A Taxonomic and Biogeographic Reappraisal of the Genus Dicksonia (Dicksoniaceae) in the Neotropics Author Noben, Sarah Institute for Systematic and Evolutionary Botany, University of Zurich, Zollikerstr. 107, 8008 Zurich, Switzerland sarah.noben@uzh.ch Author Kessler, Michael Institute for Systematic and Evolutionary Botany, University of Zurich, Zollikerstr. 107, 8008 Zurich, Switzerland Author Weigand, Anna Institute for Systematic and Evolutionary Botany, University of Zurich, Zollikerstr. 107, 8008 Zurich, Switzerland Author Tejedor, Adrian School for Field Studies, 100 Cummings Center, Suite 534 - G, Beverly, MA 01915, USA. Author Rodr´, Wilson D. Sinchi-Amazonic Institute of Scientific Research, Calle 20 Número 5 - 44, Bogotá, Cundinamarca, Colombia Author Gallego, Luis Fernando Giraldo Universidad CES, Calle 10 A Número 22 - 04, Medell´ın, Antioquia, Colombia & Herbario Universidad de Antioquia (HUA), Calle 67 Número 53 - 108, Medell´ın, Antioquia, Colombia Author Lehnert, Marcus Nees-Institut für Biodiversität der Pflanzen, Rheinische Friedrich-Wilhelms Universität Bonn, Meckenheimer Allee 170, D- 53115 Bonn, Germany & Systematische Botanik und Mykologie, Ludwig-Maximilians-Universität München, Menzinger Strasse 67, 80638 München, Germany & Institute for Systematic and Evolutionary Botany, University of Zurich, Zollikerstr. 107, 8008 Zurich, Switzerland text Systematic Botany 2018 Basel, Switzerland 2018-11-28 43 4 839 857 https://doi.org/10.1600/036364418x697634 journal article 309952 10.1600/036364418X697634 9c17e11b-1c44-4df6-a7b3-98e2d5bc098b 1548-2324 15483304 3. DICKSONIA NAVARRENSIS Christ , Bull. Herb. Boissier II, 6:188. 1906. TYPE : COSTA RICA . San José : Valley of the Rio Navarro , [ca. 09°57 ' 54"N , 83°58 ' 05"W , 1400–1800 m ] K. Wercklé 1905 ( holotype P-00642552 ! [Herb. Christ]; isotype K-000894778 !). Dicksonia ghiesbreghtii Maxon , Contr, U.S. Natl. Herb. 17:155. 1913. TYPE : MEXICO . Chiapas : Without locality, A.B. Ghiesbreght 353 ( holotype US-00066374 !, isotypes BM-000050846 !, GH-00008277 !, NY-149027 !/- 149028 ! [fragments of US ], P-00642547 [fragment of US]). Trunks to 8 m tall, 19.5 cm in diam including persistent old petiole bases, with additional varying cover of adventitious roots, these usually rusty red when young; with a loose skirt of old fronds; adventitious buds occurring regularly on the petiole bases instead of the lowest pinna pairs (ML pers. obs.; Calderón-Sáenz 2000 ). Fronds to 400 cm long, monomorphic, erect to patent, larger ones arching. Petioles mostly 10–25 cm (in immature plants to 70 cm ) long, covered with woolly, pale, yellowish to orange hairs with an outer layer of longer reddish to brown hairs, all hairs thin-walled, all catenate ( Panama to Ecuador ) or only the longest hairs turgid ( Mexico to Costa Rica ); lowest pinnae may be replaced by adventitious buds ( Panama to Ecuador ). Laminae to 330 X 153 cm , tripinnate-pinnatifid, coriaceous, obovate to oblanceolate, basally tapering. Frond axes (rachises, costae, costules) smooth, few hairs on upper part of rachis, not persistent, costae and costules covered with ciliform hairs, adaxially reddish, antrorsely curved, to 1.5 mm long, abaxially pale brown to white, appressed to matted, partially catenate, to 2 mm long. Pinnae to 76.5 X 28.5 cm , sessile, oblong-lanceolate with attenuate tips, ca. (18–)22–26 pairs per frond, lower pairs reduced to ca. 1/3–1/4 the length of longest medial pinnae, lowest ones pinnate-pinnatifid with strong costa. Pinnules to 12.0 X 2.4 cm , oblong to triangular-lanceolate, sessile to subsessile, basally truncate to weakly auriculate, apically attenuate. Segments to 13.5 X 4.8 mm , sessile, weakly falcate, lobed to weakly dissected, margins flat to weakly revolute, proximal segments larger than medial ones. Veins glabrous or midvein abaxially with few whitish hairs to 1 mm long. Sori 0.9–1.4 mm diam, oblong when closed, circular when open, on the acroscopic and basiscopic side of the segment, each at the end of a lateral vein on a separate lobes, distance receptacle to the midvein 1.5–2.2 mm . Indusia bivalved, outer one brown with entire cartilaginous margin, inner one light brown with subentire to erose margins, beset with few obovate, hyaline cells. Paraphyses catenate, pale, fragile, slightly longer than sporangia. Spores tetrahedral-globose with prolonged, depressed lobes, 51 X 36 μm , exospore smooth, perispore bacillar-granular. Figures 1E, I, J , 5D . FIG. 7. Important characters of Neotropical Dicksonia . A. Dicksonia lehnertiana . A 1 . Petiole. A 2 . Fertile pinnule, abaxially. ( Lehnert 3019, BONN). B. Dicksonia stuebelii . B 1 . Petiole. B 2 . Fertile pinnule, abaxially ( Sanchez 11111, UC). C. Dicksonia sellowiana . C 1 . Petiole, hairs only persisting near base ( Hatschbach 43455, US). C 2 . Sterile pinnule, adaxially. C 3 . Immature fertile segments abaxially, with costae still covered in woolly undercoat. C 4 . Fertile segments abaxially, sori fully developed. D. Dicksonia antarctica . D 1 . Crozier, showing easily abraded woolly undercoat. D 2 . Fertile segments abaxially, with glabrescent costae and costules. (C 2 –C 4, D: from living plant at Botanic Garden Bonn). Etymology —The name refers to the R´ıo Navarro near San José, Costa Rica , the type locality. Distribution and Habitat —Distributed from Mesoamerica to the northern Andes, in evergreen montane rainforests at 940–2450(–2800) m ( Fig. 6 ). Additional Specimens Examined — Mexico . — CHIAPAS : Pueblo Nuevo Solistahuacá , 17°11 ' 24"N, 92°54 ' 15"W, 2000 m , 16 Dec 1971 , D. E . Breedlove 23214 ( MO ) .— OAXACA : Cerro Zempoaltepetl , 17°09 ' 09"N, 96°00 ' 29"W, 2000–2438 m , 19 Sep 1970 , J . T . Mickel and S . W . Leonard 4680 ( UC ) ; Teotitlán , 17°01 ' 58"N, 96°31 ' 12"W, 2133–2255 m , 16 Oct 1969 , J . T . Mickel and R . L . Hellwig 4126 ( UC ) . El Salvador . — CHALATENANGO : E slope of Los Esesmiles , 14°21’N , 89°09’W , 2500 m , 24 Mar 1942 , J . M . Tucker 1127 ( MO ) .— SANTA ANA : Parque Nacional Montecristo , 14°23 ' 18"N , 89°23 ' 02"W , 2250 m , 08 May 1979 , R. Seiler 1143 ( UC ) . Guatemala . — HUEHUETENANGO : Todos Santos Cuchumatan , 15°31’N , 91°39’W , 2600 m , 26 Feb 2009 , M . M . J . Christenhusz , M . S . Vorontsova , J . Jiménez Barrios , K . Watson and C . Espada Mateos 5358 ( BM , MO ) . Honduras . — EL PARAÍSO : Yuscarán , 13°55 ' 50"N, 86°53 ' 43"W, 2000 m , 22–25 Mar 1975 , C . H . Nelson and E . Vargas N . 2547 ( MO ) .— FRANCISCO MORAZÁN : Distrito Central , 14°12 ' 30"N, 87°05 ' 39"W, 1800–2100 m , 25 March 1951 , C . V . Morton 7486 ( MO ) .— OCOTEPEQUE : La Labor , 14°28’N , 89°04’W , 1870 m , 12 Jun 1985 , E . M . Mart´ ınez S . and O . Téllez – V . 13001 ( GH n.v., MO ) .— SANTA BÁRBARA : Cuesta de Piedra Caliza , 10 km oeste de Lago Yojoa , 14°55’N , 88°07’W , 2200–2350 m , 28–30 April 1973 , A. F. Clewell and D.L. Hazlett 3930 ( MO ) . Nicaragua . — JINOTEGA : Municipio de Bocay. Reserva Natural Kilambé , 13°42’N , 85°42’W , 1200–1700 m , 11–19 Jan 2001 , R . M . Rueda et al. 15668 ( MO ) .— MATAGALPA : Behind La Selva Negra Hotel , slopes of Cerro Picacho , near the border with Dept. Jinotega , 13°00’N , 85°55’W , 1200–1540 m , 23 May 1985 , G. Davidse , A. Grijalva P. and M. Sousa S. 30367 ( MO ) .— NUEVA SEGOVIA : 13°58 ' 50"N, 86°11 ' 30"W, 1255–1660 m , Jul 2006 , L .D. Paguaga and N . Toval 41 ( HULE n. v., MO ) . Costa Rica . — CARTAGO : Carpintera , 1850 m , 10 Apr 1908 , Brade 119 ( P ) ; Valle de la Estrella , 09°46 ' 23"N, 83°57 ' 39"W, 1500 m , 26 January 1980 , H . Churchill 3271–2 ( STU , VT ) ; Canton de Paraiso , 09°44 ' 53"N , 84°07’W , 1600 m , 10 January 1997 , S. Salas , A. Rodrigues , and A. Stot 187 ( MO ) .— HEREDIA : Volcan Barva , 10°08.5 ' N , 84°07’W , 2450–2800 m , 28 Apr 1986 , M.H. Grayum 7471 ( MO ) ; Braulio Carillo National Park , 10°20’N , 84°10’W , 1215 m , 13 Nov 1986 , E . Hennipman , M . C . Roos , G . P . Verduyn , P . Velhoen 6872 ( MO ) ; P . N . Tapanti-Cerro de la Muerte , Reserva Tapanti , sendero “Arbol Caido”, 09°45.137’N , 83°46.985’W , 1300–1400 m , 17 Nov 2012 , M . Lehnert 2650 ( BONN ) . LIMÓN : Cantón de Limón , 09°46 ' 15"N, 83°20 ' 15"W, 1060–1330 m , 10 Aug 1995 , A . Rojas , G . Herrera , M . H . Grayum 2248 ( MO ) .— PUNTARENAS : Cantón Coto Brus , 08°57 ' 03"N , 82°45 ' 03"W , 1900–2100 m , 20 April 1999 , A. Rojas 5074 ( MO ) .— SAN JOSÉ : Cerro Tablazo , 09°50 ' 07"N, 84°01 ' 58"W, 1900 m , 20 Jul 1909 , A . C . Brade 203 ( STU ) . Panama . — PANAMÁ : Valle de Antón , road to Altos de Maria , 08°38.708’N , 80°05.654’W , 940 m , 03 Jan 2016 , M . Lehnert 3493 ( BONN , Z ) . Colombia . — ANTIOQUIA : Andes , Vereda Cascajeros , cabecera de la Quebrada La Chaparrala , 05°36 ' 15"N, 75°54 ' 10"W, 1950 m , 16 May 2000 , F . Giraldo and S . Mej´ ıa 2055 ( HUA , JAUM ) ; Anor´ı, Reserva La Forzosa , Sector Muros de Piedra , Qda , El Chaquiral , 06°58 ' 53"N, 75°08 ' 32"W, 1660 m , 28 Sep 2003 , W . Rodriguez et al. 4041 ( HUA ) ; Vereda El Roble , Finca La Forzosa , 6.9733306 , -75.1329444 , 1700 m , 12 Apr 2000 , F. Giraldo and S. Mej´ ıa 2022 ( HUA , JAUM ), Vereda Villa Fátima , Alto la Aguada , Quebrada la Soledad , 6.9860111 , -75.1190528 , 1440 m , 30 Jan 2015 , F. Giraldo 2899 ( HUA ) and M. Lehnert 2994 ( BONN , HUA , Z ) ; Corregimiento de Providencia , 07°18 ' 59"N, 75°03 ' 23"W, 1428 m , 08 Oct 2013 , P . Trujillo and R . Sabala 7020 ( HUA ) ; Amalfi , v´ıa principal hacia Medell´ ın, sector la Cascada , 06°42 ' 19"N, 75°09 ' 52"W, 1650 m , 16 Apr 2000 , F . Giraldo and S . Mej´ ıa 2035 ( HUA , JAUM ) ; carretera que conduce a la vereda El Guayabito , 06°52 ' 55.3"N , 75°05 ' 28.8"W , 1650 m , F. Giraldo 2045 ( HUA , JAUM ) ; Medell´ın, Paraje Boquerón , por la carretera al mar, 06°14 ' 27"N, 75°30 ' 49"W, 2200 m , 14 Oct 1982 , J . Hernandez et al. 519 ( HUA ) ; Yarumal , Corregimiento de Cede ~ no, Vereda La Palmita , orillas del Rio San Julián , 7.0121250N , - 75.3805861W , 1680 m , 17 Mar 2000 , F . Giraldo and S . Mej´ ıa 1983 ( HUA , JAUM ) .— CAUCA : Parque Nacional Munchique , road W towards Pacific slope, 02°42.268’N , 76°53.145’W , 2060 m , 12 Feb 2015 , M . Lehnert et al. 3122 ( BONN , HUA , Z ) ; Reserva Natural Tambito , 02°30 ' 25"N, 75°00 ' 08"W, 1590 m , 23 Aug 2000 , O . L . Casa ~ nas 495 ( COL ) .— CHOCÓ : Darién , 06°23’N , 76°31’W , 1250–1350 m , D. B . Lellinger 1980 ( COL ) .— HUILA : Palestina , Vereda La Guajira , Reserva Natural Comunitaria La Rivera , zona de amortiguación PNN cueva de Los Guacharos , 1.6563889N , - 76.1869444W , 1970 m , 23 Aug 2005 , A . León 700 ( FMB ) .— NARINO ~ : Ricaurte , 1.1333333 , -77.9666667 , 1750 m , 28 Nov 1981 , A . W . Gentry 35129 ( COL ) ; La Planada , 02°38.457’N , 77°58.942’W , 1700 m , 14 Feb 2015 , M . Lehnert et al . 3141 (BONN, HUA, Z).— RISARALDA : Cerro Montezuma , Reserva Montezuma , PN Tatama , 05°15.296’N , 76°06.559’W , 2080 m , 07 Feb 2015 , M. Lehnert et al. 3077 ( BONN , HUA , Z ) ; Mistrato , Mpio.de Mistrato , Correg.de La Leguada , 5.4075000N , - 76.0394444W , 1500 m , 28 Apr 1992 , J. Betancur 3243 ( COL ). — VALLE DE CAUCA : Dagua , cuenca del R´ıo Dagua , Km 23 de la carretera Cali-Dagua , 2050 m , 25 Sep 1991 , E . Calderon S. 103 ( COL ) ; La Cumbre , Corregimiento de Bitaco , Reserva Agua Bonita , 02°42.268’N , 76°35 ' 11"W , 1700–1900 m , 10–15 Dec 1998 , W. Vargas 5236 ( HUA , CUVC ). Ecuador . — CARCHI : Chical , 01°02 ' 37"N , 78°15 ' 00"W , 1300–1500 m , 25 Sep 1979 , A. Gentry and G. Schupp 26567 ( MO ) ; Cerro Golondrias , 00°53’N , 78°10’W , 1740–1780 m , 24 Apr–01 May 1993 , B . Boyle and L . Dalmau 1757 ( MO ) ; Reserva Ind´ıgena Awá , 00°53’N , 78°25’W , 1800 m , 17–27 Aug 1992 , G . Tipaz , M . Tirado , C . Aulestia , N . Gale and P . Ortiz 1842 ( MO ) . Notes — Christ (1906) , when describing Dicksonia lobulata and D. navarrensis , specifically refers to the specimens in his herbarium as types. According to TL-2 (http://www.sil.si. edu), Christ’ s herbarium remained largely in his hometown of Basel (BAS) except for the pteridophytes, which were transferred to Paris (P). For this reason, we regard the specimens at P with the label information “Herb. Christ, Bale ˆ” as holotypes , of which there are one for each name. Similarly, the type of D. ghiesbreghtii at US is regarded here as an originally designated holotype , since Maxon (1913) refers to the specimen at US as “type” and to material of the same gathering deposited in other herbaria as “duplicates.” A putative isotype of D. navarrensis at BM is just a small sterile frond ca. 40 cm long with 10 pinna pairs (Herbarium Christensen 679, annotated “Herb. H. Christ, Bale ˆ”). Dicksonia navarrensis corresponds with “ Dicksonia gigantea” sensu Lellinger (1989) , but the latter name appears to be more correctly placed under D. karsteniana , which see for further discussion. Dicksonia navarrensis can be separated from D. karsteniana and D. sellowiana by having larger segments (8–14 X 3.0– 4.5 mm ) with small sori ( 0.9–1.4 mm in diam) vs. smaller segments (5–11 X 2–4 mm ) with larger sori ( 1.5–2 mm in diam) in D. karsteniana , and smaller segments (3.5–7.0 X 2.3–3.2 mm ) with small sori ( 0.9–1.2 mm in diam) in D. sellowiana . Additionally, the receptacles are more distant from the segment midveins (on average 1.5–2.2 mm in D. navarrenssis vs. 0.7–1.5 mm in D. karsteniana and 0.6–1.2 mm in D. sellowiana ). Proximal segments are larger than medial ones in D. navarrensis and D. karsteniana vs. proximal segments more or less the same size as the medial ones in D. sellowiana . Plants of D. navarrensis have petioles covered with matted, woolly hairs (vs. petioles with straight hairs and soft matted undercoat in D. karsteniana ). Dicksonia karsteniana never has entirely woolly hairs, and plants that resemble D. navarrensis most in laminar dissection and sorus size come from the lower elevations of the eastern Andean slopes and have only the straight dark petiole hairs remaining in mature fronds, woolly hairs being sparse or soon shed. Axes are predominantly smooth to slightly rough in D. navarrensis , generally smooth in D. sellowiana , but variable in D. karsteniana , so we do not include it as a distinguishing character for these taxa as was done in previous studies ( Lellinger 1989 ). Lellinger (1989) reported soral paraphyses without a glandular terminal cell in D. navarrensis (as D. gigantea ) and soral paraphyses often with a dark red, nonswollen, glandular terminal cell in D. karsteniana , as a further distinguishing character. This character, however, is ambiguous because paraphyses are often broken off in older fronds and especially in dried material. We cannot confirm such a correlation and assume that all species treated here have paraphyses with clavate terminal cells and that the red color is dependent on external factors. The most outstanding distinguishing character of Dicksonia navarrensis is the presence of adventitious buds at the petiole bases ( Calderón-Sáenz 2000 ). These sprout into small plants once the rest of the leaf has rotted. Consequently, these small plants are usually not preserved in herbarium specimens, and the buds, once dry, are hard to recognize as such among the woolly hairs on the petiole. We can confirm the presence of these buds for the populations from central Panama throughout the Colombian Chocó region , but we have no such information for Costa Rica and places north, neither from our own observations nor specimen labels or the literature (e.g. Véliz and Vargas 2006 ). None of the type specimens of the names united here under Dicksonia navarrensis contains petiole material, which could show the presence of adventitious buds and the characteristic dark catenate hairs.