NEW BASIC CHROMOSOME NUMBERS for GENERA of NEOTROPICAL Fernsl
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Amer. J. Bot. 67(5): 733-738. 1980. NEW BASIC CHROMOSOME NUMBERS FOR GENERA OF NEOTROPICAL FERNSl FLORENCE S. WAGNER Department of Botany, The University of Michigan, Ann Arbor 48109 ABSTRACT Chromosome numbers for eight genera of Costa Rican ferns are reported here for the first time. These are: Neurocallis, n = 58: Marginariopsis, n = 35: Dicranoglossum, n = 36: G/y photaenium, n = 37; Cochlidium, n = 33; Ormolomu, n = 42; Peltapteris, n = 40; and Loxso mopsis, n = 46. Chromosome observations involving several other genera are included: So lanopteris, n = 37; Xiphopteris, n = 32; Hemidictyum, n = 31; P/euroderris, 2n = 80; Dictyoxiphium, 2n = 80: and Polystichum, n = 41. These numbers are compared to those of genera of presumed taxonomic affinity. OUR KNOWLEDGE of the cytotaxonomy of MATERIALS AND METHODS-Ail ofthe chro Neotropical ferns, which essentially began mosome counts to be reported here were ob with the publication of Walker's survey of Ja tained from plants growing in Costa Rica. Fer maican pteridophytes (1966, 1973a), has been tile parts of fronds collected in the field were extended in other areas ofthe New World trop placed, at the end of each day's field trip, in ics, in Mexico by Mickel, Wagner and Chen small bottles containing a saturated solution (1966) and Smith and Mickel (1977), and in ofparadichlorobenzene, and refrigerated at ca. Brazil by Tryon, Bautista and Araujo (1975). 4 C. After 24 hr or less, excess water was Individual monographic studies and taxonomic blotted from the fertile leaf fragments, which papers have further augmented the list of were then put in Newcomer's Solution, left at known chromosome numbers. A summary of room temperature overnight, and placed in a the present state of our information can be freezer in the morning. Chromosome studies found in the study of "Evolutionary Patterns were carried out at the University of Michigan and Processes in Ferns" (Lovis, 1977) which using ACH (50% acetocarmine:50% Hoyer's contains a list of the basic chromosome num Solution) as a squashing and mounting medi bers of fern genera. According to the classi um. fication by Lovis, there remain 80 genera (in All collections were made by W. H. and F. cluding a few designated as subgenera) for S. Wagner and L. D. Gomez P. (with the ex which cytological observations have not been ception of Peltapteris foeniculacea) and are published. Approximately 25 of these are New represented by duplicate vouchers (in the field World genera. This report presents chromo number series of W. H. Wagner) both in the some numbers for 11 of these previously un University of Michigan Herbarium (MICH) counted genera in Lovis' list, including for the and in the Herbario Nacional of the Museo sake of completeness several in which reports Nacional, San Jose, Costa Rica (CR). have been only recently published. I also in clude discussion of certain unusual new num OBSERV ATIONS-The observations are ar bers that differ from those previously reported ranged according to the sequence presented by in some genera. Brief comments are added, Lovis (1977, p. 261 ff.). where appropriate, on distinctive taxonomic characters. NEUROCALLIS Fee Neurocallis praestantissima (Bory) Fee 1 Received for publication 27 July 1979; revision ac cepted 16 November 1979. (Fig. 1, 13)-C. R. Heredia Provo Slopes of I acknowledge the constant and generous help afforded Volcan Barba, Zurqui, steep rainforest along me by Luis Diego Gomez P., Director of the Museo Na stream, 13 March 1979, Wagner 79058. The cional de Costa Rica, and by W. H. Wagner, Jr. Further studies of species reported here and other Central Amer monotypic genus Neurocallis is a rare and lo ican ferns in collaboration with these two are being carried cal fern of mountain rainforests in the West out as part of an NSF Latin American Cooperative Science Indies, Costa Rica and Colombia. Neurocallis Grant INT78-19909. I am also indebted to David B. Lel praestantissima has simple-reticulate venation linger of the U.S. National Herbarium for identifications of several species in this report, to Alan R. Smith for (i.e., without included veinlets), and dimor suggested revisions, and to Jean Hazen for laboratory as phic, nearly acrostichoid fertile fronds-these sistance. characters distinguishing it from most of the 733 734 AMERICAN JOURNAL OF BOTANY [Vol. 67 '. P, • -:l. + ~~ . ~ ~/" , '.. • ", ~I .... ";' '1• ~ "" 4 • ~ . e-'" .,. li- ,. 1 , q ( .II-# .. • ::~ ... ~ I ... ' . ~ f It ~ ... f. ~ \ •, ••• I ."'.. .> • 2 • 3 .~. ..... (-""-' 11 c { "'\ ... I- ~ ~/-J. 6- • e ~ ...,.- i~ ~ " I, {, ... 1 + .. r/L -- ~ ~.. 4 5 6 0 ., .~ "'" .. () If» .:) - " • -" • .. • w.. .' ".- '" 1 ... •.. .., .. - .r 7 , .If 9 Q ~ f II t) b '"~\ D .; Jt, iJ 10 , 11 12 Fig. 1-12. Photographs of chromosomes. Fig. 13-24. Corresponding interpretations of chromosome figures. 1,13. Neurocallis praestantissima (Bory) Fee, n = 58, x270. 2,14. Marginariopsis wiesbaurii (Sodiro) C. Chr., n = 35, x610. 3,15. Dicranoglossum panamense (C. Chr.) L. D. Gomez, n = 36, x670. 4,16. Glyphotaenium percrassum (Bak.) Copel., n = 37, x660. 5,17. Xiphopteris limula Christ, 2n = 64, x830. 6,18. Xiphopteris limula Christ, 2 cells, n = 32, x660. 7,19. Cochlidium rostra tum (Hook.) Maxon ex C. Chr., n = 33, x660. 8,20. Ormoloma standleyi May-June, 1980] WAGNER-CHROMOSOME NUMBERS OF NEOTROPICAL FERNS 735 ~ A~ 1.1•• , ~ A .. ~~ -. .-., " , .... .. ,~ ...... '" .. ,. K ~+ "I\ , r. • ~~ ~.. JI""..0r~.I~~..0" • • e ... (),. • 13 14 • 15 e~ I\~ ~ '\(., Ir.~Q 1""\ \'Y ) ",', \ ... ... .., ~,~ ~I-J. 6'" 1i;I~, ~~ ,\ t -11 t t.. ~ ~ ,,< \ \-~, '~h-' ~ 6~¥i: ''\.1 JJ »,» I ... t ~ 16 17 18 ¥ 0 0 4 ,)It'tI/ f .",~ .. -. • J <> .... .4-••..' • ~ .... .,.'- to • '\ 0'" It. ~ 0 .,. ", ........ , ~.,.tt • .." .. .. Df .. Jr '" #//1 ...., • ....-tt'.t6 ..Qr ~. 8 .... 0)0" U "'" ,'"A ., - e 1.~ l ~ .. ~ f; ",r 19 20 "'-Pc 21 9 #4 *- + ~ ~A.J. GO~ ~ ~~ b .... ~. J 'V~ 0 ...., ~\ ~O .. ,. ~ - #I :''St-;. :~.I ~ \ ~ ... ~ '.. ,. ••+~. JI , , .... 22 • 23 '" Jf: 24 Maxon, n = 42, x610. 9,21. Hemidictyum marginatum (L.) Presl, n = 31, x 720. 10,22. Polystichum dubium (Karst.) Diels, 11 = 41, x830. 11,23. Peltapteris foeniculacea (Hk. & Grev.) L. D. Gomez, n = 40, x670. 12,24. Loxsomopsis costaricensis Christ, n = 46, x800. 736 AMERICAN JOURNAL OF BOTANY [Vol. 67 species of its probable closest relative, Pteris. and Mickel, 1977, have found P. angusta in Meiotic material from Costa Rica has 58 pairs Mexico, however, to have 34 pairs at meiosis.) of chromosomes, a number consistent with our The chromosome number of Dieranoglossum present theories of its affinities with Pteris, in is in fact uncommon among Neotropical which x = 29. The Neotropical Pteris gran polypodies. difolia L. has fronds similar to those of Neu roeallis, although they are not dimorphic. The GLYPHOTAENIUM J. Smith venation pattern of P. grandifolia is unusual in having free veins that become reticulate in Glyphotaenlum percrassum (Bak.) Cope a band near the margin. A chromosome count, land (Fig. 4, 16)-C. R. Heredia Provo Vol n = 58, from Costa Rican material of this din Barba, above Porrosati, 13 November species (Limon Provo Moin , 16 November 1977, Wagner 77517. Glyphotaenium (includ 1977, Wagner 77577) is the same as that re ing Enterosora Baker) is a genus of small ported from Mexican and Jamaican material. epiphytic grammitid ferms of Central Amer ica and northern South America, which char MARGINARIOPSIS Christensen acteristically have thick, spongy leaf tissue and impressed or immersed sori. Glypho Marginariopsis wiesbaurii (Sodiro) C. Chr. taenium percrassum has a chromosome num (Fig. 2, 14)-C. R. Alajuela Provo Road N from ber of 37 pairs. Copeland (1947) has suggested San Ramon to Balsa, 14 March 1977, Wagner that Glyphotaenium may be derived from 77063. Marginariopsis, a rare, monotypic Grammitis trifurcata (L.) Cope I. That species genus of small epiphytes occurring in Costa also has 37 pairs (Walker, 1966). Rica and Ecuador, has strongly dimorphic fronds, the fertile ones linear and coenosoral, COCHLIDIUM Kaulfuss the sterile ones lanceolate. The chromosome number of Marginariopsis wiesbaurii is 35 Coehlidium rostratum (Hooker) Maxon ex pairs at meiosis. Relatively few Neotropical C. Chr. (Fig. 7, 19)-C. R. Cartago Prov. San members of the Polypodiaceae have this num Cristobal, 15 November 1978, Wagner ber, the more usual chromosome complement 78574. Coehlidium is a genus of small epi being 37. Smith and Mickel (1977) have re phytes of the West Indies and Mexico to South ported n = 35 for Polypodium friedriehsthal America. Coehlidium rostratum is one of two ianum Kunze from Costa Rica, and two Old out of 16 species (Bishop, 1978) to bear coe World pleopeltids, Lepisorus (usually included nosori deeply immersed in a central groove. in Pleopeltis) and Belvisia, have species with Its chromosome number is 33 pairs at meiosis. 35 pairs of chromosomes. In both of the latter The most frequently encountered number in genera, however, a tendency towards aneu members of the Grammitidaceae is 37. How ploidy results in both lower and higher num ever, Walker (1966) found 33 pairs of chro bers in Lepisorus and a lower number in Bel mosomes in Grammitis hartii (Jenm.) Proctor. visia (Lovis, 1977). At the present time, the (A still lower number for a member of the number n = 35 tends to support the separation Grammitidaceae is reported below.) of Marginariopsis from Pleopeltis, XIPHOPTERIS Kaulfuss DICRANOGLOSSUM J. Smith (syn. Es ehatogramme Trev.) Xiphopteris limula Christ (Fig. 5, 6, 17, 18)-C. R. Cartago Provo Volcan Turrialba, Dieranoglossum panamense (c. Chr.) L. D. Santa Cruz, 15 November 1977, Wagner Gomez (Fig. 3, 15)-C. R. Limon Prov. Road 77552. Bishop (1978) placed Xiphopteris to Chase, 5 km from Bribri, 17 November 1977, serrulata (Swartz) Kaulf., the type spe Wagner 77588. Dicranoglossum , a genus cies of the genus, in Cochlidium, and the comprising six species of small epiphytes of chromosome number of that species has been shaded forests, is found in the New World trop reported as 74 pairs." Other authors have in ics from Cuba and Central America to Brazil.