Three new species of ferns (Woodsiaceae and Polypodiaceae) from Mexico JOHN T, M I C K E L AND J. DANIEL T E J E R O - D I E Z

Mickel, J. T. (The New York Botanical Garden, Bronx, NY, 10458-5126, U.S.A.; e-mail: [email protected]) & J. D. Tejero-Dfez, (Universidad Nacional Autrnoma de Mrxico, Escuela Nacional de Estudios Profesionales Iztacala, Carrera de Biologfa, Apartado Postal 314, Tlalnepantla 54090, Mrxico, Mexico; e-mail: [email protected]). Three new species of ferns (Woodsiaceae and Polypodiaceae) from Mexico. Brittonia 56:115-120. 2004.~Three new species of ferns are described from Mexico: Athyrium tripinnatum, Cystopteris millefolia, and Polypodium praetermissum.

Key words: Mexico, ferns, Athyrium, Cystopteris, Polypodium.

Preparation of manuscripts on the pteridophytes of the Republic of Mexico (Mickel & Smith, submitted 2002) and on the pteridoflora of the state of M r x i c o (TejeroDfez, 1998), reveal three new species o f ferns, which are here described.

Athyrium tripinnatum Mickel & Tejero, sp. nov. (Fig. 1D-F) TYPE: M E X I C O . Mrxico: Mun. Tejupilco, desviaci6n a E1 Reparo, K m 38 a partir de la desviaci6n (carretera Tejupilco-Bejucos) a Nanchititla, 1500 m, 30 Sep 1988, Tejero-Dfez 2919 (HOLOTYPE: IZTA; ISOTYPE: NY). Ab A. palmensi et A. skinneri laminis tripinnatopinnatifidis distincta.

Rhizomes short-creeping, 3 m m diam.; rhizome scales light brown, narrowly deltate, with long-acuminate apices, the margins entire, to 4 • 0.5 m m ; fronds approximate ( 2 - 4 m m distant), to 60 cm long; stipes ca. 1/3 frond length, stramineous, adaxially grooved, with sparse linear scales similar to those of rhizomes and appressed septate hairs 0.2-0.5 m m long (mostly on abaxial surface); blades tripinnate-pinnatirid, broadly deltate, to 30 cm wide, with 15-18 pairs of free pinnae, the blade apex shortly attenuate; rachises subflexuous,

stramineous, slightly alate, with scattered hairs similar to those o f stipes mainly on abaxial surface; pinnae alternate, narrowly triangular, ascending, the basal ones to 19 • 10 cm, texture herbaceous, abaxial surface with appressed hairs 0.1-0.2 m m long, adaxial surface glabrous; veins pinnate, ending in the teeth; sori one per ultimate s e g m e n t , 1 - 1 . 5 m m long, e l o n g a t e , Jshaped, or round; indusia 0.2-0.4 m m wide, whitish, coarsely toothed. Distribution.--Terrestrial in oak woods, locally abundant on slopes of arroyos; 1500 m. K n o w n only f r o m the type collection. Athyrium tripinnatum is similar to A. palmense (H. Christ) Lellinger in having slender rhizomes, thin deltate blades, and an indusium deeply lacerated into 2 - 4 teeth, but differs in having blades that are tripinnate-pinnatifid rather than bipinnatepinnatifid as in A. palmense or pinnate-pinnatifid as in A. skinneri. With increasing dissection, there is increasing frond size: A. skinneri (Baker) C. Chr. to 44 c m long with 5 - 7 pinna pairs, A. palmense to 53 cm long with 12-15 pinna pairs, and A. tripinnatum to 60 cm long with 15-18 pinna pairs.

Cystopteris millefolia Mickel & Tejero, sp. nov. (Fig. I A - C ) TYPE: M E X I C O . Mrxico: Mun. Ocuilan, K m 18, Ocuilan-Cuernavaca, 2130 m, 9

Brittonia, 56(2), 2004, pp. 115-120. 9 2004, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

ISSUED: 7 April 2004

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hi

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i

I f

ram. :

I I _L

A /

5cm.

FIG. 1. A - C . Cystopteris millefolia (Hinton 13499, ARIZ). A. Habit. B. Pinna. C. Segments. D-F. Athyrium tripinnatum (Tejero D{ez 2929, NY). D. Pinna. E. Segment. F. Sorus.

Nov 1985, Tejero-Dfez 2234 (HOLOTYPE: R h i z o m e s s h o r t - c r e e p i n g , ca. 2 m m NY; ]SOTYPE: IZTA). diam., not protruding beyond fronds; rhizome scales brown, m e m b r a n o u s , lustrous, A C. fragili laminis tripinnato-pinnatifidis vel quad- lanceolate, entire, 1.5-3 • 0.8-1.5 m m , ripinnatis, segmentis linearibus differt, mixed with fine hairs; fronds 1 4 - 5 0 c m

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l o n g , c l u m p e d ; s t i p e s 1/3-~ f r o n d l e n g t h , s t r a m i n e o u s , r e d d i s h b r o w n at base, w i t h s c a l e s at b a s e to 3 m m long, m i x e d w i t h hairs as on r h i z o m e s , g l a b r o u s d i s t a l l y ; b l a d e s t r i p i n n a t e - p i n n a t i f i d to q u a d r i p i n nate, l a n c e o l a t e , 1 2 - 3 0 • 4 . 5 - 1 4 cm, w i t h proximal 2-3 pinna pairs slightly reduced; rachises glabrous or with scattered minute g l a n d u l a r hairs in r a c h i s g r o o v e s a d a x i a l l y ; p i n n a e 1 2 - 1 6 p a i r s , 2 - 8 c m long, p e r p e n d i c u l a r to r a c h i s e s , c h a r t a c e o u s , g l a b r o u s ; s e g m e n t s linear, 1 m m w i d e ; veins e n d i n g in s m a l l e m a r g i n a t i o n s ; indusia h o o d l i k e , g l a b r o u s , entire. D i s t r i b u t i o n . - - W e t m o s s y r o c k s in f o r e s t s t r e a m s , m o i s t b a n k s in p i n e a n d p i n e - f i r forests; 2 2 0 0 - 3 5 0 0 m. C e n t r a l M e x i c o . Additional specimens examined. MEXICO. M6xico: Side of Popocatdpetl, ca. 4100 m, 16 Jul 1947, Barkley et al. 2346 (LL); NW of Toluca, Mexico Hwy 15, Km 105, 8500 ft (ca. 2600 m), 16 Sep 1962, Barr 62-795 (ARIZ): Mum Villa de Allende, San Jos6 de Allende, 10 Oct 1985, Bartholomew 2886 (GH, MEXU); between Km 76 and 77 on Amecameca-Popoeat6petl rd., 3250 m, 2 Aug 1958, Beaman 2061 (TEX); Mun. Temascaltepec, E1 Polvorfn, Km 13 Toluca a Temascaltepec, 9 May 1988, Flores 817 (MEXU); Mun. Ocuilan, Km 14 entre Santa M6nica Ocuilan y Cuernavaca, 99~ '44"W, l 8~ Herndndez Salazar 15 (IZTA); Mun. Ocuilan, Cerro Zempoala en el Parque Nacional Lagunas de Zempoala, 22 Jul 2000, Rojas 5411 (MEXU); Mun. Villa Victoria, Laguna del Carmen en Parque Nacional Bosencheve, 19~ 100~ Tejero-Dfez 2937 (IZTA). Michoacfin: Rt. 15, ca. 0.7 km N of Km 14@ ca. 7 km W of M6xico-Michoacfin state line, ca. 2930 m, 14 Jul 1966, Cruden 1068 (GH, MICH); Mun. Angangueo, Ejido Cerro Prieto en Barranca Honda, 15 Mar 1987, Diaz Barriga 3597 (IEB, MEXU); Distrito Zit~cuaro, Zit~cuaro-La Difunta, 3000 m, 27 Nov 1938, Hinton 13499 (ARIZ, GH, LL, PH, TEX): Distrito Uruapan, Tancitaro. 3150 m, 18 Nov 1940, Hinton 15711 (TEX): Mun. Quer~ndaro, 3 kmal sur de San Jos6 de Ia Cumbre. Rzedowski 48315 (MEXU). Nlorelos: Mun. Cnernavaca, Barranca Tepeite, Dec 1947, Lyonnet [471200019] (MEXU); Mun. Cuernavaca. Barrancas rumbo a Santa Maria de Ocuilan, Pdrez-Garc{a 766 (UAMIZ). C y s t o p t e r i s f r a g i l i s (L.) Bernh. s.1. is o n e o f the m o s t w i d e s p r e a d p t e r i d o p h y t e s p e cies in the w o r l d . It is k n o w n to h y b r i d i z e w i t h o t h e r s p e c i e s in N o r t h A m e r i c a ( H a u tier et al., 1990, 1993; H a u f l e r & W i n d h a m , 1991). N e a r l y all M e x i c a n and C e n t r a l A m e r i c a n m a t e r i a l is t r e a t e d b y us and o t h ers (e.g., Stolze, 1981; M i c k e l & B e i t e l , 1988; M o r a n , 1995; R i b a et al., 1996) as C.

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f r a g i l i s , l a r g e l y b e c a u s e it has not b e e n s t u d i e d as c l o s e l y as s p e c i m e n s f r o m N o r t h America. Cystopteris membranifolia Mickel ( f r o m V e r a c r u z , O a x a c a , a n d G u a t e m a l a ) is d i s t i n g u i s h e d by its v e r y thin b l a d e s , l o n g - c r e e p i n g r h i z o m e s , and t u b e r c u l a t e e c h i n a t e spores. W i t h i n C. f r a g i l i s s.1. in M e x i c o , t h e r e is c o n s i d e r a b l e m o r p h o l o g i cal v a r i a t i o n ( e s p e c i a l l y in d e g r e e o f d i s section), s p o r e v a r i a t i o n ( b o t h r u g o s e a n d echinate), and ploidal variation, judging f r o m s p o r e sizes ( d i p l o i d 2 6 - 3 9 Ixm, tetrap l o i d 3 7 - 4 5 txm, h e x a p l o i d 4 6 - 5 3 ~ m ; M o r a n , pers. c o m m . ) . M o s t s p e c i m e n s h a v e v e i n s e n d i n g in s e g m e n t e m a r g i n a t i o n s , b u t this c h a r a c t e r s e e m s variable. M u c h , b u t c e r t a i n l y n o t all, M e x i c a n m a t e r i a l r e f e r r e d to C. f r a g i l i s m a y b e C. r e e v e s i a n a L e l l i n g e r w h i c h is c o m m o n in the s o u t h w e s t e r n U n i t e d States, b u t until d e t a i l e d studies a r e m a d e , w e p r e f e r to u s e the n a m e C. f r a g i l i s . T h e r e m a y y e t b e s e v e r a l t a x a to b e s e g r e g a t e d f r o m C. f r a g i l i s in M e x i c o , b u t C. millefolia, w i t h its finely d i s s e c t e d f r o n d s , is the m o s t distinct. P o l y p o d i u m p r a e t e r m i s s u m M i c k e l & Tej e r o , sp. n o v . (Fig. 2 A - G )

TYPE: M E X I C O . N a y a r i t : M u n . d e Tepic, a 17 k m d e Jestis M a r f a r u m b o a la M e s a del Mayab, 22~ 104~ 7 Sep 1991, R a m f r e z 1078 (HOLOTYPE: M E X U ; ISOTYPES: I Z T A , N Y ) . A P. plesiosoro squamis rhizomatis comosis, patentioribus distincta. R h i z o m e s c r e e p i n g , ca. 5 m m d i a m . ; rhiz o m e s c a l e s c o v e r i n g 8 5 - 1 0 0 % o f rhiz o m e s , o r a n g e - t a n , c o n c o l o r o u s to s o m e w h a t p a l e r at m a r g i n s , s o m e t i m e s w i t h darker point of attachment, ovate-lanceolate, c o m o s e , 2 . 0 - 3 . 5 x 0 . 8 - 2 . 1 ram, marg i n s d e n t i c u l a t e o r subentire, m o s t l y asc e n d i n g or s o m e w h a t s p r e a d i n g ; f r o n d s 2 1 45 • 6 - 1 2 ( - 1 8 ) c m , distant; stipes 1/3 f r o n d length, s t r a m i n e o u s to c a s t a n e o u s , a d a x i a l l y grooved, glabrous; blades pinnatisect, deltate to o b l o n g , a p e x p i n n a t i f i d or w i t h a s u b c o n f o r m t e r m i n a l p i n n a , hairs o f l a m i n a r t i s s u e 0 . 1 - 0 . 3 m m l o n g w i t h 2 - 3 cells; p i n nae 8 - 2 6 pairs, l i n e a r - d e l t a t e , a d n a t e a n d s o m e w h a t s u r c u r r e n t to s l i g h t l y n a r r o w e d at their b a s e s , the p r o x i m a l 1 - 2 p a i r s o f t e n

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FIG. 2. Polypodium praetermissum. A. Habit. B. Rhizome scale. C. Pinna detail. D. Pinna detail, distal pinna. E. Habit. F. Pinna detail, proximal pinna. G. Rhizome scale. (A-D, Palmer 374, NY; E-G, Reeder & Reeder 2473, US.)

discrete and not adnate, attenuate, the larger ones (at or near the base) 5 . 0 - 6 . 0 cm X 0.8-1.2 cm, margins inconspicuously crenulate; rachises a n d costae stramineous to

greenish, with scattered hairs on both surfaces but more abaxially, hairs catenate, 0.3-0.7 m m long with 5 - 1 0 cells; veins partially and irregularly anastomosing with

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more anastomoses in proximal half of b l a d e s a n d i n d i v i d u a l p i n n a e ; sori o b l o n g or o c c a s i o n a l l y r o u n d , m i d w a y b e t w e e n costae a n d m a r g i n s in s i n g l e rows, 1-1.5 • 1 . 5 - 2 . 0 ram.; s p o r a n g i a glabrous. D i s t r i b u t i o n . - - O n rocks or in r o c k y soil, rarely epiphytic, w e t n o r t h - f a c i n g cliffs a n d r o c k faces on l a v a flows a n d steep arroyos in oak, p i n e - o a k , a n d p i n e forests; 1 1 0 0 2 1 5 0 m. M e x i c o . Additional specimens examined. MEXICO. Durango: About 21 mi E of Santa Lucfa on the rd. from Villa Uni6n to Durango, 28 Sep 1953, Reeder & Reeder 2473 (MEXU, US). Jalisco: Mun. Zapopan, La Primavera, 24 Aug 1988, Cola 17004 (XAL); Barranca hr. Atequizall~in,lower slopes of Volc~inNevado, Correll 14362 (US); Mun. Tapalpa, 1-2 mi E of Tapalpa, 30 Oct 1960, McVaugh 20530 (MEXU, MICH, MO); Muff. Etzatlfin, cerca de Etzatlfin, 2 Oct 1903, Rose & Painter 7585 (GH). Nayarit: Mun. E1 Nayar, Arroyo de la Taberna NW of Mesa del Nayar, 1325 m, 13 Aug 1980, Breedlove 45550 (CAS, MEXU); Mun. Ahuacatl~in, Volcfin Ceboruco, 3.5 mi NW of Ahuacatlfin, 13 Aug 1959, Feddema 462 (GH, IEB, NY); Mun. Santa Marfa del Oro, NE of Santa Marfa del Oro, 1820 Aug 1959, Feddema 776 (MEXU, MICH); Mun. Tepic, 17 km de Jes6s Marfa, rumbo a la Mesa del Mayab, 22~ 104~ 1 Aug 1990, Flores 2155 (MEXU); Mun. Compostela, 9 mi N of Compostela, 28 Aug 1957, McVaugh 16497 (MEXU); Mun, Ahuacatl~in, arroyo 5 mi SE of Ahuacatl~n on rd. to Barranca del Oro, 27 Aug 1957, McVaugh 16353 (MEXU); 6.5 mi NW of Ahuacatl~in,new lava flow of Volcfin Ceboruco, 13 Aug 1963, Mickel 1370, 1371, 1375 (all NY): Mun. Acaponeta, 5 mi NE of La Mesa Nayar, 29 Jul 1970, Norris & Taranto 14275 (ENCB); Mun. Ixtl~in del Rio, 11 mi N of Ixtlfin del Rfo, 13 Aug 1959, Straw & Foreman 1820 (GH); Mun. Ahuacatl~in, 10 km al S de Ahacatlfin, 21~ 104~ 19 Oct 1986, Tdllez 9777 (MEXU), 9863 (MEXU); Mun. Tepic, 4 km a San Bias a partir del entronque Tepic-Mazatl~in, 6 Aug 1987, Tdllez 10739 (MEXU). Sinaloa: Mun. Badiraguato, 5 mi NE of La Cienega along rd. to Santa Rita, 2230 m, 5 Oct 1970, Breedlove 18573 (NY), Mun. Concordia, along small logging rd. hr. Loberas Microwave Station, 1570 m, 4 Aug 1980, 44963 (CAS); Mun. Tamazula, La Bajada, 500 m, Gonzdlez Ortega 4291 (MEXU); Cerro Quemado, NE of Panuco, ca. 1500 m, 30 Aug 1935, Pennell 20137 (PH). Sonora: Sierra de Alamos, upper Cation de Bolsa, 1650 m, 20 Aug 1994, Fishbein 1934 (ARIZ), 1937 (ARIZ); Cerro Seguarivo, E of San Bernardo, 15001700 m, 7-8 Aug 1935, Pennell 19657 (PH); Trigo Moreno, Sierra E1 Chuchupate, ca. 15 km (by air) SE of Yecora, 1420-1480 m, 17 Aug 1998, Van Devender 98-1025 (NY). P o l y p o d i u m p r a e t e r m i s s u m has blade surfaces with short hairs o n the costae a n d rachises adaxially, scattered o n l a m i n a e a n d axes abaxially. H o w e v e r , the S o n o r a n a n d

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S i n a l o a n s p e c i m e n s are e s s e n t i a l l y g l a b r o u s a n d have f e w e r p i n n a e ( 8 - 1 0 vs. 1 3 - 2 0 in D u r a n g o , 2 5 - 2 8 in N a y a r i t a n d Jalisco). F u r t h e r m o r e , blades are n a r r o w l y o b l o n g i n Jalisco a n d N a y a r i t (Fig. 2 A - D ) vs. b r o a d l y deltate in D u r a n g o , Sinaloa, a n d S o n o r a (Fig. 2 E - G ) . This v a r i a t i o n is p r o b a b l y clinal. P o l y p o d i u m p r a e t e r m i s s u m is close to P. p l e s i o s o r u m K u n z e i n its g e n e r a l architecture, with m a n y b r o a d l y adnate p i n n a e (at least in Jalisco a n d N a y a r i t ) a n d short l a m inar hairs, b u t differs in h a v i n g s p r e a d i n g to a s c e n d i n g , c o m o s e r h i z o m e scales a n d less r e g u l a r v e i n a n a s t o m o s e s , P o l y p o d i u m p r a e t e r m i s s u m has a p i n n a t e (vs. p i n n a t i sect) b l a d e base a n d g e n e r a l l y alternate (vs. opposite) p i n n a e . In P. p r a e t e r m i s s u m the r h i z o m e scales are clearly i n d i v i d u a l , s o m e w h a t spreading, a n d n a r r o w l y o v a t e - d e l t a t e to l a n c e o l a t e with a c l u m p of hairs at the base o f the scales, at the p o i n t o f attachm e n t ; in P. p l e s i o s o r u m the scales are s t r o n g l y a p p r e s s e d to the r h i z o m e s or to each other ( m a k i n g it o f t e n difficult to distinguish individual scales) and broadly ovate to o b l o n g (at least those o f the b o d y o f the r h i z o m e ) to b r o a d l y l a n c e o l a t e at the r h i z o m e apices. T h e n e w species also r e s e m b l e s P o l y p o d i u m s u b p e t i o l a t u m Hook., w h i c h can b e d i s t i n g u i s h e d b y its a b u n d a n t l a m i n a r hairs, toothed p i n n a m a r g i n s , a n d free veins. T h e m o r e n o r t h e r n s p e c i m e n s o f Po l yp o d i u m p r a e t e r m i s s u m r e s e m b l e P. f r a t e r n u m Schltdl. & C h a m . i n h a v i n g g l a b r o u s (or s u b g l a b r o u s ) l a m i n a e , i r r e g u l a r l y a n a s t o m o s i n g veins, f e w e r p i n n a pairs, attenuate p i n n a apices, a n d u n t o o t h e d p i n n a m a r g i n s , b u t P. p r a e t e r m i s s u m differs in h a v i n g more broadly adnate pinnae, thinner blade texture, shorter a n d c o m o s e r h i z o m e scales, s o m e short hairs o n the costae a n d l a m i n a e , a n d m o r e n o r t h w e s t e r n distribution. The more northern specimens of Polyp o d i u m p r a e t e r m i s s u m are s i m i l a r to P. dulce Poir. in Lam. in b l a d e a n d p i n n a shape a n d c o m o s e r h i z o m e scales. T h e former, though, is d i s t i n c t in h a v i n g a n a s t o m o s i n g (vs. free) veins.

Acknowledgments T h e d r a w i n g s were p r e p a r e d b y H a r u t o F u k u d a . We t h a n k Dr. A l a n S m i t h for as-

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sistance with the Latin diagnoses and Leone Brown for helping prepare the manuscript. Literature Cited H a u f l e r , C. H., R. C. M o r a n & M. D. W i n d h a m . 1993. Cystopteris. Pages 263-270. In: N. R. Morin, c o n v e n i n g editor. Flora of North A m e r i c a North of Mexico. Vol. 2. Pteridophytes and G y m n o s p e r m s . Oxford University Press. New York. - & M . D. W i n d h a m . 1991. N e w species of North A m e r i c a n Cystopteris and Polypodium, with c o m m e n t s on their reticulate relationships. Amer, Fern J. 81: 7-23. , & T. A. R a n k e r . 1990. Biosystematic analysis o f the Cystopteris tennesseensis (Dryopteridaceae) complex. Ann. Missouri Bot. Gard. 77: 314-329. M i c k e l , J. T. & J. M. Beitel. 1988. Pteridophyte flora

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of Oaxaca. Mexico. Mem. N e w York Bot. Gard. 46: 1-568. - & A. R. S m i t h . Submitted. Pteridophytes of Mexico. Mere. New York Bot. Gard. M o r a n , R. C. 1995. Cystopteris. Page 228. In: R. C. Moran & R. Riba, volume editors. Psilotaceae a Salviniaceae. h~: G. Davidse, M. Sousa S. & S. Knapp. general editors. Flora Mesoamericana. Vol. 1. Universidad Nacional A u t r n o m a de Mrxico, Ciudad Universitaria. Riba, R., L. P a c h e c o , A. Vald~s & Y. S a n d o v a l . 1996. Pteridoflora del estado de Morelos. Lista de familias, g6neros y especies. Acta Bot. Mex. 37: 45-65. Stolze, R. G. 1981. Ferns and fern allies o f Guatemala. Part II. Polypodiaceae. Fieldiana, Bot. n.s. 6: 1522. Tejero-Diez, D, 1998. Pteridoflora del occidente del estado de Mrxico, Mexico. Masters thesis, Facultad de Ciencias, Universidad Nacional A u t 6 n o m a de M r x i c o . Mdxico D.E