofeifa-Bolaños, J.B., Gigant, L.R., Nicolás-García, M., Pignal, M., Tavares-González, F.B., Hágsater, E., Salazar-Chávez, G.A., Reyes-López,D., Archila-Morales, F.L., García-García, J.A., da Silva, D., Allibert, A., Solano-Campos, F., Rodríguez-Jimenes, G.C. Paniagua-Vásquez, A., Besse, P., Pérez-Silva, A. & Grisoni, M. (2017) A new Vanilla species from Costa Rica closely related to V. planifolia (Orchidaceae). European Journal of Taxonomy 284: 1–26.
https://doi.org/10.5852/ejt.2017.284
Barros, F. de, Vinhos, F., Rodrigues, V.T., Barberena, F.F.V.A., Fraga, C.N., Pessoa, E.M., Forster, W., Menini Neto, L., Furtado, S.G., Nardy, C., Azevedo, C.O. & Guimarães, L.R.S. (2016) Orchidaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB38415 (accessed 5 October 2017)
Besse, P., Da Silva, D., Bory, S., Grisoni, M., Le Bellec, F. & Duval, M.F. (2004) RAPD genetic diversity in cultivated vanilla: Vanilla planifolia, and relationships with V. tahitensis and V. pompona. Plant Science 167: 379–385.
https://doi.org/10.1016/j.plantsci.2004.04.007
Bory, S., Lubinsky, P., Risterucci, A.M., Noyer, J.L., Grisoni, M., Duval, M.F. & Besse, P. (2008) Patterns of introduction and diversification of Vanilla planifolia (Orchidaceae) in Reunion Island (Indian Ocean). American Journal of Botany 95: 805–815.
https://doi.org/10.3732/ajb.2007332
Calderón-Sáenz, E. (Ed.) (2007) Libro Rojo de Plantas de Colombia. In: Volumen 6. Orquídeas primer parte. Instituto Alexander von Humboldt—Ministerio de Ambiente, Vivienda y Desarrollo Territorial. Bogotá, Colombia, 828 pp. Available from: <a href="http://repository.humboldt.org.co/handle/20.500.11761/31410"><span style="text-decoration: underline;">http://repository.humboldt.org.co/handle/20.500.11761/31410</span></a> (accessed 5 October 2017)
Cárdenas López, D., Castaño Arboleda, N. & Sua Tunjano, S. (2009) Flora de la Estrella Fluvial de Inírida (Guainía, Colombia). Biota Colombiana 10: 1–30.
Dawson, I.K., Guariguata, M.R., Loo, J., Weber, J.C., Lengkeek, A., Bush, D., Cornelius, J., Guarino, L., Kindt, R., Orwa, C., Russell, J. & Jamnadass, R. (2013) What is the relevance of smallholders’ agroforestry systems for conserving tropical tree species and genetic diversity in circa situm, in situ and ex situ settings? A review. Biodiversity and Conservation 22: 301–324.
https://doi.org/10.1007/s10531-012-0429-5
Etter, A. (2001) Puinawai y Nukak: Caracterización ecológica de dos reservas nacionales naturales de la Amazonía colombiana (Vol. 2). IDEADE, Pontificia Universidad Javeriana-Serie Ambiente y Desarrollo 2, Bogotá, 330 pp.
Flanagan, N.S., Otero, J.T., Molineros, F.H., Mosquera-Espinosa, A.T., Vásquez, E., Garnica, S., Rosero, D., Parra, S., Martínez, C. & Palma, R. (2012) Aprovechamiento sostenible de Recursos Biológicos Nativos del Distrito de Manejo Integrado de Atuncela. Guía Práctica. Cartilla de los Recursos Biológicos del Distrito de Manejo Integrado de Atuncela. Proyecto de CVC No 0051 y la Pontificia Universidad Javeriana, Cali, Colombia, 31 pp.
Flanagan, N.S. & Mosquera-Espinosa, A.T. (2016) An integrated strategy for the conservation and sustainable use of native Vanilla species in Colombia. Lankesteriana 16: 201–218.
https://doi.org/10.15517/lank.v16i2.26007
Flanagan, N.S., Chavarriaga, P. & Mosquera-Espinosa, A.T. (2018) Conservation and sustainable use of Vanilla crop wild relatives in Colombia. In: Havkin-Frenkel, D. & Belanger, F.C. (Eds.) Handbook of vanilla science and technology, Second Edition. Wiley-Blackwell, Oxford, UK, pp. 85–110.
https://doi.org/10.1002/9781119377320.ch6
Fraga, de C.N. (2002) Notas taxonômicas para as espécies brasileiras de Vanilla Mill. (Orchidaceae). Boletim do Museu de Biologia Prof. Mello Leitão (N. Série) 13: 45–52.
Fraga, de C.N., Couto, D.R. & Pansarin, E.R. (2017) Two new species of Vanilla (Orchidaceae) in the Brazilian Atlantic Forest. Phytotaxa 296 (1): 63–72.
https://doi.org/10.11646/phytotaxa.296.1.4
Hajjar, R. & Hodgkin, T. (2007) The use of wild relatives in crop improvement: a survey of developments over the last 20 years. Euphytica 156: 1–13.
https://doi.org/10.1007/s10681-007-9363-0
Hernández-Hernández, J. (2011) Vanilla Diseases In: Havkin-Frenkel, D. & Belanger, F.C. (Eds.) Handbook of Vanilla Science and Technology, First Edition. Wiley-Blackwell, Oxford, UK, pp. 26–39.
https://doi.org/10.1002/9781444329353.ch2
Heywood, V., Casas, A., Ford-Lloyd, B., Kell, S. & Maxted, N. (2007) Conservation and sustainable use of crop wild relatives. Agriculture, Ecosystems & Environment 121: 245–255.
https://doi.org/10.1016/j.agee.2006.12.014
Hoehne, F.C. (1944) Orchidaceae novas para a flora do Brasil, dos Herbaìrios do Inst. de Bota?nica, Jardim Bota?nico, Rio de Janeiro e ComissaÞo de Linhas Telegraìficas Estrateìgicas, de Mato Grosso ao Amazonas. Arquivos de Botânica do Estado de São Paulo 1: 125–134.
Humboldt, F.W.H.J., von Bonpland, A.J.A. & Kunth, K.S. (1821) Nova Genera et Species Plantarum (quarto ed.). Chez N. Maze Libraire, París, 512 pp.
IUCN Standards and Petitions Subcommittee (2017) Guidelines for using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. Available from: <a href="http://www.iucnredlist.org/documents/RedListGuidelines.pdf"><span style="text-decoration: underline;">http://www.iucnredlist.org/documents/RedListGuidelines.pdf</span></a> (accessed 5 October 2017)
Koch, A.K., Fraga, de C.N., Santos, J.U.M.D. & Ilkiu-Borges, A.L. (2013) Taxonomic notes on Vanilla (Orchidaceae) in the Brazilian Amazon, and the description of a new species. Systematic Botany 38: 975–981.
https://doi.org/10.1600/036364413X674706
Lindley, J. (1838) Miscellaneous notices. Edward´s Botanical Register 24 (Misc.): 1–95.
Linnaeus, C. (1781 [1782]) Supplementum Plantarum Systematis Vegetabilium Editionis Decimae Tertiae, Generum Plantarum Editiones Sextae, et Specierum Plantarum Editionis Secundae. Editum a Carolo a Linné. Impensis Orphanotrophei, Brunsvigae, 468 pp.
Martius, C.F.P. (1888) Flora Brasiliensis, 12 (1). Monachii, Frid. Fleischer, 522 pp.
Molineros-Hurtado, F.H., González-Mina, R.T., Flanagan, N.S. & Otero, J.T. (2014) Vanilla rivasii (Orchidaceae), a new species from the Colombian pacific region. Lankesteriana 13: 353–357.
https://doi.org/10.15517/lank.v13i3.14423
Orbigny, A.D. d´ (1842) Voyage dans l’Amérique Méridionale, 7 (3). Chez P. Bertrand, Paris, 140 pp.
Ospina-Calderón, N.H., Mendoza,<sup>, </sup>M., García Agapito, L.T., Mateus, H.A., Vásquez, J., Delgado Mazuera, C., Mosquera-Espinosa, A.T. & Flanagan, N.S. (2017) Guía a las Orquídeas de la Estrella Fluvial de Inírida, Guainía, norte de Amazonas, Colombia. Available from: http://fieldguides.fieldmuseum.org/guides/guide/932 (accessed 5 October 2017)
Pennington T.D. (1990) Flora Neotropica. Monograph 52. Sapotaceae. New York Botanical Garden for the Organization for Flora Neotropica, New York, 770 pp.
Presl, C.B. (1830) Reliquiae Haenkeanae seu, Descriptiones et icones plantarum quas in America Meridionali et Boreali 1: 101.
Reichenbach, H.G. (1865) Vorstudien zu einer Orchidographie der Antillen, besonders britischen Antheils. Neuheiten und eingehende Critiken bisheriger Literatur. Flora XLVIII: 273–280.
Rolfe, R.A. (1895) Vanillas of commerce. Bulletin of Miscellaneous Information Kew 104: 169–178.
Rolfe, R.A. (1896) A revision of the genus Vanilla. Journal of the Linnaean Society 32: 439–478.
https://doi.org/10.1111/j.1095-8339.1896.tb00702.x
Roux-Cuvelier, M. & Grisoni, M. (2011) Conservation and movement of Vanilla germplasm. In: Odoux, E. & Grisoni, M. (Eds.) Vanilla. CRC Press, Boca Ratón, pp. 31–41.
Sambin, A. & Chiron, G.R. (2015) Deux nouvelles espèces de Vanilla (Orchidaceae) de Guyane française. Richardiana 15: 306–316.
Schiede, C.J.W. (1829) Botanische Berichte aus Mexico. Linnaea 4: 554–583.
Schluter, P.M., Soto Arenas, M.A. & Harris, S.A. (2007) Genetic variation in Vanilla planifolia (Orchidaceae). Economic Botany 61: 328–336.
<a href="https://doi.org/10.1663/0013-0001(2007)61%5b328:GVIVPO%5d2.0.CO;2">https://doi.org/10.1663/0013-0001(2007)61[328:GVIVPO]2.0.CO;2</a>
Soto Arenas, M.A. (2003) Vanilla. In: Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (Eds.) Genera orchidacearum: Orchidoideae (part 2), Vanilloideae. Oxford University, New York, pp. 321–334.
Soto Arenas, M.A. (2010) A new species of Vanilla from South America. Lankesteriana 9: 281–284.
https://doi.org/10.15517/lank.v0i0.12020
Soto Arenas, M.A. & Cribb, P. (2010) A new infrageneric classification and synopsis of the genus Vanilla Plum. Ex. Mill. (Orchidaceae: Vanillinae). Lankesteriana 9: 355–398.
https://doi.org/10.15517/lank.v0i0.12071
Soto Arenas, M.A. & Dressler, R.L. (2010) A revision of the Mexican and Central American species of Vanilla Plumier ex Miller with a characterization of their ITS region of the nuclear ribosomal DNA. Lankesteriana 9: 285–354.
https://doi.org/10.15517/lank.v0i0.12065
Wallace, A.R. (1853) Palm Trees of the Amazon and their uses. John van Voorst, London, 129 pp.
https://doi.org/10.5962/bhl.title.11168
Xiong, W., Zhao, Q., Zhao, J., Xun, W., Li, R., Zhang, R., Wu, H. & Shen, Q. (2015) Different continuous cropping spans significantly affect microbial community membership and structure in a vanilla-grown soil as revealed by deep pyrosequencing. Microbial Ecology, 70 (1), 209–218.
https://doi.org/10.1007/s00248-014-0516-0
Andrews, H.C. (1808) Vanilla planifolia. Botanist’s Repository, for new, and rare plants 8: 552.
Aublet, J.B.C.F. (1775) Histoire des Plantes de la Guiane Françoise. Didot, Paris, 621 pp.
Azofeifa-Bolaños, J.B., Gigant, L.R., Nicolás-García, M., Pignal, M., Tavares-González, F.B., Hágsater, E., Salazar-Chávez, G.A., Reyes-López,D., Archila-Morales, F.L., García-García, J.A., da Silva, D., Allibert, A., Solano-Campos, F., Rodríguez-Jimenes, G.C. Paniagua-Vásquez, A., Besse, P., Pérez-Silva, A. & Grisoni, M. (2017) A new Vanilla species from Costa Rica closely related to V. planifolia (Orchidaceae). European Journal of Taxonomy 284: 1–26.
https://doi.org/10.5852/ejt.2017.284
Barros, F. de, Vinhos, F., Rodrigues, V.T., Barberena, F.F.V.A., Fraga, C.N., Pessoa, E.M., Forster, W., Menini Neto, L., Furtado, S.G., Nardy, C., Azevedo, C.O. & Guimarães, L.R.S. (2016) Orchidaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB38415 (accessed 5 October 2017)
Besse, P., Da Silva, D., Bory, S., Grisoni, M., Le Bellec, F. & Duval, M.F. (2004) RAPD genetic diversity in cultivated vanilla: Vanilla planifolia, and relationships with V. tahitensis and V. pompona. Plant Science 167: 379–385.
https://doi.org/10.1016/j.plantsci.2004.04.007
Bory, S., Lubinsky, P., Risterucci, A.M., Noyer, J.L., Grisoni, M., Duval, M.F. & Besse, P. (2008) Patterns of introduction and diversification of Vanilla planifolia (Orchidaceae) in Reunion Island (Indian Ocean). American Journal of Botany 95: 805–815.
https://doi.org/10.3732/ajb.2007332
Calderón-Sáenz, E. (Ed.) (2007) Libro Rojo de Plantas de Colombia. In: Volumen 6. Orquídeas primer parte. Instituto Alexander von Humboldt—Ministerio de Ambiente, Vivienda y Desarrollo Territorial. Bogotá, Colombia, 828 pp. Available from: <a href="http://repository.humboldt.org.co/handle/20.500.11761/31410"><span style="text-decoration: underline;">http://repository.humboldt.org.co/handle/20.500.11761/31410</span></a> (accessed 5 October 2017)
Cárdenas López, D., Castaño Arboleda, N. & Sua Tunjano, S. (2009) Flora de la Estrella Fluvial de Inírida (Guainía, Colombia). Biota Colombiana 10: 1–30.
Dawson, I.K., Guariguata, M.R., Loo, J., Weber, J.C., Lengkeek, A., Bush, D., Cornelius, J., Guarino, L., Kindt, R., Orwa, C., Russell, J. & Jamnadass, R. (2013) What is the relevance of smallholders’ agroforestry systems for conserving tropical tree species and genetic diversity in circa situm, in situ and ex situ settings? A review. Biodiversity and Conservation 22: 301–324.
https://doi.org/10.1007/s10531-012-0429-5
Etter, A. (2001) Puinawai y Nukak: Caracterización ecológica de dos reservas nacionales naturales de la Amazonía colombiana (Vol. 2). IDEADE, Pontificia Universidad Javeriana-Serie Ambiente y Desarrollo 2, Bogotá, 330 pp.
Flanagan, N.S., Otero, J.T., Molineros, F.H., Mosquera-Espinosa, A.T., Vásquez, E., Garnica, S., Rosero, D., Parra, S., Martínez, C. & Palma, R. (2012) Aprovechamiento sostenible de Recursos Biológicos Nativos del Distrito de Manejo Integrado de Atuncela. Guía Práctica. Cartilla de los Recursos Biológicos del Distrito de Manejo Integrado de Atuncela. Proyecto de CVC No 0051 y la Pontificia Universidad Javeriana, Cali, Colombia, 31 pp.
Flanagan, N.S. & Mosquera-Espinosa, A.T. (2016) An integrated strategy for the conservation and sustainable use of native Vanilla species in Colombia. Lankesteriana 16: 201–218.
https://doi.org/10.15517/lank.v16i2.26007
Flanagan, N.S., Chavarriaga, P. & Mosquera-Espinosa, A.T. (2018) Conservation and sustainable use of Vanilla crop wild relatives in Colombia. In: Havkin-Frenkel, D. & Belanger, F.C. (Eds.) Handbook of vanilla science and technology, Second Edition. Wiley-Blackwell, Oxford, UK, pp. 85–110.
https://doi.org/10.1002/9781119377320.ch6
Fraga, de C.N. (2002) Notas taxonômicas para as espécies brasileiras de Vanilla Mill. (Orchidaceae). Boletim do Museu de Biologia Prof. Mello Leitão (N. Série) 13: 45–52.
Fraga, de C.N., Couto, D.R. & Pansarin, E.R. (2017) Two new species of Vanilla (Orchidaceae) in the Brazilian Atlantic Forest. Phytotaxa 296 (1): 63–72.
https://doi.org/10.11646/phytotaxa.296.1.4
Hajjar, R. & Hodgkin, T. (2007) The use of wild relatives in crop improvement: a survey of developments over the last 20 years. Euphytica 156: 1–13.
https://doi.org/10.1007/s10681-007-9363-0
Hernández-Hernández, J. (2011) Vanilla Diseases In: Havkin-Frenkel, D. & Belanger, F.C. (Eds.) Handbook of Vanilla Science and Technology, First Edition. Wiley-Blackwell, Oxford, UK, pp. 26–39.
https://doi.org/10.1002/9781444329353.ch2
Heywood, V., Casas, A., Ford-Lloyd, B., Kell, S. & Maxted, N. (2007) Conservation and sustainable use of crop wild relatives. Agriculture, Ecosystems & Environment 121: 245–255.
https://doi.org/10.1016/j.agee.2006.12.014
Hoehne, F.C. (1944) Orchidaceae novas para a flora do Brasil, dos Herbaìrios do Inst. de Bota?nica, Jardim Bota?nico, Rio de Janeiro e ComissaÞo de Linhas Telegraìficas Estrateìgicas, de Mato Grosso ao Amazonas. Arquivos de Botânica do Estado de São Paulo 1: 125–134.
Humboldt, F.W.H.J., von Bonpland, A.J.A. & Kunth, K.S. (1821) Nova Genera et Species Plantarum (quarto ed.). Chez N. Maze Libraire, París, 512 pp.
IUCN Standards and Petitions Subcommittee (2017) Guidelines for using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. Available from: <a href="http://www.iucnredlist.org/documents/RedListGuidelines.pdf"><span style="text-decoration: underline;">http://www.iucnredlist.org/documents/RedListGuidelines.pdf</span></a> (accessed 5 October 2017)
Koch, A.K., Fraga, de C.N., Santos, J.U.M.D. & Ilkiu-Borges, A.L. (2013) Taxonomic notes on Vanilla (Orchidaceae) in the Brazilian Amazon, and the description of a new species. Systematic Botany 38: 975–981.
https://doi.org/10.1600/036364413X674706
Lindley, J. (1838) Miscellaneous notices. Edward´s Botanical Register 24 (Misc.): 1–95.
Linnaeus, C. (1781 [1782]) Supplementum Plantarum Systematis Vegetabilium Editionis Decimae Tertiae, Generum Plantarum Editiones Sextae, et Specierum Plantarum Editionis Secundae. Editum a Carolo a Linné. Impensis Orphanotrophei, Brunsvigae, 468 pp.
Martius, C.F.P. (1888) Flora Brasiliensis, 12 (1). Monachii, Frid. Fleischer, 522 pp.
Molineros-Hurtado, F.H., González-Mina, R.T., Flanagan, N.S. & Otero, J.T. (2014) Vanilla rivasii (Orchidaceae), a new species from the Colombian pacific region. Lankesteriana 13: 353–357.
https://doi.org/10.15517/lank.v13i3.14423
Orbigny, A.D. d´ (1842) Voyage dans l’Amérique Méridionale, 7 (3). Chez P. Bertrand, Paris, 140 pp.
Ospina-Calderón, N.H., Mendoza,<sup>, </sup>M., García Agapito, L.T., Mateus, H.A., Vásquez, J., Delgado Mazuera, C., Mosquera-Espinosa, A.T. & Flanagan, N.S. (2017) Guía a las Orquídeas de la Estrella Fluvial de Inírida, Guainía, norte de Amazonas, Colombia. Available from: http://fieldguides.fieldmuseum.org/guides/guide/932 (accessed 5 October 2017)
Pennington T.D. (1990) Flora Neotropica. Monograph 52. Sapotaceae. New York Botanical Garden for the Organization for Flora Neotropica, New York, 770 pp.
Presl, C.B. (1830) Reliquiae Haenkeanae seu, Descriptiones et icones plantarum quas in America Meridionali et Boreali 1: 101.
Reichenbach, H.G. (1865) Vorstudien zu einer Orchidographie der Antillen, besonders britischen Antheils. Neuheiten und eingehende Critiken bisheriger Literatur. Flora XLVIII: 273–280.
Rolfe, R.A. (1895) Vanillas of commerce. Bulletin of Miscellaneous Information Kew 104: 169–178.
Rolfe, R.A. (1896) A revision of the genus Vanilla. Journal of the Linnaean Society 32: 439–478.
https://doi.org/10.1111/j.1095-8339.1896.tb00702.x
Roux-Cuvelier, M. & Grisoni, M. (2011) Conservation and movement of Vanilla germplasm. In: Odoux, E. & Grisoni, M. (Eds.) Vanilla. CRC Press, Boca Ratón, pp. 31–41.
Sambin, A. & Chiron, G.R. (2015) Deux nouvelles espèces de Vanilla (Orchidaceae) de Guyane française. Richardiana 15: 306–316.
Schiede, C.J.W. (1829) Botanische Berichte aus Mexico. Linnaea 4: 554–583.
Schluter, P.M., Soto Arenas, M.A. & Harris, S.A. (2007) Genetic variation in Vanilla planifolia (Orchidaceae). Economic Botany 61: 328–336.
<a href="https://doi.org/10.1663/0013-0001(2007)61%5b328:GVIVPO%5d2.0.CO;2">https://doi.org/10.1663/0013-0001(2007)61[328:GVIVPO]2.0.CO;2</a>
Soto Arenas, M.A. (2003) Vanilla. In: Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (Eds.) Genera orchidacearum: Orchidoideae (part 2), Vanilloideae. Oxford University, New York, pp. 321–334.
Soto Arenas, M.A. (2010) A new species of Vanilla from South America. Lankesteriana 9: 281–284.
https://doi.org/10.15517/lank.v0i0.12020
Soto Arenas, M.A. & Cribb, P. (2010) A new infrageneric classification and synopsis of the genus Vanilla Plum. Ex. Mill. (Orchidaceae: Vanillinae). Lankesteriana 9: 355–398.
https://doi.org/10.15517/lank.v0i0.12071
Soto Arenas, M.A. & Dressler, R.L. (2010) A revision of the Mexican and Central American species of Vanilla Plumier ex Miller with a characterization of their ITS region of the nuclear ribosomal DNA. Lankesteriana 9: 285–354.
https://doi.org/10.15517/lank.v0i0.12065
Wallace, A.R. (1853) Palm Trees of the Amazon and their uses. John van Voorst, London, 129 pp.
https://doi.org/10.5962/bhl.title.11168
Xiong, W., Zhao, Q., Zhao, J., Xun, W., Li, R., Zhang, R., Wu, H. & Shen, Q. (2015) Different continuous cropping spans significantly affect microbial community membership and structure in a vanilla-grown soil as revealed by deep pyrosequencing. Microbial Ecology, 70 (1), 209–218.
https://doi.org/10.1007/s00248-014-0516-0