Abstract
A morphological and molecular study was carried out on two species of Carex from the former genus Uncinia from mainland South America and Juan Fernández Islands: C. firmula (Kük) J.R. Starr and C. subsacculata (G.A.Wheeler & Goetgh.) J.R.Starr. Both species share the particular character of having articulated pistillate scales that leave behind on the spike rachis a saccate stipe-like appendage after falling. We detected within C. firmula morphological differences between mainland South American specimens and Juan Fernández specimens, mainly regarding flower density and utricle size. On the contrary, the molecular analysis showed no molecular differentiation between the two groups of populations. Whereas, strong molecular and morphological support were found for C. subsacculata. As such, we recommend that the populations of C. firmula found on Juan Fernández and those found in the mainland should be recognised as different subspecies: island Carex firmula subsp. firmula and mainland C. firmula subsp. tenuis (Poepp. ex Kunth) R.Ridley & Jim.-Mejías. We discuss that the morphological differences may be a result of isolation in an oceanic island environment and have effect on seed dispersal. Taxonomic descriptions, images of diagnostic characters and a key are included.
References
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Jiménez-Mejías, P., Luceño, M., Wilson, K .L., Waterway, M.J. & Roalson, E.H. (2016) Clarification of the use of the terms perigynium and utricle in Carex L. (Cyperaceae). Systematic Botany 41 (3): 519–528. https://doi.org/10.1600/036364416X692488
Jiménez-Mejías, P. & Dorr, L.J. (2018) The Quinquín sedges: taxonomy of the Carex phleoides group (Cyperaceae). Annals of the Missouri Botanic Garden. 103: 591–603. https://doi.org/10.3417/2018125
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L’.Héritier de Brutelle, C.L. (1789) Sertum Anglicum: seu Plantae Rariores quae in Hortis juxta Londinum, Imprimis in Horto Regio Kewensi Excoluntur, ab Anno 1786 ad Annuum 1787 Observatae. Paris. https://doi.org/10.5962/bhl.title.11440
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Marticorena, C., Stuessy, T.F. & Baeza, C. (1998) Catalogue of the vascular flora of the Robinson Crusoe or Juan Fernández islands, Chile. Gayana Botánica (Chile) 55: 187–211.
Meyer, F.G. (1966) Chile-United States botanical expedition to Juan Fernández Islands. Antarctic Journal of the United States Sep-Oct: 238–242.
Ogle, D.H., Wheeler, P. & Dinno, A. (2018) FSA: Fisheries Stock Analysis. R package version 0.8.22, https://github.com/droglenc/FSA.
Persoon, C.H. (1807) Synopsis plantarum, seu Enchiridium botanicum, complectens enumerationem systematicam specierum hucusque cognitarum. Vol. 2. Parisiis Lutetiorum.
Pitman, N. (2004) Uncinia subsacculata. The IUCN Red List of Threatened Species 2004: e.T45155A10983787. https://doi.org/10.2305/IUCN.UK.2004.RLTS.T45155A10983787.en
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R Core Team. (2013) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. [http://www.R-project.org/]
Roalson, E., Columbus, J. &Friar, E. (2001) Phylogenetic Relationships in Cariceae (Cyperaceae) Based on ITS (nrDNA) and trnT-L-F (cpDNA) Region Sequences: Assessment of Subgeneric and Sectional Relationships in Carex with Emphasis on Section Acrocystis. Systematic Botany 26 (2): 318–341. [http://www.jstor.org/stable/2666710]
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Skottsberg, C. (1920–1956) The Natural History of Juan Fernández and Easter Island Vol 2: Botany. Uppsala. https://doi.org/10.5962/bhl.title.25662
Starr, J.R., Harris, S.A. & Simpson, D.A. (2004) Phylogeny of the unispicate taxa in Cyperaceae tribe Cariceae I: generic relationships and evolutionary scenarios. Systematic Botany 29 (3): 528–544. https://doi.org/10.1600/0363644041744455
Starr, J.R., Harris, S.A. & Simpson, D.A. (2008) Phylogeny of the unispicate taxa in Cyperaceae tribe Cariceae II: The limits of Uncinia in R.F.C. Naczi and B.A. Ford (editors), Sedges: Uses, Diversity, and Systematics of the Cyperaceae. Monographs in Systematic Botany from the Missouri Botanic Gardens 108: 243–267.
Starr, J.R. & Ford, B.A. (2009) Phylogeny and evolution in Cariceae (Cyperaceae): current knowledge and future directions. The Botanical Review 75 (1): 110–137. https://doi.org/10.1007/s12229-008-9020-x
Stearn, W.T. (1989) Botanical Latin. New Edition. David & Charles Inc.
Stuessy, T.F., Foland, K.A. Sutter, J.F. Sanders, R.W. & Silva, O.M. (1984) Botanical and geological significance of potassium-argon dates from the Juan Fernández Islands. Science 225: 49–51. https://doi.org/10.1126/science.225.4657.49
Stuessy, T.F., Swenson, U. Crawford, D.J. Anderson, G. & Mario Silva, O. (1997) Plant Conservation in the Juan Fernández Archipelago, Chile, Aliso 16 (2): 89–101. https://doi.org/10.5642/aliso.19971602.04
Thiers, B. (2016) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. [http://sweetgum.nybg.org/science/ih/] https://doi.org/10.1007/s12228-016-9423-7
Waterway, M.J. & Starr, J.R. (2007) Phylogenetic relationships in tribe Cariceae (Cyperaceae) based on nested analyses of four molecular data sets. Aliso 23: 165–192. https://doi.org/10.5642/aliso.20072301.13
Wheeler, G.A. (1994) The Uncinia (Cyperaceae) of Tierra del Fuego, the Falkland Islands and South Georgia. Anales Inst. Patagonia, Ci. Nat. 22: 21–31.
Wheeler, G.A. (1996) First report of Uncinia macrolepis and U. tenuis (Cyperaceae) in Colombia and new Colombian sites for U. paludosa. Rhodora 98 (893): 80–84.
Wheeler, G.A. (2007) Carex and Uncinia (Cyperaceae, Cariceae) from the Juan Fernández archipelago, Chile. Darwiniana 45 (1): 120–141.
Wheeler, G.A. & Goetghebeur, P. (1995) Four new species of Uncinia (Cyperaceae) from northern South America Aliso 14 (2): 141–146. https://doi.org/10.5642/aliso.19951402.06
Wheeler, G.A. & Goetghebeur, P. (1997) Uncinia (Cyperaceae) of Ecuador. Aliso 15 (1): 7–25. https://doi.org/10.5642/aliso.19961501.03
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Yen, A. & Olmstead, R. (2000) Molecular Systematics of Cyperaceae Tribe Cariceae Based on Two Chloroplast DNA Regions: NdhF and trnL Intron-Intergenic Spacer. Systematic Botany 25 (3): 479–494. https://doi.org/10.2307/2666691
Bernardello, G., Anderson, G., Stuessy, T. & Crawford, D. (2011) The angiosperm flora of the Archipelago Juan Fernández (Chile): Origin and dispersal. Canadian Journal of Botany 84:(8) 1266–1281. https://doi.org/10.1139/b06-092
Carlquist, S. (1966a) The biota of long-distance dispersal. II. Loss of dispersibility in Pacific Compositae. Evolution 20 (1): 30–48. https://doi.org/10.1111/j.1558-5646.1966.tb03341.x
Carlquist, S. (1966b) The biota of long-distance dispersal. III. Loss of dispersibility in the Hawaiian flora. Brittonia 18: 310–335. https://doi.org/10.2307/2805148
Carlquist, S. (1974) Island Biology. Columbia University Press, New York. https://doi.org/10.5962/bhl.title.63768
Cavanilles, A.J. (1799) Icones et Descriptiones Plantarum, vol. 5. Regia Typographia, Madrid.
Cody, M.L. & Overton, J. (1996) Short-term evolution of reduced dispersal in island plant populations Journal of Ecology, 53–61. https://doi.org/10.2307/2261699
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9 (8): 772. https://doi.org/10.1038/nmeth.2109
Flora del Conosur. (2021) Catálogo de las Plantas Vasculares. Instituto de Botánica Darwinion. Publishjed on the Internet: http://www.darwin.edu.ar/Proyectos/FloraArgentina/Familias.asp (Retrieved 15 November 2021)
Fresnillo, B. & Ehlers, B.K. (2008) Variation in dispersibility among mainland and island populations of three wind dispersed plant species. Plant Systematics and Evolution 270 (3–4): 243–255. https://doi.org/10.1007/s00606-007-0615-1
García-Moro, P., A. Otero, C. Benítez-Benítez, L. Costa, S. Martín-Bravo, R.F.C. Naczi, A.A. Reznicek, E.H. Roalson, J.R. Starr & P. Jiménez-Mejías (2022) Biogeography and Systematics of Carex subg. Uncinia: a unique radiation for the genus Carex in the Southern Hemisphere. Taxon.
Global Carex Group (2015) Making Carex monophyletic (Cyperaceae, tribe Cariceae): A new broader circumscription. Botanical Journal of the Linnean Society 179: 1–42. https://doi.org/10.1111/boj.12298
Govaerts, R. (compiler), Jiménez-Mejías, P., Koopman, J., Simpson, D.A., Goetghebeur, P., Wilson, K., Egorova, T. & Bruhl, J. (2021) World Checklist of Cyperaceae. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet: http://wcsp.science.kew.org/ (Retrieved 15 November 2021)
Hammel, B.E., Grayum, M.H., Herrera, C. & Zamora, N. (eds.) (2003) Manual de plantas de Costa Rica. Volumen II. - Missouri Botanical Garden Press.
Kudoh, H., Takayama, K. & Kachi, N. (2013) Loss of seed buoyancy in Hibiscus glaber on the oceanic Bonin Islands. Pacific Science 67 (4), 591–598. https://doi.org/10.2984/67.4.9
Idárraga-Piedrahita, A., Ortiz, R.D.C., Callejas Posada, R. & Merello, M. eds. (2011) Flora Antioquia: 2: 9–939. Universidad de Antioquia, Medellín.
Jiménez-Mejías, P, Hahn, M., Lueders, K., Starr, J.R., Brown, B.H., Chouinard, B.N., Chung, K., Escudero, M., Ford, B.A., Ford, K.A., Gebauer, S., Gehrke, B., Hoffmann, M.H., Jin, X., Jung, J., Kim, S., Luceño, M., Maguilla, E., Martín-Bravo, S., Míguez, M., Molina, A., Naczi, R.F.C., Pender, J.E., Reznicek, A.A., Villaverde, T., Waterway, M.J., Wilson, K.L., Yang, J., Zhang, S., Hipp, A.L., Roalson, E.H. The Global Carex Group. (2016a) Megaphylogenetic Specimen-Level Approaches to the Carex (Cyperaceae) Phylogeny Using ITS, ETS, and matK Sequences: Implications for Classification. Systematic Botany 41 (3): 500–519. https://doi.org/10.1600/036364416X692497
Jiménez-Mejías, P., Luceño, M., Wilson, K .L., Waterway, M.J. & Roalson, E.H. (2016) Clarification of the use of the terms perigynium and utricle in Carex L. (Cyperaceae). Systematic Botany 41 (3): 519–528. https://doi.org/10.1600/036364416X692488
Jiménez-Mejías, P. & Dorr, L.J. (2018) The Quinquín sedges: taxonomy of the Carex phleoides group (Cyperaceae). Annals of the Missouri Botanic Garden. 103: 591–603. https://doi.org/10.3417/2018125
Katoh, K., Rozewicki, J. & Yamada, K.D. (2017) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics. https://doi.org/10.1093/bib/bbx108
Kukethal, G. (1920) Cyperaceae novae. V. Repertorium novarum specierum regni vegetabilis 16: 430–435. https://doi.org/10.1002/fedr.19200162506
Kukkonen, I. (1967) Vegetative Anatomy of Uncinia (Cyperaceae). Annals of Botany 31 (3): 523–544. https://doi.org/10.1093/oxfordjournals.aob.a084161
Kunth, K.S. (1837) Enumeratio Plantarum, II. Cyperographia synoptica. Stuttgart and Tiibingen.
L’.Héritier de Brutelle, C.L. (1789) Sertum Anglicum: seu Plantae Rariores quae in Hortis juxta Londinum, Imprimis in Horto Regio Kewensi Excoluntur, ab Anno 1786 ad Annuum 1787 Observatae. Paris. https://doi.org/10.5962/bhl.title.11440
Linnaeus, C. (1753) Species Plantarum 2. Impensis Laurentii Salvii, Stockholm, pp. 972–979.
Marticorena, C., Stuessy, T.F. & Baeza, C. (1998) Catalogue of the vascular flora of the Robinson Crusoe or Juan Fernández islands, Chile. Gayana Botánica (Chile) 55: 187–211.
Meyer, F.G. (1966) Chile-United States botanical expedition to Juan Fernández Islands. Antarctic Journal of the United States Sep-Oct: 238–242.
Ogle, D.H., Wheeler, P. & Dinno, A. (2018) FSA: Fisheries Stock Analysis. R package version 0.8.22, https://github.com/droglenc/FSA.
Persoon, C.H. (1807) Synopsis plantarum, seu Enchiridium botanicum, complectens enumerationem systematicam specierum hucusque cognitarum. Vol. 2. Parisiis Lutetiorum.
Pitman, N. (2004) Uncinia subsacculata. The IUCN Red List of Threatened Species 2004: e.T45155A10983787. https://doi.org/10.2305/IUCN.UK.2004.RLTS.T45155A10983787.en
Presl, C. (1848) Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften 5 (5): 344. [Prague]
R Core Team. (2013) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. [http://www.R-project.org/]
Roalson, E., Columbus, J. &Friar, E. (2001) Phylogenetic Relationships in Cariceae (Cyperaceae) Based on ITS (nrDNA) and trnT-L-F (cpDNA) Region Sequences: Assessment of Subgeneric and Sectional Relationships in Carex with Emphasis on Section Acrocystis. Systematic Botany 26 (2): 318–341. [http://www.jstor.org/stable/2666710]
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 (12): 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
Ruiz, L.H. & Pavon, J.A. (1794) Florae Peruvianae, et Chilensis Prodromus, sive novorum generum plantrum peruvianum, et chilensium descriptiones et icones. Madrid, pp. 80, t. 15.
Skottsberg, C. (1920–1956) The Natural History of Juan Fernández and Easter Island Vol 2: Botany. Uppsala. https://doi.org/10.5962/bhl.title.25662
Starr, J.R., Harris, S.A. & Simpson, D.A. (2004) Phylogeny of the unispicate taxa in Cyperaceae tribe Cariceae I: generic relationships and evolutionary scenarios. Systematic Botany 29 (3): 528–544. https://doi.org/10.1600/0363644041744455
Starr, J.R., Harris, S.A. & Simpson, D.A. (2008) Phylogeny of the unispicate taxa in Cyperaceae tribe Cariceae II: The limits of Uncinia in R.F.C. Naczi and B.A. Ford (editors), Sedges: Uses, Diversity, and Systematics of the Cyperaceae. Monographs in Systematic Botany from the Missouri Botanic Gardens 108: 243–267.
Starr, J.R. & Ford, B.A. (2009) Phylogeny and evolution in Cariceae (Cyperaceae): current knowledge and future directions. The Botanical Review 75 (1): 110–137. https://doi.org/10.1007/s12229-008-9020-x
Stearn, W.T. (1989) Botanical Latin. New Edition. David & Charles Inc.
Stuessy, T.F., Foland, K.A. Sutter, J.F. Sanders, R.W. & Silva, O.M. (1984) Botanical and geological significance of potassium-argon dates from the Juan Fernández Islands. Science 225: 49–51. https://doi.org/10.1126/science.225.4657.49
Stuessy, T.F., Swenson, U. Crawford, D.J. Anderson, G. & Mario Silva, O. (1997) Plant Conservation in the Juan Fernández Archipelago, Chile, Aliso 16 (2): 89–101. https://doi.org/10.5642/aliso.19971602.04
Thiers, B. (2016) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. [http://sweetgum.nybg.org/science/ih/] https://doi.org/10.1007/s12228-016-9423-7
Waterway, M.J. & Starr, J.R. (2007) Phylogenetic relationships in tribe Cariceae (Cyperaceae) based on nested analyses of four molecular data sets. Aliso 23: 165–192. https://doi.org/10.5642/aliso.20072301.13
Wheeler, G.A. (1994) The Uncinia (Cyperaceae) of Tierra del Fuego, the Falkland Islands and South Georgia. Anales Inst. Patagonia, Ci. Nat. 22: 21–31.
Wheeler, G.A. (1996) First report of Uncinia macrolepis and U. tenuis (Cyperaceae) in Colombia and new Colombian sites for U. paludosa. Rhodora 98 (893): 80–84.
Wheeler, G.A. (2007) Carex and Uncinia (Cyperaceae, Cariceae) from the Juan Fernández archipelago, Chile. Darwiniana 45 (1): 120–141.
Wheeler, G.A. & Goetghebeur, P. (1995) Four new species of Uncinia (Cyperaceae) from northern South America Aliso 14 (2): 141–146. https://doi.org/10.5642/aliso.19951402.06
Wheeler, G.A. & Goetghebeur, P. (1997) Uncinia (Cyperaceae) of Ecuador. Aliso 15 (1): 7–25. https://doi.org/10.5642/aliso.19961501.03
Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. ISBN 978-3-319-24277-4. [http://ggplot2.org] https://doi.org/10.1007/978-3-319-24277-4
Yen, A. & Olmstead, R. (2000) Molecular Systematics of Cyperaceae Tribe Cariceae Based on Two Chloroplast DNA Regions: NdhF and trnL Intron-Intergenic Spacer. Systematic Botany 25 (3): 479–494. https://doi.org/10.2307/2666691