Research Article |
Corresponding author: Rita Ema Morero ( ritamorero@gmail.com ) Corresponding author: Gloria E. Barboza ( gbarboza@imbiv.unc.edu.ar ) Academic editor: Blanca León
© 2016 Rita Ema Morero, David S. Barrington, Monique A. McHenry, João Paulo Condack, Gloria E. Barboza.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Morero RE, Barrington DS, McHenry MA, Condack JPS, Barboza GE (2016) Typifications and synonymy in Polystichum (Dryopteridaceae) from Chile and Argentina. PhytoKeys 65: 91-105. https://doi.org/10.3897/phytokeys.65.8620
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Polystichum Roth is one of the largest and most taxonomically challenging fern genera. South American species have a rich and complex nomenclatural history; many of the early names are inadequately typified. Based on extensive examination of original type material, we designate eleven lectotypes (including Aspidium mohrioides, A. montevidense f. imbricata, A. montevidense f. squamulosa, A. plicatum, A. pycnolepis, Dicksonia andina, Polystichum elegans, P. mohrioides f. latifolia, P. multifidum var. autranii, P. platyphyllum var. kurtziana, and Polypodium polystichoides), and one neotype (P. brongniartianum) for Polystichum taxa. Furthermore, three new synonyms are proposed.
Polystichum Roth es uno de los géneros de helechos más grandes y de mayor complejidad taxonómica. Las especies sudamericanas tienen una rica y compleja historia nomenclatural, con muchos de los nombres inadecuadamente tipificados. En base a un minucioso examen del material tipo original, designamos lectotipos para once taxones de Polystichum (que incluyen Aspidium mohrioides, A. montevidense f. imbricata, A. montevidense f. squamulosa, A. plicatum, A. pycnolepis, Dicksonia andina, Polystichum elegans, P. mohrioides f. latifolia, P. multifidum var. autranii, P. platyphyllum var. kurtziana y Polypodium polystichoides) y un neotipo (P. brongniartianum). Además, se proponen 3 nuevos sinónimos.
Ferns, taxonomy, typification, nomenclature, South America
Helechos, taxonomía, tipificación, nomenclatura, Sudamérica
Polystichum Roth (Dryopteridaceae) is a worldwide, taxonomically complex genus of ± 360–400 species (
Taxa in the genus Polystichum can be recognized morphologically by stems usually ascending or erect, short internodes, and a dictyostelic stele. The monomorphic leaves bear several kinds of scales (hairs are rare); the ultimate segments are mostly asymmetric at base, often acroscopically auriculate, and ordinarily spinulose or at least dentate-mucronate at the margins. The sori are indusiate or not, when present the indusia are rounded-peltate.
South America, with ~40 species (43% of species in the New World), is an important world center of diversity (
Recent work has revealed that the polystichums of Chile and Argentina pertain to two different monophyletic lineages, an exindusiate tropical Andean clade (
In the course of work on revisions of Polystichum from Argentina and Chile we became aware that critical nomenclatural work with original materials was needed for a number of the taxa. In this manuscript we designate lectotypes for eleven taxa, one neotype, and propose three new synonyms for Central Andean and Southern Andean Polystichum which inhabits Argentina and Chile. We report the novelties for the two regions separately, alphabetically by basionym species name.
Review of types was based on examination of specimens in herbaria (
Information about botanical publications, authors, dates, collectors and their herbarium and types, were taken from
1. Aspidium mohrioides d´Urv., Flore des Iles Malouines: 26. 1825. Type: [Islas Malvinas] “I. Soledad”, n.d., D. d’Urville 92. Lectotype (designated here): P! (P00636426 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p00636426]). Syntypes remaining: [Islas Malvinas] “Iles Malouines, D. d’Urville s.n. – P! (P00636427 [https://science.mnhn.fr/institution/mnhn/collection/p/item/p00636427]). Iles Malouines, [year] 1815, M. Lesson s.n. – P! (P0036428 [https://science.mnhn.fr/institution/mnhn/collection/p/item/p00636428]).
= Polystichum mohrioides (d´Urv.) C.Presl, Tent. Pterid.: 83. 1836.
Three specimens are stored at P from Islas Malvinas, two collected by J. S. C. Dumont d´Urville (P00636426 and P00636427) and the third by R. P. Lesson (P00636428). The ferns were collected during a circumnavigation in the corvette “Le Coquille” (1822-1825) under the command of Captain L. Duperrey, in which Dumont d’Urville was a second officer and Lesson surgeon and naturalist (
2. Aspidium plicatum Poepp. ex Kunze, Linnaea 9: 94. 1834.Type: [CHILE. Bio-Bio:] “Chile austr. in rupibus mont. Pico de Pilque in Cordillera de Antuco”, Dec 1828. E. F. Poeppig (Diar. 745). Lectotype (designated here): W! [W-0003927]; isolectotypes:
= Polystichum plicatum (Poepp. ex Kunze) Hicken ex Hosseus, Trab. Inst. Bot. Farmacol. (Buenos Aires) 33: 9. 1915.
When
3. Dicksonia andina Phil., Anales Univ. Chile 94: 359. 1896. Type: [CHILE. Valdivia:] “Cuesta de Lipela, Cord. de Ranco”, Jan 1887, O. Philippi s.n. Lectotype [designated here (second step lectotypification after Rodríguez Ríos 1987: 51)]:
= Polystichum multifidum (Mett.) T. Moore, Index Fil. (T. Moore): 84. 1857. Basyonym: Aspidium multifidum Mett., Fil. Lechl. 1: 20. 1856.
Rudolph A. Philippi misapplied the name Dicksonia (Fam. Dicksoniaceae), a genus of tree ferns, to specimens of Polystichum. We found three sheets at
4. Polystichum brongniartianum J.Rémy, in Fl. Chil. [Gay] 6: 518. 1854. Type: [CHILE. Concepción]: “Chile Austral, Concepción” C. Gay s.n., not localized. Neotype (designated here): CHILE. Llanquihue: Peulla. 13 Jun 1953, A. Pfister s.n.
= Polystichum subintegerrimum (Hook. & Arn.) R.Rodr., Gayana. Bot. 44: 48. 1987. Basyonym: Aspidium subintegerrimum Hook. & Arn., Bot. Beechey Voy.: 52. 1832.
The type specimen of P. brongniartianum (C. Gay s. n.) cited by
5. Polystichum elegans J.Rémy, in Fl. Chil. [Gay] 6: 514. 1854. Type: [CHILE. Prov. Colchagua: “Cordillera de Talcaregué, pie le Volcán de Cordier”, Feb 1831, C. Gay 29. Lectotype [designated here (second step lectotypification after Rodríguez Ríos 1987: 49)]: P! (P00636434 [http://plants.jstor.org/stable/10.5555/al.ap.specimen.p00636434?searchUri=filter%3Dname%26so%3Dps_group_by_genus_species%2Basc%26Query%3Dpolystichum%2Belegans]); isolectotypes: P! (P00636432 [https://science.mnhn.fr/institution/mnhn/collection/p/item/p00636432], P00636435 [https://science.mnhn.fr/institution/mnhn/collection/p/item/p00636435]).
= Polystichum plicatum (Poepp. ex Kunze) Hicken ex Hosseus, Trab. Inst. Bot. Farmacol. (Buenos Aires) 33: 9. 1915. Basyonym: Aspidium plicatum Poepp. ex Kunze, Linnaea 9: 94. 1834.
There are three sheets of Gay at P, two sheets are housed as Gay 29 (P00636434! and P00636432!) with data of the collection place, while the third sheet (P00636435!) lacks collector number and only “Chili” is indicated as the collection location. Rodríguez Ríos (1987) wrote “Se halla en las altas cordilleras de Talcaregue, provincia de Colchagua, en las orillas de los arroyos, cerca del volcán en donde es algo rara (P!)”. The Gay 29’s collection has a label with the same information cited by Rodríguez Ríos. As a second step lectotypification is necessary, we select P00636434 as the second step lectotype because it includes the diagnostic character for this taxon and it is the most complete accession (three fronds and the petiole scales are intact).
6.Polystichum mohrioides f. latifolia Hicken, Anales Soc. Ci. Argent. 62: 171. 1906. Type: [ARGENTINA. Mendoza. Dpto. San Carlos:] “Cordillera del Portillo de La Llareta entre El Paso del Portillo (4300 m.s.m.) y la Laguna del Diamante (3324 m.s.m.). La Resguardia del Portillo, in petros, umbrosis passim. Leg. Arturo Lemos”, Mar 1900, F. Kurtz 11173. Lectotype (designated here):
= Polystichum plicatum (Poepp. ex Kunze) Hicken ex Hosseus, Trab. Inst. Bot. Farmacol. (Buenos Aires) 33: 9. 1915. Basyonym: Aspidium plicatum Poepp. ex Kunze, Linnaea 9: 94. 1834.
We examined the three syntypes of P. mohrioides f. latifolia. All three are of good quality; two are preserved at
7.Polystichum multifidum var. autranii Hicken, Anales Soc. Ci. Argent. 62: 172. 1906. Type: [ARGENTINA] “Chubut: Laguna Blanca, J. Koslowsky n° 240”. Lectotype (designated here): SI000112!; isolectotypes: SI000113!
= Polystichum multifidum (Mett.) T. Moore, Index Fil. (T. Moore): 84. 1857. Basyonym: Aspidium multifidum Mett., Fil. Lechl. 1: 20. 1856.
Although there are two sheets of this variety at
8.Polystichum multifidum var. dusenii C.Chr., Ark. Bot. 10 (2): 19. 1910. syn. nov. Type: [CHILE, Aysén:] “Patagonia Occ. in valle fluminis Aysén”, 15 Jan 1897, P. Dusén 489 Holotype: S! (S05-10906 [http://plants.jstor.org/stable/10.5555/al.ap.specimen.s05-10906])
= Polystichum chilense (H.Christ) Diels, Nat. Pflanzenfam. [Engler & Prantl] 1: 192. 1899. Basyonym: Aspidium aculeatum var. chilense (H.Christ) Ber. Schweiz. Bot. Ges. 3: 39. 1893.
Christensen (1910) recognized P. multifidum var. dusenii based on lamina division (twice pinnate-pinnatifid lamina), and shape and margin of pinnules (pinnules obovate with slightly dentate margins). Looser (1968) suggested that this set of plants should be considered a variety of P. chilense; later, Rodríguez Ríos (1987) agreed and added another diagnostic character (coriaceous pinnules), to support this position and formalized the recognition of the variety: P. chilense var. dusenii (C.Chr.) R.Rodr. We observed frequent variation in the lamina division among P. chilense populations, even within the same population. This variability is often associated with the size and age of the plant (Barrington 2012). Consequently, we synonymize var. dusenii under P. chilense.
9. Polystichum pearcei Phil., Linnaea 33: 305. 1865. syn. nov. Type. [CHILE] Valdivia, Cordillera de Ranco, s.f., R. Pearce s.n. Holotype:
= Polystichum multifidum (Mett.) T. Moore, Index Fil. (T. Moore): 84. 1857. Basyonym: Aspidium multifidum Mett., Fil. Lechl. 1: 20. 1856.
The main diagnostic characters for P. pearcei are the 3-pinnate frond and the rachis slightly scaly (Philippi 1865). Rodríguez Ríos (1987) considered the same diagnostic characters but subordinated this species as a variety under P. multifidum. Analyzing herbarium specimens and field observations of P. multifidum, we found high variability in the lamina division and density of scales on the rachis. In addition, when P. multifidum grows in warmer and/or drier conditions, the leaves are smaller, less divided, and not as scaly (Morero, pers. obs.). Since the type of Polystichum pearcei falls within the variation of P. multifidum, we propose it as a synonym. Molecular studies in progress support this proposal (Barrington pers. comm.).
10.Aspidium montevidense f. imbricata Hieron., Bot. Jahrb. Syst. 22 (3): 366. 1897. Type: [ARGENTINA, Tucumán:] “Siambón, Sierra de Tucumán”, 8 May 1872, P. G. Lorentz 158. Lectotype (designated here):
= Polystichum montevidense (Spreng.) Rosenst., Hedwigia: 46: 111, 1906. Basyonym: Polypodium montevidense Spreng., Syst. Veg. (ed. 16) [Sprengel] 4 (1): 59. 1827.
This name was based on two syntypes both from Tucumán hills (Argentina): Lorentz 158 and Lorentz & Hieronymus 801. Hieronymus described this taxon with the specimens deposited at B, but a careful search of B does not yield them. Four duplicates of Lorentz 158 exist in herbaria: two housed at
11.Aspidium montevidense f. squamulosa Hieron., Bot. Jahrb. Syst. 22 (3): 366. 1897. Type: [ARGENTINA, Córdoba:] “Las Ramadas cerca de San Miguel, Sierra Achala de Córdoba”, 14 Mar 1876, G. Hieronymus 479. Lectotype (designated here):
= Polystichum pycnolepis (Kunze ex Klotzsch) Hieron., Bot. Jahrb. Syst. 34 (4): 452. 1904. Basyonym: Aspidium pycnolepis Kunze ex Klotzsch, Linnaea 20: 365. 1847.
We located two of the three syntypes cited by
12. Aspidium pycnolepis Kunze ex Klotzsch, Linnaea 20: 365. 1847. Type: [VENEZUELA] “Columbia, Col. Tovar”. 1856. J. W. K. Moritz 296 b. Lectotype (designated here): B! (B200148046-b [http://ww2.bgbm.org/herbarium/specimen.cfm?SpecimenPK=83712&idThumb=279937&SpecimenSequenz=1&loan=0]); isolectotypes: B! (B200148045-b [http://ww2.bgbm.org/herbarium /specimen.cfm?SpecimenPK=166846&idThumb=279936&SpecimenSequenz=1&loan=0]); K! (K000512992 [http://apps.kew.org/herbcat/detailsQuery.do?imageId=231125&pageCode=1&presentPage=1&queryId=2&sessionId=E0DC715CEF51CFBB8FB85F938E0D6780&barcode=K000512992]. Syntypes remaining: [VENEZUELA] “Columbia”. 1856. J. W. K. Moritz 296 – B! (B200148177 [http://ww2.bgbm.org/herbarium/specimen.cfm?SpecimenPK=83714&idThumb=279972&SpecimenSequenz=1&loan=0], B200148176 [http://ww2.bgbm.org/herbarium/specimen.cfm?SpecimenPK=83713&idThumb=279971&SpecimenSequenz=1&loan=0]); K! (K000512989 [http://apps.kew.org/herbcat/detailsQuery.do?imageId=231122&pageCode=1&presentPage=1&queryId=1&sessionId=E0DC715CEF51CFBB8FB85F938E0D6780&barcode=K000512989]), M! (not available on line),
= Polystichum pycnolepis (Kunze ex Klotzsch) Hieron., Bot. Jahrb. Syst. 34 (4): 452. 1904.
The type material of Aspidium pycnolepis is a mixed collection. In order to stabilize the application of this name in the sense that it has been used by most other botanists, we choose Moritz 296 b (B200148046!) since most closely represents the original description. The remaining syntypes, after the designation of the lectotype of A. pycnolepis, pertain to at least three species, only one of which is P. pycnolepis. We have seen the following: “Columbia, Páramo de la Culata”, Moritz 296 (B! [det. P. orbiculatum and P. gelidum, see annotation labels]; K! [det. P. gelidum]; M! [det. P. pycnolepis];
13. Polypodium polystichoides Klotzsch, Linnaea 20: 383, 1847, syn. nov. Type: [COLOMBIA] “Columbien 1846 (Col. II)”, s.d. H. Karsten 5. Lectotype (designated here): B! (B200148238 [http://herbarium.bgbm.org/object/B200148238]); isolectotypes: B! (B200148237 [http://herbarium.bgbm.org/object/B200148237], B200148239 [http://herbarium.bgbm.org/object/B200148239]);
= Polystichum platyphyllum (Willd.) C.Presl, Tent. Pterid. 84. 1836. Basyonym: Aspidium platyphyllum, Sp. Pl. ed. 4, 5 (1): 255. 1810.
There are three sheets of Karsten at B (one more, not found, may be housed at
14. Polystichum platyphyllum var. kurtziana Hicken, Anales Soc. Ci. Argent. 63: 169. 1907. Type: [ARGENTINA] “Prov. Córdoba: Departam. San Alberto. Sierra Grande: Cuesta del Tránsito: Región del Tabaquillo. An feuchten Orten in grossen Gruppen; zerstreut”, 6-13 Jan 1895, F. Kurtz 8352. Lectotype (designated here):
= Polystichum montevidense (Spreng.) Rosenst., Hedwigia: 46: 111. 1906. Basyonym: Polypodium montevidense Spreng., Syst. Veg. (ed. 16) [Sprengel] 4(1): 59. 1827.
A single collection was cited in the protologue of this varietal name. Of the four duplicates of the type collection, two are housed at
We are indebted to the curators and staff of A,