Research Topics1. Cytogenetic

To plant biologists interested in patterns of genetic variation and evolution, analyses of chromosome numbers represent a fundamental step in the study of any group of organisms.

2. Floristic Studythe inventory and characterization of botanical diversity.

Chromosome Counting－ 1983. The ferns of Elden Mountain, Arizona. Am. Fern J.73: 85-92. － and C. G. Schaack. 1983. Miscellaneous chromosome counts for Adiantaceae, Aspleniaceae, Asteraceae, and Poaceae. Taxon 32: 664-665.

Reticulate EvolutionHaufler, C. H., － and E. W. Rabe. 1995. Reticulate evolution in the Polypodium vulgare complex. Systematic Botany 20: 89-109. Haufler, C. H., － and T. A. Ranker. 1990. Biosystematic analysis of the Cystopteris tennesseensis complex. Annals of the Missouri Botanical Garden 77: 31

4-329.Haufler, C. H. and － . 1991. New species of North American Cvstopteris and Polypodium, with comments on their reticulate relationships. Am. Fern J.81: 7-2

3.

Biosystematic Gastony, G. J and － . 1989. Species concepts in pteridophytes: The treatment and definition of agamosporous species. Am. Fern J.79: 65-77.－ 1986. The use of fern herbarium specimens in biosystematic research: Introduction. Am. Fern J.76: 101. － and C. H. Haufler. 1986. Biosystematic uses of fern gametophytes derived from herbarium specimens. Am. Fern J.76: 114-128. － , P. G. Wolf and T. A. Ranker. 1986. Factors affecting prolonged spore viability in herbarium collections of three species of Pellaea. Am. Fern J.76: 141-14

8.Werth, C. R. and － . 1991. A model for divergent allopatric speciation of polyploid pteridophytes resulting from silencing of duplicate-gene expression. The A

merican Naturalist 137: 515-526. Haufler, C. H., － , D. M. Britton and S. J Robinson. 1985. Triploidy and its evolutionary significance in Cystopteris protrusa. Canadian Journal of Botany 63: 1

855-1863. Paris, C. A. and － . 1988. A biosystematic investigation of the Adiantum pedatum complex in eastern North America. Systematic Botany 13: 240-255. Yatskievych, G., － and E. Wollenweber. 1990. A reconsideration of the genus Pitvrogramma Link (Adiantaceae) in the southwestern United States. Am. Fer

n J.80: 9-17.Benham, D. M. and － . 1992. Generic affinities of the star-scaled cloak ferns. Am. Fern J.82: 47-58.Ranker, T. A., S. F. Floyd, － and P. G. Trapp. 1994. Historical biogeography of Asplenium adiantum-nigrum (Aspleniaceae) in North America and implicatio

ns for speciation theory in homosporous pteridophytes. American Journal of Botany 81: 776-781. － & G. Yatskievych. 2003a. Chromosome studies of cheilanthoid ferns (Pteridaceae, Cheilanthoideae) from the western United States and Mexico. America

n Journal of Botany 90: 1788-1800.

TaxonomyYatskievych, G. and － . 1986. Notes on Arizona Pteridophyta. Journal of the Arizona Nevada Academy of Sciences 21: 19-21. － . 1987. Argyrochosma, a new genus of cheilanthoid ferns. Am. Fern J.77: 37-41. － . 1993. New taxa and nomenclatural changes in the North American fern flora. Contributions from the University of Michigan Herbarium 19: 31-61. Mendoza, A., － , B. Perez-Garcia and G. Yatskievych. 2001. Una nueva especie de Pellaea (Pteridaceae) del estado de San Luis Potosi, Mexico. Acta Bota

nica Mexicana 57: 15-21. － & G. Yatskievych. 2003b. Revised nomenclature and a new North American record for the Villose Cliff Brake (Pellaea. Pteridophyta). Novon 13: 358-362. － & G. Yatskievych. 2005. A novel hybrid Polypodium (Polypodiaceae) from Arizona. Am. Fern J.95: 57-67.

Author or coauthor of Flora of North AmericaPteridaceae : Astrolepis, Notholaena, Argyrochosma, Pellaea, Cheilanthes, Pityrogramma, PentagrammaPolypodiaceae : Pleopeltis, PolypodiumWoodsiaceae : Cystopteris, WoodsiaCoauthor of The Pteridophytes of Mexico.Woodsiaceae : Woodsia

TaxonomyYatskievych, G. and － . 1986. Notes on Arizona Pteridophyta. Journal of the Arizona N

evada Academy of Sciences 21: 19-21. － . 1987. Argyrochosma, a new genus of cheilanthoid ferns. Am. Fern J.77: 37-41. － . 1993. New taxa and nomenclatural changes in the North American fern flora. Contrib

utions from the University of Michigan Herbarium 19: 31-61. Mendoza, A., － , B. Perez-Garcia and G. Yatskievych. 2001. Una nueva especie de Pel

laea (Pteridaceae) del estado de San Luis Potosi, Mexico. Acta Botanica Mexicana 57: 15-21.

－ & G. Yatskievych. 2003b. Revised nomenclature and a new North American record for the Villose Cliff Brake (Pellaea. Pteridophyta). Novon 13: 358-362.

－ & G. Yatskievych. 2005. A novel hybrid Polypodium (Polypodiaceae) from Arizona. Am. Fern J.95: 57-67.

Author or coauthor of Flora of North AmericaPteridaceae : Astrolepis, Notholaena, Argyrochosma, Pellaea, Cheilanthes, Pityrogramm

a, PentagrammaPolypodiaceae : Pleopeltis, PolypodiumWoodsiaceae : Cystopteris, Woodsia

Coauthor of The Pteridophytes of Mexico.Woodsiaceae : Woodsia

UNA NUEVA ESPECIE DE PELLAEA (PTERIDACEAE) DEL ESTADO DE SAN LUIS POTOSÍ, MÉXICO

MENDOZA, A., M. WINDHAM, B. PÉREZ-GARCÍA and G. YATSKIEVYCHActa Botanica Mexicana (2001), 57: 15-21

A new species of Pellaea sect. Pellaea is described and illustrated, Pellaea ribae Mendoza & Windham, which is clearly distinguished from other species in the genus by the abundant white trichomes present on the petiole, rachis, and abaxial surface of the leaves. Presently P. ribae is known only from two localities in San Luis Potosí, Mexico. Possible relationships of this new species to other cheilanthoid ferns are discussed.

Argyrochosma, a New Genus of Cheilanthoid FernsM. D. Windham

American Fern Journal 77: 37-41. (1987)

• Genus Type: Notholaena nivea (Poir.) Desv.• Treated as Notholaena, Acrostichum, Cheilanthes, Gymnogramma and o

thers no longer in use.• Most taxonomist placed N. nivea group in Notholaena or Pellaea by m

orphological characters and recognized it as a natural group as sect. Argyrochosma also including some Pellaea species.

• Several distinct sporophyte morphological characters: rhizome scales, leaf architecture, sporangial distribution, spore morphology, plus chromosome number, gametophyte morphology, chemical composition of the farinose indument and patterns of variablity at conservatie enzyme loci, lead the author raised sect. Argyrochosma as a genus.

Windham and Yatskievych. 2003. Chromosome studies on cheilanthoid ferns (Pteridaceae: Cheilanthoideae) from the Western United States and Mexico. Amer. J. Bot. 90: 1788-1800.

2. Pityrogramma trifoliata (Linnaeus) R. M. Tryon, Contr. Gray Herb. 189: 68. 1962. Goldenrod fern Acrostichum trifoliatum Linnaeus, Sp. Pl. 2: 1070. 1753; Trismeria trifoliata (Linnaeus) DielsLeaves 50--150 cm, rarely longer. Petiole reddish brown, shiny, scaly. Blade linear-lanceolate, 1-pinnate apically to 2-pinnate proximally; fertile pinnae white-farinose abaxially. Pinnae palmate to pinnate, 2--3(--7)-foliolate proximally, with flat surfaces parallel to ground, undivided and linear apically. Pinnules short-stalked, linear, occasionally lobed proximally, margins serrulate. Spores with perispore sparsely granular, lacking equatorial flange. 2 n = 116.

Roadside ditches and canal banks; 0--20 m; Fla.; Mexico; West Indies in Greater Antilles; Central America; South America.

The range of Pityrogramma trifoliata is apparently actively expanding in Florida, perhaps in response to increasing disturbance of wetland habitats. A good case can be made for segregation of this species into the monotypic Trismeria Fée, based on differences in dissection of leaves and spore ornamentation noted in the descriptions of this and the previous species. Because naturally occurring hybrids with four other Pityrogramma taxa are known to exist in the tropics, P . trifoliata is retained in Pityrogramma for the present, following R. M. Tryon (1962), who viewed P . trifoliata as a specialized offshoot of the core group of species in the genus. Elsewhere, plants are known with yellow farina on the fertile leaves, but these are nowhere common.

Chromosome Counting－ 1983. The ferns of Elden Mountain, Arizona. Am. Fern J.73: 85-92. － and C. G. Schaack. 1983. Miscellaneous chromosome counts for Adiantaceae, Asple

niaceae, Asteraceae, and Poaceae. Taxon 32: 664-665.

Reticulate EvolutionHaufler, C. H., － and E. W. Rabe. 1995. Reticulate evolution in the Polypodium vulgare

complex. Systematic Botany 20: 89-109. Haufler, C. H., － and T. A. Ranker. 1990. Biosystematic analysis of the Cystopteris ten

nesseensis complex. Annals of the Missouri Botanical Garden 77: 314-329.Haufler, C. H. and － . 1991. New species of North American Cvstopteris and Polypodiu

m, with comments on their reticulate relationships. Am. Fern J.81: 7-23. Haufler, C. H., － , D. M. Britton and S. J Robinson. 1985. Triploidy and its evolutionary

significance in Cystopteris protrusa. Canadian Journal of Botany 63: 1855-1863.

The Ferns of Elden Mountain, Arizona.M. D. Windham

Amer. Fern J. 73: 85-93. (1983)

• Elden Mountain Fern Flora: 20 species representing 11 genera• Chromosome count: 15 species• Spore count: 19 speciesAdiantum pedatum L. s64; Asplenium adiantum-nigrum L. s64, n72II; A. platyneuron (L.) B.S.P. s64, n36II;A. resiliens Kunze s32-64*, n108I; A. septentrionale (L.) Hoffm. s64, n72II;A. trichomanes L. subsp. trichomanes s64, n36IICheilanthes eatonii Bak. s32; C. feei Moore s32; C. fendleri Hook. s64, n30II; C. wootonii Maxon s32, n90II; Cystopteris cf. tennesseensis Shaver s64-32*, n84II = C. utahensis Windham & Haufler (1991)Dryopteris filix-mas (L.) Schott s64, n82IIPellaea truncata Goodding s64-32*, n29II; P. wrightiana Hook. s64-32*, n58II;Pityrogramma triangularis (Kaulf.) Maxon s64;Polypodium hesperium Maxon s64, n74II;Polystichum scopulinum (D.C. Eaton) Maxon s64, n82II;Pteridium aquilinum (L.) Kuhn var. pubescens Underw extinct.Woodsia mexicana Fee s64, n82; W. oregana D.C. Eaton s64, n76II

•C. tennesseensis first recognized by Shaver (1950) who described it as a hybrid between C. bulbifera and C. protrusa.

•Disagreement: McGregor et Weatherby reduced it as variety of C. fragilis.•Blasdell (1963) revealed C. tennesseensis as tetraploid: hybridization between C. protrusa and C. bulbifera

In this paper, the authors reported a series of analyses designed to investigate the patterns and the processes behind the systematic confusion in C. tennesseensis. The authors described studies of biogeography, morphology, chromosomes, and isozymes that clarify the origin and current status of C. tennesseensis and its progenitor diploids.

Annals of the Missouri Botanical Garden, Vol. 77, No. 2. (1990), pp. 314-329.

Materials and Methods

• Morphological variability, geographic distribution and habitat diversity from herbarium specimens.

• Spore lengths (max.) were recorded

For chromosomal, isozymic and garden morphological comparisons, transplants into Univ. Kansas greenhouses.

• Analyses morphological variation based on plants cultivated in the greenhouse: 21 qualitative characters

• The transplants are also prepared for chromosome counting and isozyme experiments.

Materials and Methods

• Morphological variability, geographic distribution and habitat diversity from herbarium specimens.

• Spore lengths (max.) were recorded

• For chromosomal, isozymic and garden morphological comparisons, transplants into Univ. Kansas greenhouses.

• Analyses morphological variation based on plants cultivated in the greenhouse: 21 qualitative characters

• The transplants are also prepared for chromosome counting and isozyme experiments.

Results: Spores

Materials and Methods

• Morphological variability, geographic distribution and habitat diversity from herbarium specimens.

• Spore lengths (max.) were recorded

• For chromosomal, isozymic and garden morphological comparisons, transplants into Univ. Kansas greenhouses.

• Analyses morphological variation based on plants cultivated in the greenhouse: 21 qualitative characters

Meiotic Chromosome Behavior

C. bulbiferaN=42

C. tennesseensisN=84

C. protrusaN=42

Meiotic Chromosome Behavior

Isozymes22 enzymes were examined and 9 provided consistent, interpretable results.HK, IDH, LAP, PGI-2, PGM-1, PGM-2, SkDH, TPI-1, TPI-2

Gametophytes express the same isozymes as do sporophytes

Materials and Methods

• Morphological variability, geographic distribution and habitat diversity from herbarium specimens.

• Spore lengths (max.) were recorded

• For chromosomal, isozymic and garden morphological comparisons, transplants into Univ. Kansas greenhouses.

• Analyses morphological variation based on plants cultivated in the greenhouse: 21 qualitative characters

Gametophytes

• None of the 100 isolated gametophytes of C. bulbifera and C. protrusa produced sporophytes (4 months) but both are hermaphroditic.

• 25 of 100 isolated gametophytes of C. tennesseensis did form sporophytes and lower genetic load than diploides ones.

C. bulbiferaBB 2x = 42II

C. tennesseensisBBPP 4x = 84II

C. protrusaPP 2x = 42II

C. utahensisBBRR 4x = 84 II

C. laurentianaBBNNRR 6x = 126 II

C. reevesianaRR 2x = 42IIRRRR 4x = 84 II

C. fragilisNNRR 4x = 84 II

C. “hemifragilis”NN 2x = 42II

C. tenuisNNPP 4x = 84 II

BBP

BPPx wagneriBNPP 3x = 42I + 42II

x illinoensisBNP 3x = 126I

fragilis x tenuisNNPR 4x = 84I + 42II

BNMR

NRR

Cystopteris fragilis complex (Haufler & Windham, 1991; Lellinger, 1985)