WRIGHTIA A BoTrANICAL JOURNAL 1945-1951 VouuME I EDITOR Cyrus LoNGwortH LUNDELL University Press IN DALLAS SOUTHERN METHODIST UNIVERSITY Dauuas, TEXAS CONTENTS OF VOLUME I Complete in Five Numbers Number 1, August 15, 1945 The genus Garcia Vahl, a potential source of superior hard quick- aryme ou. By Cyrus Loneworth Lundell... 3.3... 5 eo ss The genus Triodanis Rafinesque, and its relationships to Specularia and Acompanuia: fy Hogers MeVaugh... 2. 2... os ee oe ce New phanerogams from Texas, Mexico and Central America. By Cyrus Longworth Lundell Notes and News Number 2, April 30, 1946 Revision of the genus Chaetopappa DC. By Lloyd H. Shinners Revision of the genus Leucelene Greene. By Lloyd H. Shinners..... The genus Dichaetophora A. Gray and its relationships. By Lloyd H. EI os a Revision of the genus Aphanostephus DC. By Lloyd H. Shinners... Revision of the genus Kuhnia L. By Lloyd H. Shinners............ Plants of Mexico and Central America—I. By Cyrus Longworth RAN 6 a a ee A useful method for drying plant specimens in the field. By Cyrus Longworth Lundell Notes and News Number 3, December 15, 1947 A new species of Dracocephalum L. from Texas. By Cyrus Longworth ee ee ee Additions to the orchids of Texas. By Donovan S. Correll.......... Two anomalous new species of Erigeron L. from Texas. By Lloyd H. Oe ee es Number 4, June 30, 1948 Wild Rose Pass of Texas, an obscure locality and its actual position. By Rogers MeVaugh oe re ees Curly mesquite grass in Texas and Northern Mexico. By V. L. Cory A new species of Chrysopsis Nuttall from the driftless area of Wiscon- ee ee i ee ee awe ct kee Additional notes on the Eriocaulaceae, Avicenniaceae and Ver- benaceae of Texas—I. By Harold N. Moldenke........... I22 145 161 163 165 166 183 187 206 207 214 218 220 iv CONTENTS Number 5, May 15, 1949 A preliminary survey of the distribution of Texas Pteridophyta. Dy nee ON, a a 247 Notes INDEX TO VOLUME I, WRIGHTIA December 1, 1951 ! Abronia, 53 Ameliae, 54 fragrans, 55 Acacia, 242 Acalypha, 58 Achaetogeron, 63, 80, 91 Acrodiclidium, 145 glaberrimum, 147 lucidum, 147 mexicanum, 147 misantlae, 147 Adder’s-tongue family, 268 Adenophora, 14 Adiantum, 255 Capillus-Veneris, 252, 253, 255, 261 modestum, 255 pedatum, 278 tricholepis, 253, 256 tricholepis f. glabrum, 253, 256 Adopogon, 187 amplexicaule, 194 carolinianum, 1908 Dandelion, 190 montanum, 192 occidentale, 200 virginicum, 194 Aleurites, 7, 12 cordata, 9 Fordii, 9 Aloysia ligustrina, 242 ligustrina var. Schulzii, 242, 243 macrostachya, 242 Wrightii, 242 Ames, Oakes, 166 Amphiachyris, 63 Anchistea virginica, 267 Anemia mexicana, 253, 267 Antennaria plantaginifolia, 70 Aphanostephus, 74, 90, 91, 92, 95, 121, 186 arizonicus, 95, 96, 98, TIO, FEY, 143, 1 F4 FOO, 104, 1On March 1, 1948, Dr. C. L. Lundell resigned as Professor of Botany and Director of the University Herbarium of Southern Methodist University to devote full time to his duties as Director and Head Botanist of the Texas Research Foundation. He had served in his position at Southern Methodist University on a dollar a year basis since July 1, 1946, the date the Foundation became an independent non-profit research and educational institution, chartered by the State of Texas. The Lundell Herbarium, which had been deposited on loan at Southern Methodist University, was transferred to the Texas Research Foundation at Renner in 1948. Com- prising principally tropical American collections of the Lundells, Perey H. Gentle, Eizi Matuda, and George B. Hinton, it contains types or isotypes of most of the species described by Lundell. This herbarium contains approximately 25,000 specimens from Texas, Mexico, and Central America, and about 5,000 specimens from other areas. Aside from the strictly tropical groups, grasses are being given primary consideration in the development of the collection. With the withdrawal of the Lundell Herbarium, the University Herbarium of Southern Methodist University is largely restricted to collections from temperate North America, with Texas specimens predominating. On July 1, 1949, Dr. Lloyd H. Shinners was appointed Director of the University Herbarium. All communications regarding exchanges with the Lundell Herbarium, as well as requests for loans, should be sent to the Texas Research Foundation, P. O. Box 43, Renner, Texas. 281 282 arkansanus, 102 arkansanus var. Hallii, 102 humilis, 96, 98, 111, 114, 116 jaliscensis, 99, 115, 116 Kidderi, 96, 100, 107, 110 pachyrrhizus, 99, 117 perennis, 119 pilosus, 99, 101 pinulensis, 121 potosinus, 98, 108 pulchellus, to1 ramosissimus, 74, 96, 98, 100, 102 104, 107, 108, 109, 113, I19 ramosus, 99, 114, 116 Riddellii, 98, 99, 101, 119 skirrhobasis, 74, 95, 96, 100, 102 104, 110, 413 skirrhobasis var. Hallii, 102 skirrhobasis f. incisifolius, 103 skirrhobasis f. quasigigantiusculus, 103 skirrhobasis 100, 106 Apogon, 188, 200 gracilis, 205 humilis, 188, 202 lyratum, 202 Wrightii, 202 Aptilon, 188 Arethusa racemosa, 170 Aspidium juglandifolium, 265 trifoliatum, 267 Asplenium, 256 asplenioides, 257 ebeneum, 256 Palmeri, 253, 256 parvulum, 256 pinnatifidum, 278 platyneuron, 249, 253, 256 resiliens, 252, 253, 256 trichomanes, 253, 257 Aster, 63, 82, 90, 96 arenosus, 85 bellus, 84, 85 ericaefolius, 82, 85 ericaefolius var. tenuis, 84, 85 hirtifolius, 85 Leucelene, 82, 85 scopulorum, 89 Astranthium,63, 74, 90, 91, 92, 95, 121 integrifolium, 74, 90 ? > var. thalassius, 098, WRIGHTIA xylopodum, 81, 121 Asyneuma, 14 Athyrium, 257 acrostichoides, 275 asplenioides, 249, 253, 257, 275 Filix-femina, 257, 275 Avicenniaceae, 220, 221 Avellano, 4 Avicennia nitida, 221 Azolla, 269 caroliniana, 253, 269 Azorina, 14 Bellis, 63 Bernardia, 53 aurantiaca, 57, 58 interrupta, 58 Biche rayo, 57 Blephariglottis lacera, 167 Boltonia, 90, 91, 92, 93 campestris, 94 Bommeria, 257 hispida, 253, 257, 276 Botrychium, 268 dissectum var. 268 obliquum, 268 obliquum var. tenutfolium, 268 virginianum, 253, 268 Bouchea linifolia, 244 spathulata, 244 Bourdonta, 64, 65 bellidifolia, 63, 71 effusa, 64, 68 Brachyactis, 63 Brachychaeta, 63 Brazoria, 53 arenaria, 59, 60 truncata, 60 Brickellia, 122 grandiflora, 122 hymenochlaena, 122 monocephala, 122 Buffalo grass, 217 tenuifolium, Callicarpa americana, 245 americana var. lactea, 245 Calopogon pulchellus, 177 Campanula, 13, 51 americana, 20 amplexicaulis, 34 253) INDEX angulata, 51 biflora, 31, 33 canescens, 19, 20 coloradoense, 25, 27, 28 colorata, 19, 20 dimorphantha, 19, 20 Erinus, 17, 20 falcata, 29 fastigiata, 20 flagellaris, 34 intermedia, 31, 33; 34 leptocarpa, 47 ludoviciana, 31, 33 montevidensis, 31 pentagonia, 17, 20 perfoliata, 33, 34 ramosissima, 17, 20, 28 Reverchoni, 28 syriaca, 28 Campanulaceae, 13 Campanulastrum, 14 Campylocera, 14, 21 leptocarpa, 47, 49, 5° leptocarpa var. glabella, 48, 5° Celastrus, 155 lenticellatus, 156 longipes, 156 siltepecanus, 155, 156 Centaurium, 18 Chaetanthera, 65 Chaetopappa, 63-81, 82, 90, 121 asteroides, 63, 64, 66, 67, 72, 73, 74 asteroides var. grandis, 67, 76, 77 asteroides var. imberbis, 63, 67, 73, 74, 79, 77 bellidifolia, 66, 71, 74, 121 bellioides, 67, 76, 77, 82 bellioides var. hirticaulis, 67, 79 effusa, 66, 68, 70, 121 keerlioides, 66, 69 Hersheyi, 66, 67 modesta, 76, 77 Parryi, 66, 70 pulchella, 67, 79 Chaetophora, 65 asteroides, 66, 73 Chanekia caudata, 146 coriacea, 146 Chaperno, 155 blanco, 154 negro, 155 283 Cheilanthes, 251, 257 aemula, 253 alabamensis, 252, 253, 257, 275 aspera, 258 castanea, 258 Eatoni, 252, 253, 257, 258 Eatoni f. castanea, 253, 258 Feei, 253, 258 Fendleri, 253, 258 gracilis, 258 horridula, 253, 258 horridula f. compacta, 248, 258 Kaulfussii, 253, 250 lanosa, 253, 259 lanuginosa, 258 lendigera, 253, 259 leucopoda, 253, 259 Lindheimeri, 253, 259 microphylla, 248, 275 Moritziana, 257 myriophylla, 275 tomentosa, 252, 253, 2590 tomentosa var. Eatoni, 257 villosa, 253, 259 Wootoni, 253, 260 Wrightii, 253, 260 Chilopsis, 242 Chrysanthemum Parthenium, 121 Chrysopsis, 82, 90, 218, 219 Ballardii, 219 ertcoides, 84 hispida, 219 wisconsinensis, 218, 219 Cinnamon-fern family, 268 Citharexylum Berlandieri, 244 brachyanthum, 244 brachyanthum var. glabrum, 244 spathulatum, 244 Clerodendrum frangrans var. pleni- florum, 246 indicum, 246 Clubmoss family, 272 Coleosanthus, 122 Conyza, 63 Corallorhiza, 177, 178 grandiflora, 178 Wisteriana, 177 Corallorrhiza, 178 Correll, Donovan 8., 166, 247 Helen B., 166, 248 Cory, V. L., 163, 206, 214 284 Critonia, 123, 134 Kuhnia, 133 Cryosophila, 53 bifurcata, 53, 54 Cookii, 54 Warscewiczil, 54 Cuatlaguilocuatl, 4 Curly-grass family, 267 mesquite grass, 214 Cylindrocarpa, 14 Cymbia, 187, 188, 200 occidentalis, 188, 200 Cynthia, 187, 188 amplexicaulis, 194 Boscti, 190 Dandelion, 188, 190 Dandelion var. montana, 192 falcata, 194 Griffithii, 194 lyrata, 190 montana, 192 virginica, 188, 189, 194 viridis, 194 Cypripedium Calceolus var. pubes- cens, 166 Cystopteris, 260 bulbifera, 278 fragilis, 253, 260 fragilis var. protrusa, 253, 260 fragilis f. simulans, 253, 260 Dalea, 123 Dichaetophora, 63, 90, 91, 92, 93, 95 campestris, 90, 93, 94 Diosphaera, 14 Diplazium acrostichoides, 275 Diplopappus, 82 ericoides, 84 ertcoides var. hirtella, 84 Diplostelma, 64, 65 bellioides, 64, 77 filiformis, 73, 74 pumila, 73 radians, 73 Distasis, 64, 65 asteroides, 73 concinna, 80 heterophylla, 80 modesta, 64, 76, 77 Parryi, 70 Downingia, 51 WRIGHTIA Dracocephalum, 165 Correllii, 165 virginianum, 165 Dryopteris, 260 augescens var. Lindheimert, 248, 261 cristata, 253, 260 dentata, 253, 260, 261 Filix-mas, 253, 260 floridana, 275 hexagonoptera, 254, 260 intermedia, 276 ludoviciana, 248, 275 marginalis, 278 normalis, 253, 254, 255, 261 normalis var. Lindheimeri, 261 noveboracensis, 276 parasitica, 260 patens, 261 spinulosa var. intermedia, 276 thelypteris var. Haleana, 254, 261 versicolor, 254, 261 Duranta repens, 244 repens var. alba, 244 Dysmicodon, 21 californicum, 31, 33 ovatum, 31, 33 perfoliatum, 34 Egletes, 90, 91, 92, 186 arkansana, 102 humilis, 114 ramosissima, 109 Endlicheria, 145 Browniana, 146 zapoteoides, 145 Epidendrum conopseum, 178 Epipactis gigantea, 169 Equisetaceae, 271 Equisetum, 271 hyemale, 272 hyemale var. affine, 272 hyemale var. intermedium, 271 hyemale var. robustum, 272 - kansanum, 254, 271 kansanum f. caespitosum, 254, 271 kansanum f. variegatoides, 254, 274 laevigatum, 253, 254, 271 prealtum, 252, 254, 271, 272 prealtum f. Drummondii, 254, 272 prealtum f. texana, 254, 272 robustum, 272 Pe ae ee Eremiastrum, 63 Erigeron, 63, 80, 90, 91, 92, 94, 95; 183-186 concinnus, 80 philadelphicus, ot Bigelovii, 91 coronarius, 91, 184 Geiseri, 183, 184 Geiseri var. calcicola, 184 mimegletes, 184, 186 pumilus, 91 strigosus, 186 tenuis, 184 Eriocaulaceae, 220, 221 Eriocaulon compressum, 221 decangulare, 221 Kornickianum, 221 texense, 221 Euaster, 82 Eucephalus ericoides, 84 Eupatorium, 123 alternifolium, 123, 132, 133 altissimum, 123 canescens, 123, 143 cubense, 143 oblongifolium, 143 Euthamia, 63 Fallugia, 242 paradoxa, 213 Fassett, N. C., 61 Favratia, 14 Ferns, Texas, 247-278 Filix fragilis, 260 Fimbriella lacera, 167 Flora of Texas, 166, 220, 247 Fondren, Mrs. W. W., 163 French mulberry, 245 Garcia, I-12 Mayana, 3,4 nutans, I-12 parviflora, 6, 12 Garcia oil, 7-12 Giant spiral-orchid, 175 Give-and-take, 54 Glandular Ponthieva, 170 Glandulous Neottia, 170 Golden dewdrop, 244 Green fringed-orchis, 169 Gutierrezia, 63 Gymnogramma Ehrenbergiana, 276 INDEX Gymnopteris Ehrenbergiana, 276 hispida, 257 Gyrostachys brevifolia, 173 Habenaria Andrewsii, 168 Chapmanii, 166 clavellata, 167 cristata, 167 flava, 167 lacera, 167 nivea, 169 quinqueseta, 169 repens, 169 Heterocodon, 14, 19, 20 rariflorum, 19 Hexalectris grandiflora, 178 mexicana, 178 nitida, 179, 180 revoluta, 179, 180, 181 spicata, 179, 181 Warnockii, 178, 181 Hilaria Belangeri, 214-217 cenchroides, 215, 216 cenchroides var. texana, 216 mutica, 217 Swalleni, 215-217 Hildreth, A. C., 61 Hoffmannseggia, 127 Horsetail family, 271 Huevo de gato, 4 Hyoseris amplexicaults, 193, 194 angustifolia, 190 biflora, 193 caroliniana, 197, 198 major, 190 montana, 192 prenanthoides, 193 ramosissima, 197 virginica, 194, 197 Hypericum aphyllum, 53, 58 lbidium longilabris, 173 285 Institute of Technology and Plant Industry, 206 Inula, 82 ericoides, 84 Ionactis alpina, 80 Isoetaceae, 274 Isoetes, 274 lithophila, 254, 274 melanopoda, 254, 275 Isopappus, 90 286 WRIGHTIA Isotria verticillata, 170 James, Mrs. B. W., 206 Jaumea linearis, 143 Keerlia, 65, 97, 117, 121 bellidifolia, 63, 71, 73, 121 effusa, 68, 121 linearifolia, 121 mexicana, 81, 121 ramosa, 116 skirrhobasis, 102, 104 Krigia, 63, 187-206 amplexicaulis, 194 bellidioides, 200 biflora, 188, 189, 193-197 biflora f. glandulifera, 194 caroliniana, 197 caroliniana var. leptophylla, 197 Dandelion, 188, 189, 190-192 dichotoma, 197 gracilis, 189, 205, 206 integrifolia, 194 leptophylla, 197 montana, 189, 192-193 nervosa, 200 occidentalis, 187, 188, 189, 200 occidentalis var. mutica, 200 occidentalis f. mutica, 200 oppositifolia, 187-189, 201, 202-205, 206 petiolaris, 202 tenella, 197 virginica, 77, 187, 188, 1 89, 193, 194, 107 virginica var. dichotoma, 197 Kuhnia, 122-144 adenolepis, 122, 127, 141, 142 albicaulis, 133 altaica, 123, 133 arabica, 142 arguta, 142 baccharoides, 142 chlorolepis, 125, 126, 120, 131, 238 139, 144 cinerea, 133 Critonia, 133 dasypia, 133 divaricata, 133 elliptica, 133 elliptica var. montana, 133 ? eupatorioides, 123, 124, 132, 133, 134 eupatorioides var. angustifolia, 133, 134 eupatorioides var. corymbosa, 133, 134 eupatoriodes var. corymbulosa, 123, 125, 131, 136, 137-139, 141 eupatorioides var. glutinosa, 133, 138 eupatorioides var. gracilis, 133, 134 eupatorioides var. gracillima, 128 eupatorioides var. ozarkana, 122, 125, 136, 137 eupatorioides var. pyramidalis, 123, 125, 132, 134, 138, 139, 141 eupatorioides var. texana, 122, 125, 136, 137, 139 Fitzpatricki, 138 frutescens, 142 fruticosa, 142 fulva, 133 glabra, 133 glabrata, 143 glomerata, 133 glutinosa, 123, 133, 138 Gooddingi, 138 hirtiflora, 143 Hitchcocki, 138 icosantha, 143 jacobaea, 138 Kuhnia, 133 latifolia, 133 leptophylla, 124, 128 leptophylla var. mexicana, 126, 128 lineana, 133 linearifolia, 143 podocarpa, 143 macrantha, 138 Marximiliani, 138 media, 133 microphylla, 126, 127, 144 Mosieri, 125, 132 multiramea, 143 oreithales, 126, 127, 131 paniculata, 133, 134 pinnata, 143 pubescens, 133 pyrifolia, 143 reticulata, 138 rosemartaefolia, 132 . rosmarinifolia, 123, 129, 142, 143 INDEX 287 rosmarinifolia var. chlorolepis, 129 rosmarinifolia var. gracillima, 128 Schaffneri, 127, 144 spicata, 144 suaveolens, 123, 138 triplinervis, 131 tuberosa, 123, 133 villosa, 144 virgata, 133 Lachnocaulon anceps, 221 Lantana, 236 achyranthifolia, 237 camara, 236 camara var. mista, 236 citrosa, 236, 237 horrida, 235 macropoda, 236 rubra, 235 Larrea, 242 Lauraceae, 145 Laurus Cerventesit, 146 Lazxanon, 188 diversifolium, 202 heterophyllum, 202 parviflorum, 188, 202 Legousia coloradoensis, 25 falcata, 29 Juliani, 51 Legouzia biflora, 31 coloradoense, 25 leptocarpa, 48 perfoliata, 34 Leontodon Dandelion, 190 Leucelene, 63, 82, 90 alpina, 89 alsinoides, 84, 85 arenosa, 84, 85 ericoides, 82, 84, 99, I19 ericoides var. serotina, 84, 85 ericoides var. tenius, 85 hirtella, 85 serotina, 85 Leucopsidium, 95, 97 arkansanum, 102 humile, 114 Liatris, 123 spicata, 144 Licaria, 145 Cufodontisit, 147 excelsa, 147 Lippia alba, 237 graveolens, 237 javanica, 237 reptans, 241 scabra, 237 strigulosa, 241 Lonchocarpus, 53, 145 apricus, 57 belizensis, 55 chiapensis, 152, 153 eruentus, 55 Gillyi, 56, 57, 154 hidalgensis, 153, 154 latifolius, 55 Michelianus, 153 nicaraguensis, 153, 154 rugosus, 57, 154 sericeus, 56 Whitei, 154 Lorinseria, 261 areolata, 249, 254, 261 Lundell, Amelia A., 206 Cyrus Longworth, 1, 53, 145, 161, 165, 206, 247, 281 Lundell Herbarium, 281 Luthera, 188 virginica, 194 Lycopodiaceae, 272 Lycopodium, 272 adpressum, 254, 272 alopecuroides, 254, 272 alopecuroides var. adpressum, 272 alopecuroides var. pinnatum, 254, 272 earolinianum, 249, 254, 272 inundatum var. Bigelovii, 276 lucidulum, 276 Lygodium, 268 japonicum, 254, 268, 276 palmatum, 276 MeVaugh, Rogers, 13, 61, 207 Machaeranthera, 90 Malaxis unifolia, 178 Mangle negro, 221 Many-flowered Ponthieva, 170 Marsileaceae, 269 Marsilea, 269 macrocarpa, 270 macropoda, 254, 269 mucronata, 270 288 tenuifolia, 254, 270 uncinata, 252, 254, 270 vestita, 252, 254, 270 vestita var. tenuifolia, 270 Maytenus, 145 belizensis, 156 chiapensis, 156 guatemalensis, 156 Purpusii, 156 Schippii, 156 Michauxia, 14 Microtropis, 145 areolata, 156, 157 contracta, 157, 158 guatemalensis, 157, 158 occidentalis, 158, 160 stipitata, 145, 158, 159 Misanteca, 145 capitata, 145 caudata, 146 Cervantesii, 146 coriacea, 146 Cufodontisii, 146 excelsa, 147 glaberrima, 147 lucida, 147 mexicana, 147 misantlae, 147 Moldenke, Harold N., 220 Monoptilon, 63 Morton, C. V., 278 Muller, C. H., 61 Munnell, Mrs. Ruth G., 206 Nectandra, 145 albiflora, 147, 148 glandulosa, 148 globosa, 148 Lundellii, 148, 149 Matudai, 148, 149 rubriflora, 149 salicifolia, 149 Skutchii, 149 Standleyi, 149 Neottia glandulosa, 170 grandiflora, 178, 179 Nephrodium floridanum, 275 patens, 261 thelypteris, 261 Nephrolepis, 261 exaltata, 254, 261 WRIGHTIA Notholaena, 251, 262 Aschenborniana, 254, 262 aurea, 254, 262 bonariensis, 262 candida, 254, 262 dealbata, 254, 262 Fendleri, 276 ferruginea, 262 Grayi, 254, 262 Greggii, 254, 262 Hookeri, 263 limitanea, 254, 262 neglecta, 254, 262 nivea var. dealbata, 262 Pringlei, 262 Schaffneri, 263 Schaffneri var. Nealleyi, 254, 263 sinuata, 254, 263 sinuata var. cochisensis, 254, 263 sinuata var. integerrima, 253, 254, 263 Standleyi, 254, 263 Ocotea, 145, 149 rubriflora, 149 Onoclea, 263 sensibilis, 249, 254, 263 Ophioglossaceae, 268 Ophioglossum, 269 crotalophoroides, 254, 269 Engelmanni, 254, 269 nudicaule var. tenerum, 254, 269 pusillum, 269 tenerum, 269 vulgatum, 254, 269 Ophryosporus triangularis, 143 Orchids, 166-182 Orchis lacera, 167, 168, 169 psychodes, 167 Orcujuela, 244 Osmundaceae, 268 Osmunda, 268 ; cinnamomea, 249, 254, 260, 268 regalis, 268 regalis var. spectabilis, 249, 254, 268 Ostrowskia, 14 Palo de sangre, 56 Panicum euchlamydeum, 219 Parathesis, 53, 145 membranacea, 58, 59 platyphylla, 160 pleurobotryosa, 59 serrulata, 59, 160 Pascualito, 4 Pegolettia senegalensis, 142 Pellaea, 251, 262, 264 aspera, 258 aspera f. compacta, 258 atropurpurea, 252, 254, 264 cardiomorpha, 254, 264 cordata, 264 dealbata, 262 flexuosa, 265 glabella, 254, 264 intermedia, 264 intermedia f. pubescens, 254, 264 longimucronata, 255, 264 microphylla, 255, 264 mucronata, 276 ovata, 255, 265 pulchella, 264 ternifolia, 255, 265 Wrightiana, 252, 253, 255, 265 Pentagonia biflora, 31 coloradensis, 25 falcata, 29 leptocarpa, 48 perfoliata, 34 Pepperwort family, 269 Pepita del indio, 4 Persea, 145, 150 chiapensis, 150 chrysobalanoides, 151 flavifolia, 150 Liebmanni, 150 pallida, 151 rigens, 151 Petalostemum corymbosum, 143 Petromarula, 14 Phanerophlebia, 265 auriculata, 255, 265 umbonata, 255, 265 Phegopteris hexagonoptera, 260 Phlox, 165 Phoebe, 145 acuminatissima, 152 siltepecana, 151, 152 Phyla cuneifolia, 239, 240 incisa, 238, 240 lanceolata, 238 nodiflora, 240 INDEX 289 nodiflora var. reptans, 240, 241 strigulosa, 241 strigulosa var. parvifolia, 241 yucatana, 241 yucatana var. parvifolia, 241 Phyteuma, 14 Phytologia, 220 Pilularia, 270 americana, 255, 270 Pifion, 4 Pifoncillo, 4 Platanthera psychodes, 167 Plant dryer, 161, 162 Platanthera lacera, 167 Pluchea, ot Polyactidium, 94 Seemannii, 94 Polypodiaceae, 255 Polypodium, 265 erythrolepis, 255, 265 furfuraceum, 276 incanum, 266 polypodioides, 178, 266 polypodioides var. Michauxianum, 253, 255, 265 thyssanolepis, 255, 266 virginianum, 277 vulgare f. pygmaeum, 248, 277 Polystichum, 266 acrostichoides, 249, 255, 266 acrostichoides f. incisum, 255, 266 Polytrichon, 276 Ponthieva Brittonae, 170 glandulosa, 170 racemosa, 170, 171 Prismatocarpus falcatus, 28, 30 perfoliatus, 34 scaber, 29 Priva lappulacea, 244 Psoralea, 127 Pteridium, 266 aquilinum var. lanuginosum, 266 -aquilinum var. pseudocaudatum, 255, 266 aquilinum var. pubescens, 255, 266 latiusculum var. pseudocaudatum, 266 Pteridophyta, Texas, 247-278 Pteris, 266 aquilina var. pseudocaudata, 266 multifida, 255, 266 290 WRIGHTIA Quillwort family, 274 Sicyocodon, 14 Sideranthus, 90 j Ricinus communis, 1 Solidago, 63, 96 Ragged fringed-orchis, 169 Soxman, G. M., 247 Ragged-orchis, 169 Specularia, 13, 21 Rosa blanda, 213 Rotantha, 14 Sabal texana, 225 Salgado, Eduardo, 145 Salvinia family, 260 Salviniaceae, 269 Sapodilla, 146 Sarothra, 53 aphylla, 58 Schizaeaceae, 267 Scorzonera lanata, 188 Scutellaria, 53 petenensis, 60 Selaginellaceae, 273 Selaginella, 273 apoda, 255, 273 arenicola, 277 arizonica, 255, 273 Coryi, 255, 273 densa, 277 lepidophylla, 255, 273 ludoviciana, 273 mutica, 255, 273 mutica var. limitanea, 255, 273 mutica var. texana, 255, 273 neomexicana, 255, 273 pilifera, 255, 274 pilifera var. Pringlei, 248, 274 Pringlei, 274 Riddellii, 255, 274 rupestris, 277 rupincola, 255, 274 Sheldoni, 255, 274 Sherwoodii, 277 biflora, 31 californica, 31 castellana, 29, 31 coloradoensis, 25, 28 falcata, 17, 29, 31, 51 falcata var. pusilla, 29, 30 falcata var. scabra, 29, 31 hybrida, 17, 19 Juliani, 51 Lindheimeri, 25, 27, 28 Linsecomia, 48, 50 leptocarpa, 48 ovata, 31 pentagonia, 17 perfolrata, 18, 19, 23, 33, 34 perfoliata var. ramosa, 31 perfoliata f. rigida, 31 Speculum-Veneris, 13, 18 Spikemoss family, 273 Spiranthes, 166 Beckti, 172, 173 brevifolia, 173 cernua, 172 gracilis, 172, 173 Grayi, 172, 173 longilabris, 173, 174, 175 montana, 172 ovalis, 172, 175 Parksii, 175, 176 praecox, 175, 177 simplex, 173 tortilis, 174 tuberosa, 173 tuberosa var. Gray, 173 vernalis, 177 tortipila, 277 Streptanthus linearifolius, 207 Underwoodii, 255, 274 Stylodon carneus, 235 Wrightii, 255, 274 Swallen, Jason R., 215 Serinia, 63, 187, 188, 200 Symphyandra, 14 cespitosa, 188, 202 Symphyopappus viscosus, 142 gracilis, 205 oppositifolia, 202 Tectaria, 267 Wrightit, 202 heracleifolia, 255, 267 Shadow-witch, 170 trifoliata, 267 Shinners, Lloyd H., 61, 163, 206, 218 Tetraclea Coulteri, 246 Shiorsh, 158 Coulteri var. angustifolia, 246 INDEX Texas Research Foundation, 281 Texas State Research Foundation, 206 Tillandsia usneoides, 178 Tlacualote texixtl, 4 Tobosa grass, 217 Tracheliopsis, 14 Trachelium, 14 - Tragopogon Dandelion, 190 virginicum, 193, 194 Trichogonia hirtiflora, 143 podocarpa, 143 villosa, 144 Trifolium, 18 Triodallus, 21 rupestris, 34 Triodanis, 13-51 biflora, 25, 31, 33, 34, 37, 39) 49, 41, 46, 51 coloradoensis, 18, 21, 22, 24, 25, 28 faleata, 18, 21, 22, 24, 28 Holzingeri, 24, 41, 45, 46, 47 lamprosperma, 25, 42 leptocarpa, 22, 24, 47,51 perfoliata, 18, 21, 23, 25, 34, 36, 37; 39, 40, 43, 45, 46, 47, 51 rupestris, 23, 34 scabra, 47, 50 texana, 25, 40, 41, 43 Triorchis longilabris, 173 Troximon, 188 Dandelion, 190 virginicum, 193 Tung oil, 7-9 University Herbarium, 8.M.U., 281 Verbenaceae, 220, 222-246 Verbena ambrosifolia, 233 bipinnatifida, 231 bipinnatifida var. latilobata, 232 bonariensis, 222 bracteata, 229 cameronensis, 229 canadensis, 230 canescens var. Roemeriana, 228 ciliata, 234 ciliata var. longidentata, 231, 234 ciliata var. pubera, 234 Cloveri, 227 Cloveri var. lilacina, 227 delticola, 230 2Q1 elegans var. asperata, 230 Halei, 222 hastata, 224 hybrida, 231 Lundelliorum, 229 Macdougalii, 225 Matthesti, 225 neomexicana, 225, 227 neomexicana var. hirtella, 227 neomexicana var. xylopoda, 228 perennis, 228 plicata, 226 plicata var. Degeneri, 227 pumila, 234 quadrangulata, 230 racemosa, 234 rigida, 222 Runyoni, 225, 226, 227 seabra, 224 stricta, 225 tenera, 233 tenuisecta, 232, 233 tumidula, 231 urticifolia, 224 urticifolia var. leiocarpa, 224 Wrightii, 233 xutha, 225 Vitex Agnus-castus, 245 Agnus-castus var. alba, 245 Agnus-castus f. rosea, 246 Wahlenbergia, 14 Weatherby, C. A., 61, 278 Whitehouse, Eula, 163, 206 Wild pear, 148 Wild Rose Pass, 207-213 Woodsia, 267 mexicana, 255, 267 obtusa, 255, 267 Plumerae, 255, 267 Woodwardia, 267 angustifolia, 261 areolata, 261 virginica, 255, 267 Wright, Charles, 61, 163 Zapote, 146 Zuelania, 53 belizensis, 58 Guidonia, 58 Xanthocephalum linearifolium, 121 a i WRIGHTIA EDITOR Cyrus Lonewortsa LunpELL WRIGHTIA, a Sotanical journal, is a publication of the Institute ee anology and Plant Industry of Southern Methodist University. The — tions are by staff members and collaborators. Unsolicited manu- will not be considered. — price is $5.00 per volume. Each volume will contain a series of numbers, to be issued at irregular intervals, each number to vary price according to size. All communications regarding exchanges and et aa CHARLES WRIGHT Dedicated to the Memory of CHARLES WRIGHT 1811-1885 Pioneer Botanical Explorer WRIGHTIA VOLUME I AUGUST, 1945 NUMBER I ly SE ae ee eR ee ee Ee aS ee epee Tee pet Eee ne THE GENUS GARCIA VAHL, A POTENTIAL SOURCE OF SUPERIOR HARD QUICK-DRYING OIL Cyrus LoNGwortH LUNDELL Ee Te ee a ee ee ae ee ee ee INTRODUCTION Among the host of tropical American plants yielding products long utilized by the natives for medicinal purposes is Garcia nutans Vahl. In Mexico the seeds of this tree are locally employed as a purgative. Accord- ing to reports, the purgative action resembles that of castor oil which is obtained from the seeds of Ricinus communis L. In the wartime search for new vegetable oils undertaken by the U. 8. Government, seeds of G. nutans were gathered in Mexico and sent to laboratories in the United States for examination. The first published re- port, issued by Gardner and Westgate’, indicated that the seeds are a potential source of a new superior, hard, quick-drying oil. Shortly after the analyses of the seeds were obtained in 1943, a survey was organized by the Board of Economic Warfare under the direction of the writer to determine the extent of the native stands of the species in Mexico ‘and Central America. Although reports made locally in Mexico City indicated that a large supply of seeds could be readily obtained, these were unfounded. The first native stand of G. nutans was discovered on July 14, 1943, near Pujal in eastern San Luis Potosi. Subsequent surveys extended the range south of Tamazunchale. Throughout the area, the tree grows in abundance on alluvium along streams with the densest stands observed in the vicinity of Huichihuayan. Through survey parties, inquiries, and an examination of records of the species in American herbaria, attempts were made to locate additional native stands. The search has been futile. Although G. nutans is known from widely separated parts of tropical America, and was originally described from South America, an examination of data on specimens and published reports indicate that a high percentage, if not all, of the collections outside of eastern Mexico were made either 1 Gardner, H. A. and M. W. Westgate. National Paint, Varnish and Lacquer Asso- ciation, Inc. Scientific Section, Cire. 662: 173-178. 1943. ne ee ee ee ee eee 2 WRIGHTIA [Vou. 1, No. 1 from cultivated trees or from trees growing in or around inhabited places. Even the type was taken from a solitary evidently introduced tree growing at Santa Marta, Colombia. Since no stands of the tree have been found elsewhere, the evident con- clusion is that G. nutans is native to eastern Mexico, and that the scattered trees found throughout tropical America are the result of introductions and naturalization. Seeds with purgative and other medicinal properties, long used by the natives, are widely distributed by dealers in medicinal plant material (yerberos) in their extensive commerce. Hence the sporadic distribution of the species can be accounted for through this channel. During the survey, Charles L. Gilly, Sr., and Efraim Hernandez Xolo- cotzi of the staff of the Board of Economic Warfare (now the Foreign Economie Administration), discovered an undescribed species of Garcia in northern Tabasco, further evidence that the species of the genus are endemic to Mexico. In this study, the specimens cited are in the herbaria of the Arnold Arboretum of Harvard University, University of Michigan, New York Botanical Garden, and Southern Methodist University. Acknowledgment ‘is made to Dr. A. C. Smith of the Arnold Arboretum for his cooperation in supplying a copy of the original description of G. nutans along with a sketch of the original plate. TAXONOMY OF THE GENUS Garcia Vahl in Rohr, Skrivt. Naturh. Selsk. (Kjoebenhayn) 2: 217, pl. 9. 1792; Benth. & Hook. f., Gen. Pl. 3: 292. 1880. Shrubs or small trees up to 15 m. high, 35 em. in diameter. Leaves alternate, petiolate, penninerved, entire. Flowers monoecious, unisexual, usually with 1 pistillate and several staminate borne at the apex of a twig. Staminate flowers: calyx rupturing irregularly into 2 or more valvate seg- ments, persistent; petals 6-13, narrow, long villous within, appressed hir- sute on the outer surface, pink to deep maroon, longer than calyx; stamens numerous, borne on a pilose, glandular receptacle; filaments free; rudi- mentary ovary absent. Pistillate flowers: calyx as in staminate flowers, caducous; petals as in staminate flowers; disk glandular, deeply lobed; ovary 3-celled; style short, stout; stigma lobed, laminated, fleshy, usually deep maroon; ovules solitary in the cells. Capsules usually deeply 3-lobed, 3-seeded. Seeds globose, without caruncle. The original description of the genus appears in a paper entitled “Plantae-Slaegter beskrevne af Hr. Oberst-Lieutenant von Rohr, med tilfoide Anmaerkninger af Hr. Professor Vahl.’”’ In this paper, most of the generic descriptions (in jarger type) are signed by Rohr, and most of the specific (in smaller type) by Vahl. However, the name ‘‘Vahl” appears at the end of the description of G. nutans, and he must be considered the author of both the genus and the species. v Type species: G. nutans Vahl. | 2) 15° : 1945] LuNDELL: GARCIA 3 Glands in staminate flowers distributed over the entire surface of the re- ceptacle; petals ro-17 mm. long; stamens 63-160; ovary very conspicuous gc as AGS eae Oe ae NG Lae ce ee cee kG 1. G. nutans. Glands in staminate flowers present only in an outer series on the receptacle; petals 5.5. mm. long; stamens 29; ovary minute.......... 2. G. parviflora. 1. Garcia NuTANS Vahl in Rohr, Skrivt. Naturh. Selsk. (Kjoebenhavn) 2: 217, pl. 9. 1792; Pax in Engler, Pflanzenr. IV. 147: 14. 1910; Standl., Contr. U. S. Nat. Herb. 23: 620. 1923. Fig. 1, a-g. Garcia Mayana Britton, Sci. Surv. Porto Rico & Virgin Isl. 6: 357. 1926. An evergreen shrub or small tree, up to 15 m. high, 35 em. in diameter; twigs striate-angled, rather slender, sparsely appressed hispidulous to densely fulvous-tomentulose. Leaves petiolate, the petioles slender, 1—5 cm. long, essentially glabrous to fulvous-tomentulose; leaf blades paler on undersurface, membranaceous to subcoriaceous, oblong, oblong-elliptic, elliptic, oblanceolate-elliptic or obovate-elliptic, 8-19 em. long, rarely smaller, 2.5—7 em. wide, apex abruptly acuminate to obtuse, the acumen acutish to obtuse, usually short, base usually obtuse, sometimes rounded, midvein slightly elevated above, prominent on undersurface, primary veins 7—1o on each side, conspicuous, glabrous on upper surface except for oc- casional scattered hairs along the midvein, varying from densely soft pilose to sparsely subappressed hispidulous on lower surface. Flowers monoecious, usually with 1, sometimes with 2, pistillate flowers borne terminally on a twig, these subtended by several staminate flowers in various stages of development. Staminate flowers borne on slender peduncles 1—3.7 em. long, the peduncles sparsely subappressed hispidulous to densely fulvous-to- _mentulose; calyx 6—13.5 mm. long, opening irregularly, fulvous-tomentulose externally, glabrous within except at apex; petals 6-13, linear, linear- oblong or oblong-elliptic, variable in size in flowers from the same tree, 10-17 mm. long, 2—5.5 mm. wide, apex subulate-acuminate, base attenuate, externally densely hirsute with long straight appressed hairs, the inner surface usually villous with long hairs, but sometimes nearly glabrous, pinkish to dark red; receptacle pilose, bearing between the filaments nu- merous lanceolate to clavate glands up to 1.5 mm. long; stamens numerous, 63-160; filaments glabrous or bearing a few scattered long straggly hairs, minutely granulose, 2.4~6 mm. long; anthers apiculate, o.7—1 mm. long. Pistillate flowers: peduncles 1~3 em. long, tomentulose ; calyx 8-9 mm. long, tomentulose externally; petals linear, oblong-elliptic or elliptic, 11-13 mm. long, 3-6.5 mm. wide, apex abruptly cuspidate, villous within, hirsute on outer surface with long appressed hairs; glandular disk lobed, about 2 mm. high; ovary 3-celled, densely fulvous-tomentulose; style thick, 1.5-2 mm. long; stigma large, laminated, lobed, fleshy, black-purple to deep maroon. Capsules usually 3-lobed, 3-seeded, 2~—2.5 cm. high, 3-4 em. in diameter, fulvous-tomentulose, elastically dehiscent. Seeds globose, up to 1.7 em. in diameter. MEXICO: Sawn Luis Porosf, Finca de Porfirio Castellanos, 4 kilometers east of Pujal, on alluvial soil, July 14, 1943, C. L. Lundell 12241, tree, 6 in, 4 WRIGHTIA [Vou. 1, No. 1 diameter, petals reddish, stigma black-purple; same locality and date, Lundell 12242, tree, 4 in. diameter, petals reddish, stigma black-purple; Huichihuayan, along brook, July 15, 1943, Lundell 12247, small tree, 9-12 ft. high, corolla reddish; same locality and date, on river bank, Lundell 12248, tree, 8 in. di- ameter, 30 ft. high, petals dark red; same locality and date, Lundell 12249, tree, 8 in. diameter, 35 ft. high, petals dark red; between Huichihuayan and Tamazunchale, in forest, July 15, 1943, Lundell 12250, tree, 2 in. diameter, 12 ft. high, petals dark red; south of Tamazunchale, alt. 500 ft., along ravine, Lundell 12255, small tree, petals dark red, stigma maroon; 2 kilometers north of Huichihuayan, alt. 4oo ft., in virgin forest, July 16, 1943, Stephen S. White 5060, slender tree, 8 ft. high; same locality and date, White 5063, tree, 8 in. diameter, 25 ft. high; between Tamazunchale and Valles, kilometer 401, on bank of Axtla River, Aug. 21, 1943, C. L. Lundell & Amelia A. Lundell 12424, tree, 3 in. diameter, 12 ft. high; Huichihuayan, kilometer 409, on river bank, Aug. 21, 1943, Lundell & Lundell 12429, tree, 5 in. diameter, 27 ft. high, petals reddish; Huichihuayan, kilometer 403.5, on river bank, Nov. 5, 1943, Lundell & Lundell 12653, small tree, petals reddish; Huichihuayan, on river bank, Nov. 5, 1943, Lundell & Lundell 12655, small tree, petals reddish; Taman, about 15 kilometers southwest of Tamazunchale, along highway, at base of steep bank of small stream, April 16, 1944, Efraim Hernandez Xolocotzi 164, shrub. Curapas, Escuintla, Dec. 17, 1937, EHizi Matuda 2061; same locality, alt. 160 m., July, 1938, Matuda 2619, tree in plaza, 3 m. high, 15 cm. diameter. Tasasco, Boca del Cerro, near Tenosique, Usumacintla River, riverside, July 1-5, 1939, Matuda 3568, tree, 10 m. high, 35 em. diameter. YucaTAN, Chichen Itza, edge of Dzadz Aguada, June 25, 1932, W. C. Steere 1569, tree. PUERTO RICO: Mayaguez, planted in Agricultural Experiment Station, Mar. 10, 1925, N. L. Britton & Kenneth R. Boynton 8331 (type of G.“Mayana Britton), tree, 12 m. high, flowers pinkish. GUADELOUPE: Jardin d’Essais de Pointe 4 Pitre, July 15, 1936, H. Stehlé 403, introduced and naturalized tree, 15 m. high. TRINIDAD: Botanic Gardens, planted, Nov. 8, 1933, W. E. Broadway 9283, a small but an aged tree, fruits abundantly. COLOMBIA: Dept. Bolivar, vicinity of Turbaco, alt. 200-300 m., in thicket, Nov. 6-22, 1926, EH. P. Killip & A. C. Smith 14327, shrub, 12-15 ft., petals pink, stamens red; Torrecilla, near Turbaco, alt. 150-300 m., on limestone soil in dense woods, Nov. 7-19, 1926, Killip & Smith 14679, tree, 30-40 ft., fruit 3-celled. Vernacular names: ‘“cuatlaguilocuatl,’’ “tlacualote texixtl,’”’ ‘pifion,” “pifioncillo,” “huevo de gato,’ “avellano,” ‘“‘pascualito,” ‘‘pepita del indio.” Recorded by Xolocotzi from Tamazunchale, San Luis Potosf, the first two vernacular names are Indian and these are significant for they indicate endemism of the tree. All of the other names are Spanish, and widely applied to various plants with similar fruits. In eastern San Luis Potosf, the variation in G. nutans is considerable. In pubescence the leaves range from essentially glabrous to densely pilose on the undersurface. As a character of taxonomic significance, flower size is of minor importance, for the extremes in the species, large and small, may be found on a single tree. G. Mayana Britton represents the phase of the species with leaves pilose on the undersurface. In the large series of specimens from the vicinity of 1945] LUNDELL: GARCIA 5 Fig. 1. Garcia nutans Vahl (a-g). a, leaf, X 34; b, fruiting inflorescence, X 34; e, segment of capsule, showing mode of dehiscence, with seed still in place, x 34; d and e, seeds, showing various types of mottling on surface, X 1; f, pistillate flower (note buds of two staminate flowers), 1; g, staminate flower (note that calyx segments and petals have fallen from pistillate flower and that ovary has started to develop), X 1. Garcia parviflora Lundell (h-j). bh, leaf, X 34; i, pistillate flower after calyx seg- ments and petals have dropped, in approximately same stage of development as that in g, X 1; ], staminate flower, < 1. a-e, drawn from Efraim Hernandez Xolocotzi 164; f-g, drawn from C. L. Lundell & Amelia A. Lundell 12429; h-j, drawn from Charles L. Gilly, Sr. & Xolocotzi 297. Mlustrated by Charles L. Gilly, Sr. 6 WRIGHTIA [VoL. 1, No. 1 Huichihuayan, all of the variations found in specimens from the West Indies and South America are evident, and these do not appear to be sufficiently stable to warrant formal varietal recognition. 2. Garcia parviflora Lundell, sp. nov. Fig. 1, h-j. Arbor parva, 2.5-4 m. alta, ramulis gracilibus, novellis adpresse pubescentibus. Folia longe petiolata, membranacea, oblonga, oblongo- elliptica vel oblanceolato-oblonga, 10-16.5 em. longa, 3-5.8 em. lata, apice abrupte cuspidato-acuminata, basi obtusa vel rotundata, sub- glabra. Flores $ pedunculati, 1.3 diam.; calyx 4 mm. longus; petala 6, lineari-oblonga, 5.5 mm. longa, 1.6 mm. lata, subabrupte acuminata, intus villosa, extus adpresse hirsuta; stamina 29; filamenta glabra. Flores @ pedunculati; ovarium 3-loculare, 2.5 mm. longum, hirsutum. Small tree, 2.5-4 m. high, 3-9 em. in diameter; twigs slender, at first densely appressed hispidulous with short curved hairs, drying brown, striate-angled. Leaves with slender petioles 2-5 em. long, the petioles appressed hispidulous at first, the hairs densest at base and apex; leaf blades membranaceous, dark green above, paler on lower surface, oblong, oblong- elliptic or oblanceolate-oblong, 10-16.5 em. long, 3~5.8 em. wide, apex abruptly cuspidate-acuminate, the cusp often obtusish, base obtuse or rounded, midvein slightly elevated on upper surface, prominent on under- surface, primary veins 8—10 on each side, slender but conspicuous, anasto- mosing submarginally, the secondary veins openly reticulate, margin en- tire, slightly wavy, at first sparsely appressed hispidulous with short hairs on midvein above, and along veins on undersurface. Flowers monoecious. Staminate flowers borne on slender appressed hispidulous peduncles 7 mm. long; calyx 4 mm. long, sparsely appressed hispidulous; petals 6, rose-pink, linear-oblong, 5.5 mm. long, 1.6 mm. wide, subabruptly acuminate, short subulate, densely villous on inner surface with long hairs, appressed hairy on the outer surface with long hairs; receptacle pubescent, bearing few small glands in an outer series; stamens 29; filaments stout, 2~-2.5 mm. long, smooth; anthers 2-celled, broadly cordate, about 0.6 mm. long, incon- spicuously apiculate. Pistillate flowers borne on slender appressed his- pidulous peduncles 1.2 em. long; calyx and corolla not seen; ovary hirsute, 3-celled, 2.5 mm. long; stigma lobed, laminated; fruits unknown. MEXICO: Taxpasco, on limestone ridge just north of Cerro de Azufre, about 5 kilometers southwest of Teapa, in rain forest, alt. too-150 m., Sep- tember 20, 1944, Charles L. Gilly, Sr. & Efraim Hernandez X olocotzi 297 (type in the herbarium of Southern Methodist University), small tree, 2.5 m. tall, 3 cm. diameter, staminate flowers 1.3 em. wide; same locality and date, Gilly & Xolocotzi 304, tree 4 m. tall, 9 em. diameter. Although closely resembling G. nutans, the thin membranaceous leaves abruptly cuspidate-acuminate, the slender short peduncles, the small flowers only 1.3 em. in diameter, the rose-pink petals only 5.5; mm. long, the few stamens (29), and the small ovary distinguish the species. In G. nutans glands are present over the surface of the entire receptacle, dis- tributed among the filaments. In G. parviflora glands are present on the ie a i ie 1945] LUNDELL: GARCIA 7 receptacle only in an outer series. The new species evidently represents a localized form in which a reduction in the size and number of the floral, _ parts has taken place. Since no seeds have been available for analyses, nothing is known re- garding the possible economic value of the oil from this species. CHARACTERISTICS AND PHYSICAL PROPERTIES OF GARCIA OIL The genus Garcia is closely related botanically to the genus Aleurites, the source of tung oil. Both are referable to the JATROPHEAR. Although botanical affinity does not necessarily mean that the oil of seeds from species of two genera would have similar properties, the preliminary analyses indicate that Garcia oil and tung oil are much alike in various characteristics and properties. Since the availability of the publication containing the analyses of Garcia oil is limited, the following data have been abstracted from the report by Gardner and Westgate (l.c.). “The first step in the investigation was to shell the nuts and heat-press the kernels to obtain sufficient oil for examination. The shell was very thin and brittle and easily removable. The oil was easily expressed, was very pale, and resembled in appearance American tung oil. The oil was examined for its characteristics and its physical properties. Table 1 CHARACTERISTICS OF GARCIA NUTANS U8. 0. A. THIS CHARACTERISTIC LABORATORY LABORATORY SEED Per cent Kernel 80.52 78.7 Per cent Shell 19.48 a7.3 KERNEL Per cent Oil 53.76 56.7 Per cent Moisture SG fe OIL Refractive Index (25° C.) Hot Pressed Oil from Kernel 1.5260 5254 § Solvent Extracted Oilfrom Pomace ~~ _....... I Solvent Extracted Oilfrom Kernel —~_....... I Momace Carayity (i561 5G5 Oe) ee ee ° Saponification Number 192.4 189.2 Acid Number (Aleohol-Benzol) ee I fe) 7 Unsaponifiable Matter (percent) = # ~__...... .76 Iodine Number (Wijs, 1 hr.) 176.8 177.9 Diene Value (Ellis- Jones) Beet ie. Carbonyl Value (Leith) OD Re en ee Hexabromide Test POs cs 5 8 WRIGHTIA [Vou. 1, No. 1 ‘“‘A study of these characteristics is of considerable interest. In the first place, the nut contains a very high percentage (80%) of kernel. The kernel contains in the neighborhood of 55% oil, approximately that of tung kernel. It will be noted in Table 1 that the refractive index of the oil is very high (1.525)—substantially higher than that of American tung oil (1.517). This is true not only of the hot-pressed oil but of solvent-extracted oil from the pomace or directly from the kernel. The specific gravity is of the same order as that of tung oil. The acidity is very low, approaching that of American tung oil. The unsaponifiable matter is normal for vegetable drying oils, and the iodine value is higher than that of American tung oil. The very high diene value, the negligible carbonyl value, and the negative hexabromide test indicate that the oil is highly unsaturated but that it contains no licanic or linolenic acids. In- dications from the analytical determinations are that the oil might consist almost entirely of elaeostearic glycerides. In Table 1 it will be noted that an analysis of the nuts recently made by the laboratories of the Bureau of Agri- cultural Chemistry and Engineering, Department of Agriculture, Washington, gave results very similar to those obtained at this laboratory. Table 2 PuysicaL TEsts ON EXPRESSED OIL FROM GARCIA NUTANS TEST OBSERVATION Color (Gardner Tubes) 4 Viscosity (Gardner Tubes) Between J and K Heat-Bodying (Browne Apparatus) 282° down to 225° C.in 5 minutes 8 282° down to 175° C. in 21 s x 282° down to 100° C. in 60 ae ¥ Gelatin Time (Browne Heat Test) 7 min., 45 sec. Drying Time Without driers Under 18 hours q With 0.5% Pb, 0.05% Co driers added ipa) geo sx 1 Nature of Film on Glass Frosted and wrinkled “The next step in the investigation was to determine some of the physical properties, and these are presented in Table 2. The purpose of the heat body- ing tests was to determine the bodying effects of heat, which might possibly simulate those of kettle processing. The Browne heat test apparatus was used to secure a set of standard conditions. The oil bath was brought to a tempera- ture of 282° C., the flame was removed, and immediately comparative 5 ml. samples of the Garcia nutans oil and of the American tung oil were placed in test tubes and suspended in the oil bath until a definite lower temperature was reached. The time necessary to secure this temperature difference was noted. The process was repeated for lower temperatures. The test tubes containing the bodied oils were then cooled and the viscosities determined. The viscosities of the bodied oils obtained by this treatment were 8, X, and Y for Garcia nutans oil, as compared to M, U, and V for pure American tung oil. The much greater speed of bodying of the Garcia nutans oil was apparent, thus indicating that this oil, even in the presence of softer oils such as linseed oil, could be used to obtain quick bodying varnish. “Samples of the two oils were heated to 282° C. until gelatinization took 1945] LuNDELL: GARCIA 9 place, and the solid polymerized masses examined. That of Garcia nutans oil seemed to be somewhat firmer than that of tung oil. Also Garcia nutans oil polymerized to a solid in less than 8 minutes as compared to 12 minutes for American tung oil. “Varnishes were then prepared with the Garcia nutans oil, using a modified phenolic resin. The resin and the oil were heated quickly together for about 10 minutes to a temperature of 250° C., and held at that temperature for an- other ro to 15 minutes. Good string and body were obtained, and the varnishes were thinned to about 60% non-volatile. It was found, however, that better results were obtained when the varnish was thinned to a 50%- non-volatile content. The thinner varnish dried within a period of one to two hours, with the production of clear films. After aging overnight, a flowed-out film of this varnish could be immersed in boiling water for one hour, followed by cold water for 150 hours, without showing any blushing effects. “As a matter of general interest, there was included in this investigation of physical properties the effects of different chemicals upon the oil. For in- stance, it is well known that tung oil may be gelatinized almost immediately by the addition of stannic chloride in the cold. It was found that Garcia nutans oil gave almost the same reaction, resulting in possibly an even firmer mass. “Samples of Garcia nutans and of American tung oils were treated with a solution of a few drops of iodine dissolved in mineral spirits. These were placed in front of an ultraviolet lamp for a few minutes, and then allowed to stand in the laboratory for a few hours. Both samples were changed to a solid white opaque mass, indicating the formation of beta elaeostearin. “The rates of saponification of these two oils in alcoholic potassium hy- droxide were very different. It took at least five times as long to saponify completely the Garcia nutans oil as it did the American tung oil. This would suggest its use for varnishes of high alkali resistance. “The appearance, size and shape of the Garcia nutans nuts resembled very closely that of Japanese tung oil nuts which were investigated by the Scientific Section in November, 1921 (see Scientific Circular 138). It would appear, how- ever, that the oil from the nut of Garcia nutans is far superior to that from the nut of the Japanese tung oil tree (Aleurites cordata). As a matter of fact, it appears to be even much superior to oil from the nut of the tung oil tree (Aleurites Fordit). “A comparison of the properties of Japanese tung oil, as reported upon in Scientific Section Circular 138 referred to above, with those of Garcia nutans oil, is of interest. For instance, the Japanese tung oil has an iodine value of about 150 and a refractive index of only 1.4891. It is a much softer oil than Chinese tung oil and can be heat-treated for a longer period of time without the formation of as viscous a product and without any great danger of gelatini- zation. In other words, Japanese tung oil is much less satisfactory than Chinese tung oil as a varnish making material. These data are included here to point out the differences between Japanese tung oil and Garcia nutans oil, because at first glance the appearance of the seed of the latter product seemed much like that of Japanese tung oil seed. “In conclusion, it would appear that the seed of Garcia nutans should re- ceive immediate consideration by agricultural experts to determine whether plantings, made under proper climatic conditions, could develop quickly an available source of this oil which appears to be very valuable for the manu- facture of protective coatings.” 10 WRIGHTIA [Vou. 1, No. 1 A further report on subsequent studies of the oil from seeds of Garcia nutans has been made by Westgate? in which the following additional data are presented. “The investigation of this oil, as reported in Scientific Section Circular 662, has been supplemented with additional tests of a control nature. It was found that the oil in thin films dried on glass somewhat more rapidly and more firmly than American tung oil treated in a similar fashion, with or without driers. Comparative varnishes were made with these two oils, and it was again found that Garcia nutans oil conferred much more body in the same cooking time than did tung oil. This was true even when 50% of either of these oils in a varnish was replaced by heavy bodied soybean oil. It would appear, therefore, that on a laboratory basis the preliminary optimistic report to the varnish industry on the quality of Garcia nutans oil, which was contained in Scientific Circular 662, has been amply confirmed.’ POSSIBILITIES FOR COMMERCIALLY EXPLOITING GARCIA NUTANS The only known wild stands of G. nutans occur in eastern San Luis Potosi in the region extending from Valles to Tamazunchale. The ex- tension of the stands from Tamazunchale into Veracruz appears probable, for the species has been collected in the northern part of that state. G, nutans in this sector occurs principally on heavy alluvial soils along streams and dry stream beds at an altitude of less than 300 meters above sea level. All of the area in which the species has been found is covered with secondary vegetation, for the virgin forest has been felled from time to time for agricultural use. Where the forest has been cleared recently and abandoned, the species forms local thickets. Trees up to 20 centimeters in diameter and 12 meters high were found along the river near Huichihuayan. The Huastecan Indians report that the tree reaches a diameter of 45 centimeters in old forest. Along the river at Huichihuayan, seedlings and stump sprouts are abundant. Some of the seedlings, not over three years old, were in flower in August, 1943. Since the widespread clearing of the alluvial lands for agriculture has decimated the wild stand of G. nutans, the available crop of seeds for com- mercial purposes is very limited. A liberal estimate is that not more than 10 long tons of seeds can be obtained annually from the native stands in eastern San Luis Potosf. The seeds which can be gathered from San Luis Potos{ should be ade- quate to permit experimental tests of the oil on a pilot plant basis. Until the results of initial laboratory experiments have been confirmed by adequate commercial tests, no large scale plantings of the tree can be recommended. In order to obtain an estimate of the potential yield of seeds, a count was made of all the young fruits on a mature tree in the riparian forest at Huichihuayan. Since the dry weather in the summer of 1943 had caused * Westgate, M. W. National Paint, Varnish and Lacquer Association, Inc. Scientific Section, Cire. 672: 129-132. 1944. 1945] LUNDELL: GARCIA II the shedding of many flowers, a count was also made of all twigs which would bear fruits in a favorable season. Since the inflorescence of G. nutans consists of one, rarely two, pistillate flowers subtended by several staminate flowers, usually only one capsule with three seeds is to be found on a twig. The tree selected measured 12.5 centimeters in diameter and 9 meters tall. On the basis of the annual rings, a doubtful method for determining age in the tropics, the tree appeared to be approximately 25 years old. There was a total of 1762 twigs capable of bearing fruits. Due to the dry summer in 1943, only 203 twigs bore young capsules in August from the July flowering period. Typical of the reaction of many plants of semi-arid regions, flowering is determined to a high degree by moisture conditions. In the fall of 1943, all of the Garcia trees in the vicinity of the one felled flowered again; hence, the count of 203 for the tree is not indicative of its potential yield that year, for the set of fruits from the second flowering was considerable on other trees. In plantation culture, the spacing of trees 3 meters apart would give a stand of about 441 trees per acre. On the basis of the twig count from the mature tree growing under forest conditions at Huichihuayan, with all twigs bearing capsules, the potential maximum production would be 2697 kilograms (5946 pounds) of seeds per acre in favorable years. This calculation is based upon the potential yield of only a single wild tree under competitive forest conditions. The yield from cultivated trees should be higher. The fruits mature in April and May at which time the seeds are dis- tributed by the elastically dehiscent capsules. Under plantation culture, trees should begin to yield a limited harvest within three years. As recorded on herbarium specimens of G. nutans, the species has been planted in Puerto Rico, Guadeloupe, and Trinidad in experimental gardens. In all places it has borne fruits, and Broadway makes the following nota- tion on the label of his collection from Trinidad: ‘‘fruits abundantly.”’ The species appears to be adapted to cultivation over a large part of the low- lands of tropical America. From seeds obtained in Mexico by H. A. Gardner and sent to the Sub- Tropical Experiment Station at Homestead, Florida, test plantings were made in Florida in 1943 by 8S. J. Lynch (Westgate l.c.). Seed germination was better than 95 per cent, producing vigorous rapidly-growing seedlings. These seedlings have been set out in Florida and in the Lower Rio Grande Valley of Texas. From seeds obtained in San Luis Potosi in 1943, a test planting was made at the Agricultural Research Station of the Institute of Technology and Plant Industry at Dallas on March 17, 1944. At the same time, other seeds were planted in the greenhouse on the campus of Southern Methodist University. Although the germination was excellent, as in the Florida planting, the cotyledons were not able to throw off the hard seed coat under the dry open field conditions, resulting in a stand of only 20 per cent. Since the species is a forest tree where seeds germinate in moist conditions, 12 WRIGHTIA [Vou. 1, No. 1 growers must avoid open field plantings unless adequate shade and water can be supplied. G. nutans is not hardy, for the frost of November 27, 1944, at Dallas, when the temperature dropped to 29° F., killed all of the plants not pro- tected. Under greenhouse conditions, the seedlings thrive. From seeds planted in the greenhouse at Dallas in March, 1944, seedlings had reached a height of over 1 meter on May 17, 1945. One of the seedlings, 14 months old, flowered in May, 1945. Since the species is not hardy as far north as Dallas, test plantings are being made along the Gulf Coast of Texas. The seedlings in the plot at McAllen, Texas, have made excellent growth showing no setback during the mild winter. If commercial planting of G. nutans prove to be feasible, the sector be- tween Mante and Tamazunchale in eastern Mexico, where ample river water is available for irrigation, appears to be favorable for plantations. Puerto Rico, Cuba, and other similar tropical areas should prove equally well adapted to its culture. SUMMARY 1. From preliminary investigations, G. nutans appears to be endemic in eastern Mexico, for the only known wild stands of the species occur in the region between Valles and Tamazunchale, San Luis Potosf. 2. The genus Garcia, closely related botanically to the Asiatic genus Aleurites, the source of tung oil, contains two species native to Mexico, G. nutans and G. parviflora. Only G. nutans is at present known to bear seeds of potential commercial value as a source of quick-drying oil. 3. Initial laboratory tests indicate that Garcia oil has characteristics and physical properties similar to those of tung oil. In some ways, it ap- pears to be equal, if not superior, to the best tung oil of commerce. 4. Pilot plant tests are necessary to demonstrate that Garcia oil is as valuable commercially as laboratory results indicate. 5. The known wild stands of G. nutans are estimated to have a potential production of not more than ro long tons of seeds annually. 6. The cultivation of the tree on a plantation basis will be necessary to obtain substantial commercial quantities of the oil. Initial estimates, based upon a study of wild trees under competitive forest conditions, indicate a low yield. However, through cultivation it should be possible to substanti- ally increase the per acre production of seeds. 7. In test plantings in Florida and Texas, seedlings have made vigorous growth, some flowering within 14 months. 8. The tree is widely adapted to the lowlands of tropical America. The area in eastern Mexico between Mante and Tamazunchale is suggested as the most favorable locality for growers to make test plantings on a com- mercial scale. THE GENUS TRIODANIS RAFINESQUE, AND ITS RELATIONSHIPS TO SPECULARIA AND CAMPANULA Rocers McVaucu INTRODUCTION The genus Specularia (CAMPANULACEAE) has been more or less gen- erally accepted by botanists only since the publication of Alphonse De- Candolle’s Monographie des Campanulées in 1830, although the name was proposed in 1748 by Heister for the European plant which, as DeCandolle said, ‘“‘vulgo in omnibus linguis et omni tempore Speculum Veneris dicta fuit.”” Although four American and at least four European species are currently referred to Specularia, no critical evaluation of the generic or specific limits in the group has been published since the work of De- Candolle. Study of American material shows that at least 3 species remain undescribed, and since various workers have cast doubt upon the generic standing of the group as distinct from Campanula, it has seemed best to examine all the supposed species of Specularia before proposing additions to a genus of somewhat doubtful standing. The relationship between Specularia Speculum-Veneris (L.) A.DC. (the type species of the genus) and most of the other species is not a particularly close one, and apparently the biological status of all the species concerned may best be expressed by the grouping of all the American species and at least one European into another genus, T'riodanis Raf. Specularia as properly applied and restricted then comprises but 2 species, both European. At least a part of Specularia as currently interpreted is probably best returned to Campanula. These conclusions are set forth after examination of the generic lines in the entire CAMPANULINAE, the subtribe to which the above genera belong. The CAMPANULACEAE proper (CAMPANULACEAE~CAMPANULOIDEAE) are traditionally and apparently justifiably divided into two principal groups of genera distinguished by the manner of dehiscence of the fruit. In the first of these, CAMPANULINAE, the fruit is a capsule opening by lateral pores, and there is a clear distinction between these plants and the WAHLENBERGINAE, in which the capsule is apically dehiscent. Within the CAMPANULINAE, however, generic lines are exceedingly difficult to draw; 7~10 genera have been recognized by most conservative workers, but some of these owe their continued existence perhaps rather to tradition than to biological discontinuity. Most of the several hundred species of the sub- tribe are customarily assigned to the genus Campanula, which is thus made to include a very great variety of forms; this inclusive genus may be regarded as a diverse assemblage of species from which no one has been 13 14 WRIGHTIA [VoL. 1, No. 1 able to segregate any additional plausible genera, or the traditional genera may be thought of as small groups split off, with varying degrees of justi- fication, from the great Campanula complex. This was recognized by De- Candolle (1830, p. 43) and by Buser (1894, p. 508). Similar phenomena are familiar to students of all groups of plants involving large genera. Both Specularia and Triodanis may be regarded as segregate genera removed from Campanula. As the European species are still referred to Campanula in some recent works! and as the maintenance of T'riodanis, distinct from both Specularia and Campanula, may be questioned by some workers, it is thought desirable to develop the argument for this position in some detail. Many supposed genera have been removed from large and natural but not entirely homogeneous groups in a semi-mechanical way. Segregate genera of this kind are usually smaller than the inclusive groups from which they are taken, since the species often have in common not more than a single, arbitrarily chosen, distinguishing character. Segregation of such genera is not necessarily unwise, but the criteria must be well chosen? lest one recognize as many genera as species! It may be sound practice formally to delimit groups of species founded upon single or few characters, but the wisdom of the maintenance of these as independent genera is another matter, for at present we have no test, except that of individual and col- lective botanical opinion, by which generic limits may be determined. Ideally it may be supposed that genera constitute natural (that is, bio- logical) units; in practice they often become conveniences, proposed per- haps because of exaggerated importance attached to some striking charac- ter, and maintained because of tradition and inertia. Linnaeus (1753, 1754), in what we now call CAMPANULINAR, recognized Campanula and two additional genera, Phytewma and Trachelium. Al- though the present concept of Campanula differs from that held by Lin- naeus, chiefly because of the removal of various discordant elements imperfectly known to him, and now placed in Wahlenbergia and other distantly related groups, it seems that his ideas of classification (mostly borrowed from earlier authors!) have hardly been improved upon. Cam- panula, Phyteuma, and Trachelium remain the strongest genera, with Adenophora almost equally strong and Asyneuma but little less so. The remaining genera, without exception, are either monotypic, and accord- ingly unique or not according to the opinions of individual systematists (Azorina, Campanulastrum, Campylocera, Cylindrocarpa, Favratia, Hetero- codon, Ostrowskia, Petromarula, Rotantha, Sicyocodon and some 20 others) or are small (up to about 6 species) and based upon characters of question- able importance (Diosphaera, Michauxia, Specularia, Symphyandra, Tracheliopsis, Triodanis). ‘cf. Bentham, George, Handbook of the British Flora, ed. 7, 287. 1924. ? In Rafinesque’s Flora Telluriana (1; 81), in his “Fifty rules of generic nomenclature by Linneus and Rafinesque” is a sound bit of wisdom: “3. None but skilful Botanists ought to form and name Genera.” 1945] McVaucu: TrIoDANIS 5 In an attempt to prepare a formal statement of generic criteria in the CAMPANULINAE, the following recommendations have been drawn up. It is thought that the separation of Triedanis from Specularia and Cam- panula is justified according to these recommendations. Although the latter have not been applied to any groups of species except among the CAMPANULINAE, it should be possible to make use of them in other families. Most of the suggested procedures are fairly obvious, but lam not aware of any previous attempt to evaluate and integrate them. RECOMMENDATION 1. Primary consideration should be given by the systematist to strong morphological characters (that is, ordinarily, qualita- tive characters, or those involving changes in the nature of plant-parts, or the presence or absence of some distinctive attribute) rather than to the weaker characters (those involving changes in number, shape, position or attachment of parts). Unfortunately for the systematist, when there is a question of generic distinctness there are usually no strong morphological characters involved; determination of obscure generic lines often must depend in approximately equal parts upon: 1), tradition resulting from the combined forces of inertia and botanical opinion; and 2), evaluation of weak morphological characters that appear to indicate some biological discontinuity. Occasionally a “strong” character is not at the same time an obvious one; here recognition of actual generic limits may be delayed. In Adeno- phora, for example, all the species have a distinctive annular gland about the base of the style; this feature is pot present in any other species- group of the Campanula complex, and the standing of the genus has not been challenged. The numerous species are mostly Asiatic, however, and the gland is inconspicuously located, so that the genus was not recognized as such until after 1820, although many of the species were known to earlier workers. RECOMMENDATION 2. The erection, maintenance, or submergence of any segregate genus upon the basis of characters of minor import, or upon single characters, should be advocated only after full consideration of usage, which is the historical expression of botanical opinion. — The recognition of the power of tradition in determining generic lines may seem biologically unsound to some workers, but many genera that are not particularly distinct biologically are currently maintained because of tradition; this is often a satisfactory solution to a problem since there is no final or conclusive test for what is or is not a “good” genus. Except for the relatively rare cases involving conclusive morphological evidence, usage should be considered as of primary importance in determining the limits of “borderline” genera. RECOMMENDATION 3. The most important criterion of any supposed genus is not the width of the gap between it and another, but its own biological unity. Homogeneity in many characters, regardless of the degree of overlapping of these characters with those of other genera, 1s the best 16 WRIGHTIA [Vou. 1, No. 1 indicator of this unity. When homogeneity has been achieved, a segregate genus should be expected to form a group, any species of which will differ from those of other genera by some combination of perhaps insignificant characters, all or most of which may appear individually or in other com- binations in the other genera. Genera of the nature of satellites, if also comprising two or more series of species, invite suspicion especially if united by a single character which is also the only character separating them from a more inclusive genus. Even weaker is the segregate genus which exhibits among its species varia- tions parallel to those found in the related but more inclusive genus. RECOMMENDATION 4. Those proposing or advocating changes in status involving segregate genera should be guided by the diagnostic features of the more inclusive genera as they occur throughout the complete range of the group, and not by more or less narrow, regional similarities or dis- similarities. While those writing local floristic works must of necessity be slow in accepting proposed changes in nomenclature in order to avoid in- stability in popular usage, systematists should be wary of the taint of provincialism. RECOMMENDATION 5. The systematist must make an independent de- cision in regard to the status of each proposed genus, basing his decision upon the relationships between the proposed segregate and some more inclusive genus, but not upon the supposed equality between it and some other segregate. It is impracticable to attach relative degrees of importance to characters used in determining generic limits, or to combinations of these characters, so that it is clearly impossible to say that any two or more genera are equally well founded. Generic segregation, when advocated for the sake of supposed consistency within some major plant-group, leads to arbitrary ~ rather than natural delimitation of genera. RECOMMENDATION 6. Any segregate genus should be sharply de- limited; that is, any species which is intermediate in one or more respects toward a more inclusive genus should be relegated to the latter. The re- tention of the anomalous species in the more inclusive genus will change its limits, if at all, but very slightly, and only in this way can the segregate genus be precisely defined. RECOMMENDATION 7. While many plant-genera are segregated on the strength of single characters, it is clear that the security of the position of any genus increases rapidly in proportion to the number of differentiating characters that can be adduced, particularly if more than one species be involved. The relative importance of any single character varies even among genera of the same family, and is ultimately determined in any case by individual judgment. RECOMMENDATION 8. The generic position of any segregate group, especially if it comprises two-or more species, will probably be strengthened 1945] McVauau: TRIopANIS 17 if the group have a distinctive geographical range together with more or less distinctive morphological features. Workers may find that species which show anomalies in their geographical distribution also possess un- suspected structural peculiarities. GENERIC STATUS OF SPECULARIA When the criteria embodied in the above recommendations are applied to the species currently referred to Specularia, it appears that the genus in its traditional circumscription cannot logically be maintained apart from Campanula. It is not sharply delimited or homogeneous; the traditional features of the type species (a “rotate” corolla and a much elongated capsule) disappear entirely or partly in all the other species, and in addition the genus comprises two well-marked series of species, one having cleistog- amous flowers and a spicate inflorescence and the other having normally open flowers and an expanded inflorescence. The corolla- and capsule- characters separating Specularia from Campanula are relatively weak ones, hardly sufficient in themselves to justify a generic segregation, even if they were of universal occurrence in the species of the segregate and wanting entirely in the other genus. As some undoubted species of Cam- panula (including annual species like C. ramosissima Sibth. & Sm., among the Campanulas which are presumably most closely related to the inclusive Specularia) have a rotate corolla, and most of the American species of Specularia have a short capsule not essentially different from that of many Campanulas, it is evident that these characters do not establish the genus very firmly, and do not necessarily indicate any considerable degree of biological affinity among the species. It appears, however, that the inclusive Specularia may be broken up into relatively homogeneous biological units by the following steps: 1), re- striction of Specularia as a genus to include not more than the type species and perhaps also S. hybrida; 2), removal of all American species and also the European S. falcata to Triodanis; and 3), return of S. pentagonia to Campanula. The two remaining species of Specularia as thus restricted are both natives of western and southern Europe, with deeply divided open corollas, long capsules constricted beneath the calyx, glabrous and gradually dilated filaments, and branched inflorescences. The desirability of recognizing Specularia as a genus is still open to question, but the combination of divided corolla, elongated capsule and glabrous filaments is unique among the annual species of the Campanula complex, and these similarities may indicate some generic affinity between the two entities involved, although these are superficially dissimilar in aspect, degree of branching, and size and shape of flowers. Feer (1890, p. 609) went so far as to suggest that not only these species but also Campanula Erinus L., and C. fastigiata A.DC., which also have glabrous filaments, should be removed from Campanula on this account. Campanula pentagonia L., in spite of its elongate capsule and the almost 18 WRIGHTIA [Vou. 1, No. 1 spike-like inflorescence of some plants (the latter superficially very like Triodanis coloradoensis), I should not refer to Triodanis or to Specularia. It has the lobed (not deeply parted) corolla and dilated ciliate filaments of most Campanulas, and does not produce cleistogamous flowers. Apparently it has affinities with T'riodanis falcata (Ten.) MeVaugh and perhaps with T. coloradoensis (Buckl.) MeVaugh but to refer it to Triodanis would be to break down completely the already meager distinctions between that genus and Campanula. The remaining species of “‘Specularia’”’ form a group which seems to constitute a genus on the basis of the recommendations given above, ex- cept that it has not been traditionally accepted. This is Triodanis Raf. Its homogeneity is unquestioned; the morphological features which dis- tinguish it appear to be relatively strong ones in the Campanulaceae. The combination of annual habit, deeply divided corolla, capsule longer than wide and opening near the apex, spicate inflorescence, and regular produc- tion of cleistogamous flowers at the lower nodes is a unique one, and when found in a relatively large number of species surely points to a genetic dis- continuity that may be called a generic distinction. The geographical dis- tribution of the genus is not an unnatural one, the affinity between the flora of western arid North America and that of the Mediterranean region having been pointed out with respect to other genera (ef. Centaurium, Trifolium, ete.). The cleistogamous condition in Triodanis is not peculiar to this group, but seems to be of importance there because of its universal occurrence in all the species, because of its correlation with the spikelike inflorescence and other characters, and because of the dominance it seems to exhibit in many species over the condition normal for most genera, chasmogamy or the production of open flowers. Cleistogamy, with its accompanying reduction in size and number of parts of the flower, was early noted in the American species of Triodanis, particularly in 7. perfoliata, in which it was seen by Linnaeus and his stu- dents (1748, 1756, 1792). Many subsequent European authors have con- trasted this species of ““Specularia’’ with the European species in which cleistogamy does not occur, but few have considered the character as of more than passing taxonomic interest. Apparently Specularia Speculum-Veneris does not produce cleistog- amous flowers, unless in exceptional cases. I have found but one reference to such a condition in this species, and that a questionable one. Hegi (Il. Fl. Mit.-Eur. 6(1):367] says: “Bei dieser und verwandten Arten (besonders bei S. perfoliata DC.) sind von Kirchner und Helene Ritzerow kleistogame Bliiten . .. nachgewiesen worden,” but the reference to “dieser” [i.e. to S. Speculum-Veneris| seems to be in error. I ean find no such reference in the works of Kirchner, and Ritzerow (1907) discussed the morphology of Specularia perfoliata but not that of any other species. Darwin (1895) discussed the results of crossing and those of selfing in Specularia Speculum-Veneris. He made no mention of cleistogamy in this 1945] McVauau: TRIODANIS 19 species, but mentioned that of S. perfoliata, as if by contrast, in a footnote. Von Mohl (1863) wrote on the morphology of S. perfoliata without any mention of cleistogamy in any European species. Von Mohl’s work was adapted without any significant additions by Knuth (1898). Ascherson, in two papers on cleistogamy (?1880, ?1889), alluded to the phenomenon in various genera and species, including the Egyptian Campanula dimor- phantha, but not in the European Specularias with which he was pre- sumably familiar. Imperfect or abortive flowers in Specularia hybrida have been reported by various writers, including Linnaeus (1756), DeCandolle (1830) and Bromfield (1849), but Asa Gray (1876) denied the existence of such a condition in this species. Like Gray, “I have not found any trace’’ in Specularia hybrida of the extremely reduced flowers, with vestigial corollas, stamens, and style, which are universally present in Troidanis. Some of the flowers of S. hybrida, however, although apparently open and functional, are considerably reduced in size and apparently represent a step in the direction of truly cleistogamous flowers. I have not been able to determine the sequence, if any, in which these flowers are produced, but they seem not to be formed as a part of the regular flowering succession as in 7'riodanis, and presumably the two conditions have arisen independently. Cleistogamy in the genus Campanula is rare but has been reported not only in C. dimorphantha (mentioned above) but also in 2 Indian species, C. canescens and C. colorata; the condition in these latter was noticed by DeCandolle and was elaborated upon by Hooker and Thomson (1858) and by Oliver (1862). The morphological changes induced by cleistogamy in the individual flowers of these Indian species are closely parallel to those in the American species of Triodanis, but the differences in general mor- phology between the two groups seem to preclude the possibility that these are closely related otherwise. Another genus in which cleistogamy is of regular occurrence is the monotypic Heterocodon, of western United States and British Columbia. It appears that I was overly hasty in transferring H. rarzflorum Nutt. to Specularia (MeVaugh, 1941), for in spite of a superficial resemblance to the American species of Triodanis, it seems to have little in common with these. The general morphology of Heterocodon suggests that it has been derived rather from some ancestor more like the conventionally accepted species of Campanula; tradition, indeed, is apparently the strongest reason for keeping it out of Campanula at the present time. The flowers are usually said to be axillary but are actually terminal, and displaced laterally by the axillary branches which successively constitute the ‘‘main axis’ of the plant. The corolla is tubular as in most Campanulas, and the very short and broad capsule is apparently closely similar to those of Campanulas with basally dehiscent capsules, rather than to those of Triodanis, which funda- mentally are apically dehiscent. The placentae are very short and stalked, like those of many Campanulas with similar capsules, not elongate and sessile like those of Triodanis. 20 WRIGHTIA [Von. 1, No.1 The capsule of Heterocodon was originally described by Nuttall (1843) as “dehiscing irregularly at the base’’; Gray (1878) said “Capsule bursting indefinitely on the sides by the giving way of the thin walls,” as opposed to the condition in Campanula and Specularia, in which the capsule was said to be ‘dehiscent by one or more small valvular openings on the sides, usually over a partition.’’ Authors of floristic works including Heterocodon have without exception described the capsule as dehiscing irregularly; actually the mechanism is identical in principle with that prevailing in Campanula and related genera; an elastic portion curls outward and breaks out of the capsule, leaving an opening into two adjacent locules. In Hetero- codon the margins of the opening are less clearly defined than in Triodanis, but the opening is similar to the type found in several species of Cam- panula with basally dehiscent capsules. The pertinent characters of the genera discussed above are set forth in the following synopsis. It will be noted that the generic characters of the groups segregated from Campanula are all weak ones, in no case involving strong morphological features which absolutely separate them from the more inclusive genus, but always depending upon combinations of weaker characters which reappear individually in unrelated species of Campanula. Some workers may consider it a sound biological policy to submerge all of the segregates in Campanula. Of the three in question, 7'’riodanis appears to be the strongest, comprising as it does 8 closely similar species. There is thus some morphological justification for it, but far less for Specularia and Heterocodon. 1. Campanula L. Plants perennial, biennial or annual, if annual usually dichotomously branched, rarely with spiciform inflorescence (cf. C. americana L., C. pentagonia L.) and never with both spiciform inflores- cence and the lower flowers mostly cleistogamous; flowers normally all open, with expanded campanulate corolla, but occasionally partly cleistog- amous (C. canescens, C. colorata, C. dimorphantha), if cleistogamous then pedunculate; open corollas usually not divided as far as the middle, if deeply parted in annual species then the ovary never greatly elongate and linear (ef. C. ramosissima); filaments usually abruptly dilated and ciliate at base, if gradually dilated and glabrous then the capsule short and broad and the corolla not deeply parted (ef. C. erinus, C. fastigiata) ; capsule various, opening at base, near middle or near apex (at base in annual species with cleistogamous flowers and short capsules, near apex in C. pentagonia, which has a greatly elongated capsule). A large and heterogeneous genus, widely distributed in the Northern Hemisphere. 2. Heterocodon Nutt. Plants annual, with short-pedunculate falsely lateral flowers, the earlier ones cleistogamous; open corollas not divided as far as the middle; filaments ciliate at base; capsules short and broad, open- ing near base. One species, western North America. 3. Specularia Heist. ex. Fabr. Plants annual, branching above the middle, the flowers nearly sessile, clustered near the tips of the branches or 1945] McVauaGu: TrIoDANIS 21 corymbosely aggregated at the summit of the plant; flowers all open, or some imperfectly developed but open and not vestigial; corollas divided well below the middle, more or less rotate; filaments gradually widened to base, glabrous; capsule much elongate, linear, contracted at apex beneath the calyx-lobes, dehiscent at apex. Two species, western and southern Europe. 4. Triodanis Raf. Plants annual, the branches, if any, from base or middle of the plant; flowers axillary, sessile or essentially so, the inflorescence spiciform; flowers from the lower nodes normally cleistogamous, with the corolla and androecium vestigial; some of the upper flowers or at least the terminal one usually open (all corollas sometimes open in T. coloradoensis), with expanded corollas divided below the middle; filaments abruptly dilated and ciliate at base; capsule ovoid or clavate to linear or subulate, usually not abruptly contracted at apex, opening at the apex or (in 7’. perfoliata) at the middle or a little above it. Eight species, one chiefly Mediterranean, the others American. SYSTEMATIC TREATMENT TRIODANIS Raf., N. Fl. N. Am. 4: 67. 1837. Specularia, sect. Dysmicodon Endl., Gen. 518. 1838. Dysmicodon Nutt., Trans. Am. Phil. Soc. n. s. 8: 255. 1843. Triodallus Raf. ex Nutt., Trans. Am. Phil. Soc. n. s. 8: 255, in syn. 1843. Campylocera Nutt., Trans. Am. Phil. Soc. n. s. 8: 257. 1843- Specularia sect. Triodallus Raf. ex Torr., Fl. N. Y. 1: 428. 1843. Specularia sect. Campylocera (Nutt.) A. Gray, Proc. Am. Acad. 11: 82. 1876. Specularia American authors, not Heist. Plants annual, erect or reclining, simple or with ascending subordinate lateral branches (these except in 7. coloradoensis chiefly from the nodes near the base of the plant, sometimes 10-12 in vigorous plants and almost equaling the main axis, the plant then with a rosette-like cluster of erect wand-like stems). Stems green, or at base yellowish to reddish- or purplish- brown, with about s subalate angles, these continuous with the decurrent leaf-bases; stems, at least the basal portion, vestite on the angles but only sparingly or not at all so between them. Plant consisting of a single straight axis having up to about roo nodes, of which the lowest IO~15 OF 25 produce foliage (sterile) leaves and the remainder produce sessile bracts and axillary sessile flowers (or in T. coloradoensis some axillary flowering branches). Leaves and bracts glabrous above (except in 7. coloradoensis and T. falcata) or the lower leaves with a few hairs near the tip; lower surface of the leaves usually vestite like the base of the stem (pilose to hispidulous or scabrous), at least on the principal veins near the base of the blade; bracts often smooth and glabrous on both surfaces. Inflorescence spiciform, often dense and the bracts more or less imbricate, the flowers of the main axis and subordinate axes, if any, developing in succession from base to- ward apex except for the flower terminating the axis, which develops be- 22 WRIGHTIA [Vou. 1, No. 1 fore those immediately below it; flowers in each axil 1-3 (rarely as many as 7-8), the axillary flowers of the spike terminating repressed lateral branches which under sufficient stimulus elongate slightly and produce 1 or 2 lateral flowers from this secondary axis or even 1~4 additional later flowers from these tertiary axes, all the flowers essentially sessile (in 7’. coloradoensis some of the branches regularly elongate and produce each 1-3 flowers near the tip); flowers at the lower fertile nodes regularly cleistogamous; flowers at some of the upper nodes, at ieast the primary ones terminating the axes, producing an open (chasmogamous) corolla; axillary axes, at least when 2- or more-flowered, bracteolate, the bracteoles minute or foliaceous, at the base of the primary flower, usually narrow, up to 3-4 mm. long or sometimes larger. Calyx-lobes green, sometimes much ex- panded and foliaceous, entire except usually for one callose-glandular tooth on each edge near base, the tooth occasionally enlarged and the calyx-lobe then serrate; lobes normally 5 and all alike in the open flowers, usually either 3 or 5, less often 4 in the cleistogamous flowers (except in T. falcata and T. coloradoensis), those of the eleistogamous flowers smaller, often relatively broader, 1 or 2 often smaller than the others. Corolla of the open flowers usually lavender-blue, glabrous or with a few bristles without near the tips of the lobes, with short narrowly funnel-form tube which is mostly less than one-third as long as the whole corolla, and 5 elliptic and abruptly acuminately pointed lobes which are mostly about twice as long as wide. Stamens normally 5s, the filaments flattened, divided about equally into a narrow linear glabrous distal portion and a gradually dilated or abruptly rounded broad ciliate base; anthers linear, glabrous, longer than the filaments. Style pubescent distally half its length or more, and divided at tip into short branches equal in number to the locules of the ovary (that is, usually 3 except in 7’. leptocarpa). Cleistogamous flowers smaller than the open ones, the calyx and capsule somewhat smaller, the corolla, stamens and style reduced to minute rudiments. Capsules usually erect or ascending, linear, oblong or ellipsoid to clavate (subulate in the cleistogamous flowers of 7’. leptocarpa), at maturity tipped by the calyx- lobes, those of the cleistogamous flowers usually more nearly terete (rather than angled or suleate), tending to be ellipsoid rather than oblong or clavate as in the open flowers, and bi- or trilocular (unilocular in 7’. lepto- carpa); pores of the capsule equal in number to the locules (except some- times in 7’. leptocarpa), usually distal, opening from base toward apex (except in 7’. coloradoensis), the opening brought about by the curling out- ward of an indurated cartilaginous process which is apparently derived from a portion of the ovary (or hypanthium) wall and a portion of the septum; this process breaks cleanly outward at one end, carrying with it rather regularly defined and cleanly dehiscent portions of the thin ovary wall of adjoining locules; the process (except sometimes in 7’. leptocarpa) remains attached at one end, its actual length obscured on external ex- amination by a network of fine ribs. Seeds indefinitely numerous, mostly less than 1 mm. long, mostly chestnut- or dark brown, plump- or flattened- Poe ere 1945] McVauau: TrIopanis 23 lenticular, usually longer than wide, without prominent surface markings. Placentae axile (except in the unilocular ovary in 7. leptocarpa), extending essentially the whole length of the capsule and attached to the axis along most of its length. Type species: Triodanis rupestris Raf. |=’. perfoliata (L.) Nieuwl.]}. In the interpretation of the keys and descriptions which follow, it should be remembered that measurements, and to some extent descriptive terms, are taken from representative individuals of the several species. All the species of Triodanis, and especially T. perfoliata, are exceedingly variable and fruitful, and the plants vary so greatly in size and appearance that it is impracticable to make a single description cover all individuals, and very depauperate plants have been omitted from consideration in making up the keys and descriptions. As in many other genera in the CAMPANULACEAE, flowers produced near the end of the growing season or at other periods unfavorable to the plant may be much reduced in size; such flowers are often half or less than half as large as those normal for the species. Furthermore, as in other (especially annual) species of the same family, plants growing under adverse conditions of habitat may flower and fruit when extremely small. These very dwarf plants may be quite different in aspect from plants developed under conditions more suitable for growth, and their parts may be abnormally reduced in size, so that one can hardly give the lower limits of variation. It should likewise be remembered that the relative numbers of cleistog- amous and open flowers are probably determined in part by environment, although the basic ratios in the different species are evidently determined by hereditary factors. In a series of papers dealing with the life-history of T. perfoliata, Trent (1940a, 1940b, 1942) has shown that in this species greater numbers of flowers of both kinds are produced upon exposure to light of greater intensity or of longer daily duration, cleistogamous flowers only being produced in the weaker light or that of short duration. He sug- gests that the relation is a nutritional one, the number of flowers produced being roughly proportional to the photosynthetic activity in the plant. This agrees with the general observation that plants growing under ad- verse conditions of habitat, excluding conditions of light, are smaller and with fewer, mostly cleistogamous, flowers. Trent (1940a) has also described the development of the inflorescence in 7. perfoliata. He points out that not all the flowers at any one node develop at the same time, but that there are several (1-4) successions of developing flowers, the number of successions depending upon the vigor of the plant and so indirectly upon the habitat, the growing season and other factors of the environment. The earliest flowers to develop are cleistogamous; these appear singly at the lowest fertile nodes and under unfavorable conditions they then appear successively at all the nodes. Under favorable conditions open flowers are produced at the upper nodes during this first succession. A plant collected early in this first cycle would 24 WRIGHTIA [Vou. 1, No. 1 thus have a single cleistogamous flower in each of the lower axils and opening corolliferous (chasmogamous) flowers in the upper axils. At about the time of the appearance of the first open flowers, however, a second succession of cleistogamous flowers begins to appear at the lower nodes. These most commonly appear to be in pairs; that is, at the base of the first flower they appear to be opposite. Presumably each terminates a branch of the primary lateral axis represented by the earliest flower. The second and further successions of flowering may continue to the upper nodes of the plant under favorable conditions; in the upper nodes most of the flowers are cleistogamous after the first succession. KEY TO THE SPECIES OF TRIODANIS 1. Foliage leaves (not flower-bracts!) usually strigose or hispidulous on the upper surface; calyx-lobes mostly 8-15 mm. long, 5 in number and all those in any flower essentially alike, both in open and cleistogamous flowers; capsule trilocular (as shown by the number of pores) in both types of flowers, oblong-linear to clavate, usually 10 mm. long or more. . 2. 2. Capsule opening from apex toward base; corollas mostly 9-15 mm. Wong; Southern Pesags so el ee cs cen 1. 7’. coloradoensis. 2. Capsule opening from base toward apex; corollas mostly 6-10 mm. long; Mediterranean region and east to Iran..........2. T. falcata. 1. Leaves glabrous on the upper surface (sometimes with a few bristles near the tips); calyx-lobes usually less than 1o mm. long (usually much less than this, at least in the cleistogamous flowers), often 3 in number in the cleistogamous flowers or with 1 or 2 notably smaller than the others; capsule usually bilocular (sometimes unilocular) in the cleistogamous flowers, trilocular in the open ones, variously shaped................. a. 3. Flower-bracts lanceolate to linear, usually 6-8 times as long as wide; capsules of the cleistogamous flowers terete, subulate, curved and with more or less spreading tips, 8-12(-20) mm. long, dehiscent by longitudinal apical fractures or a single apical pore; seeds o.7-1 mm. long; prairies of the central United Sinton seo er ae 8. T. leptocarpa. 3. Flower-bracts ovate or broader, sometimes as much as 2-3 times as long as wide, but often as wide as or wider than long; capsules of the cleistogamous flowers terete or flattened, ob- long to ellipsoid or ovoid, straight and more or less appressed, usually less than 8 mm. long (occasionally to 12 mm. long), dehiscent by 2 (or 3) apical or lateral pores; seeds various... . 4. 4. Openings of the capsule linear, o.2-0.4 mm. wide, the elastic cartilage before dehiscence occupying most of the pore (i.e. with very narrow scarious margins); pores about midway between base and apex of capsule; seeds o.4—0.7 mm. long, minutely low-tuberculate in longitudinal lines; prairies, central United States.......... 00.6.5. 7. T. Holzingert. 4. Openings of the capsule broadly elliptic, oval, or rounded, 0.5-1.5 mm. wide, the narrow cartilage with broad 1945] McVauau: TRIODANIS 25 rounded scarious margins; pores about midway between base and apex, or near apex of capsule; seeds various. . s. 5. Seeds more or less quadrangular, o.4—0.6 mm. long, the surface roughened with low anastomosing longi- tudinal ridges; pores about the middle of the capsule; enstern Texas... vas ccs os sitar gs Gea SOILS 5. Seeds lenticular, often plump (occasionally umbonate), the surface smooth and more or less polished, or muriculate, but never ridged; pores near the apex or- near the midaies 2 soe re ees 6. 6. Seeds o.8-1 mm. long, smooth and highly pol- ished, biconvex but rather flat; pores at the apex of the capsule; bracts ovate to reniform, rather coarsely toothed, often broader than long, to-25 mm. broad; Missouri, Arkansas, and Oklahoma... 545.5: 5. T. lamprosperma. 6. Seeds o.5—0.6 mm. long, polished or dull, or mu- riculate, plumply lenticular; bracts and pores VATIONS soa i es ee ea Me hs 7. Pores at or very near the apex of the capsule; seeds smooth and highly polished; bracts usually longer than broad, not promi- nently veined beneath, only the midrib or 1 additional pair of veins evident; open corolla usually 1 (the terminal) only, or seattered axillary flowers with open Boras, re ee 3. T. biflora. 7. Pores well below the apex of the capsule (usually 1-1.5 mm.), usually about mid- way between base and apex; seeds mu- riculate or smooth and lustrous; bracts usually as broad as or broader than long, often with one or two pairs of veins in addition to the midrib prominent beneath; several of the upper nodes normally pro- ducing open corollas......4. 7. perfoliata. 1. Triodanis coloradoensis (Buckl.) McVaugh, comb. nov. Campanula coloradoense Buckl., Proc. Acad. Phila. 13[1861]: 460. 1862. Specularia Lindheimeri Vatke, Linnaea 38: 713. 1874- Pentagonia coloradensis Kuntze, Rev. Gen. 381. 1891. Legouzia coloradoense Heller, Contr. Herb. Frankl. & Marsh. Coll. 1: 99. 6F 1895. Specularia coloradoensis Buckl. ex Small, Fl. Southeast. U. 5. 1142. 1903. Legousia coloradoensis Briq., Candollea 4: 332. 1931- Erect, the stems 25-60 or often roo em. high and up to 3 mm. in diame- ter, prominently angled, simple or usually with ascending subordinate branches 5—15 (rarely 30) em. long from the nodes just above the middle of the plant (usually not with basal branches), the stem-angles near base 26 WRIGHTIA [Vou. 1, No. r retrorsely scabrous or hispid (occasionally smooth or with long hairs im- mediately below the leaf-bases), the upper parts usually smooth and glabrous. Leaves and bracts more or less hispid or scabrous at least on the veins beneath, their margins ciliate with numerous short, broad-based hairs, these often much longer near the base of the blade; leaves up to the first branch usually hispid above, the succeeding bracts less so, often smooth above; upper and middle foliage leaves oblanceolate or elliptic, sessile, attenuate to base and to a blunt or acute and mucronate apex, inconspicuously crenate especially on the distal half with 6-10 callose teeth on each side, the teeth often hardly visible among the stout marginal hairs, the blades mostly 5—8 times as long as wide, 0.4—1(—1.5) em. wide, 2.5-5(—7) em. long; lower and basal leaves relatively broader and more conspicuously toothed, the lowest often elliptic, rather abruptly narrowed to a blunt tip and a margined or slender petiole up to 2.5 cm. long; bracts, including those subtending flowering branches, lanceolate, sessile, narrowed and somewhat rounded at base, narrowly acute, attenuate to the callose- glandular tip, the median ones (those of the lowest sessile or nearly sessile flowers) often to—12 times as long as wide, 2-5 mm. wide, 2—4 em. long, minutely callose-denticulate. Flowers in a loosely spiciform arrangement on the main axis above the flowering branches, the flowers sessile or nearly so, 1-3 at each of the uppermost 4—6(—9) nodes on the axis and in groups of 1-3 near the tips of the flowering branches, all (including the terminal ones) prevailingly fertile and open, but those of the lower branches and/ or the lowest ones of the main spike sometimes cleistogamous; bracteoles evident even in the uppermost flowers; corolla of the cleistogamous flowers obovoid or cylindroid, 1-2 mm. long; opening corollas ‘‘blue purple” (Bebb), 9—15(—18) mm. long, the tube 2—4(—5) mm. long, the lobes 3.s—6(—8) mm. wide. Filaments mostly 2—2.5 mm. long, the distal third slender and glabrous, the proximal two-thirds dilated, oblong, ciliate. Anthers (2.7—-)3- 4.3(-5) mm. long. Capsule scabrous-papillose at least near base, oblong- linear or clavate, usually abruptly narrowed distally to the base of the calyx, 2-2.5(—3) mm. in diameter and 11~—18 mm. long (in the cleistogamous flowers) or up to 23 mm. long (in the open flowers), deeply suleate between the locules; pores opening toward the base, broadly elliptic or rounded, 1.5-2.5 mm. below the base of the calyx, o.8-1.7 mm. wide, 2-4 mm. long (the proximal end often ill-defined), the narrow central cartilage with broad rounded scarious margins, curled outward and downward into a half circle, locules and pores 3 in both types of flowers; style 6-10 mm. long, the branches mostly 1-1.5 mm. long. Calyx-lobes 5 in both types of flowers, lanceolate, loosely spreading in fruit, with slender subulate tips, usually with 1 or 2 capillary bristles at tips, the margins scabrous-ciliate, in the open flowers all alike (o.7—)1-1.5(—1.8) mm. wide, (6~)10-17 mm. long, in the cleistogamous flowers a little shorter and often unequal in width. Seeds brown, smooth and lustrous, biconvex and more or less flattened, usually about 0.7 mm. wide and 0.8—-1 mm. long.—Dry rocky hillsides and bluffs, limestone ledges, rich woods, flood plains and gravel bars, flowering from gag = Nad ph a aD ee ae Po BSUS RS SPN ne ee 1945] McVaucu: TRIODANIS 27 late April to early June, Edwards Plateau and adjacent territory, south- central Texas. Representative specimens examined*: TEXAS: Bell Co., Salado, S. E. Wolf 2966 (TA&M, US); Bexar Co., Leon Springs, Mr. & Mrs. J. Clemens 878 (Mo,R), San Antonio, V. Havard in 1884 (US), E. H. Wilkinson 87, 152 (Mo), R. Bebb 2343 (Okla); Blanco Co., Blanco, E. J. Palmer 33955 (Mo); Comal Co., bei Braunfels, F. Lindheimer 65 [III 450, isotype of S. Lind- heimeri] (Mo); Crockett Co., Ozona, V. L. Cory in 1925 (TA&M); Gillespie Co., Crab Apple Cr., G. Jermy 468 (Mo); Kendall Co., Lindendale, Palmer 9885 (Mo, US); Kerr Co., Kerrville, A. A. Heller 1731 (ANS, IA, Mo, R, US); Kimble Co., J. Reverchon 582 (ANS, Mo, TA&M, US); Menard Co., Menard, Palmer 11855 (Mo); Real Co., Thousand Springs, H. B. Parks & V. L. Cory 8568 (TA&M); Travis Co., Austin, Z. Hall 384 (Mo, US), A. A. Armer (UTex 5520] (Tex, US), Palmer 13652 (Mo, US); Uvalde Co., Uvalde, Palmer 33712 (ANS, Mo); Valverde Co., Devils River n. of Del Rio, H. A. Pilsbry in 1903 (ANS); Wichita Co., Red River, B. C. Tharp in 1922 (IA); Williamson Co., Georgetown, G. B. Wolcolt 300 (USNA); County un- determined, ‘“‘On the upper Colorado, Texas,’ S. B. Buckley in 1861 (ANS, type!); without locality, Lindhetmer III 450 (ANS, Mo, US, all doubtless isotypes of S. Lindheimeri); Blanco, C. Wright s.n. (Mo); without locality, Wright 1432 (ANS); Valley of the Rio Grande below Dofiana, C. C. Parry et al. [Mex. Bound. Surv. 698, in part] (US); Clearwater, Oregon, Spalding s.n. (ANS). One of the above records needs confirmation, namely the collection from Wichita County, Texas, which is well out of the known range of the species. The specimen alleged to have been collected by Spalding at Clear- water, Oregon [i.e. Idaho], is evidently erroneously labelled. The type of Campanula coloradoense, collected by 8. B. Buckley in June, 1861, on the “upper Colorado, Texas,’ now at the Academy of Natural Sciences, is a mature flowering specimen with numerous open flowers. The type of Specularia Lindheimeri was at Berlin; the only speci- men cited by Vatke was Lindheimer’s no. 450, of Fascicle III, collection of 1846. I have not seen the type, which may now have been destroyed, but specimens bearing this number and date are widely distributed in herbaria and all represent the same species. Lindheimer’s number 450 seems °In the preparation of this paper I have examined material from the following herbaria, indicated below in the citation of specimens by appropriate abbreviations. To those who have made this material available for study, I am very grateful: Academy of Natural Sciences, Philadelphia (ANS); Colorado State College, F ort Collins; Her- barium of C. C. Deam, Bluffton, Indiana; Gray Herbarium, Harvard University (GH); Universitetets Botaniske Museum, Copenhagen (Haun); Herbarium of L. C. Hinckley, Marfa, Tex.; Iowa State College, Ames (IA); Kansas State College, Manhattan (Kans); Louisiana State University, Baton Rouge (LSU); Missouri Botanical Garden, St. Louis (Mo) University of Oklahoma, Norman (Okla); Rocky Mountain Herbarium, University of Wyoming (R); Herbarium of Robert Runyon, Brownsville, Tex. eae yon); Southern Methodist University, Dallas, Tex. (SMU); Sul Ross State pee ers College, Alpine, Tex.; University of Texas, Austin (Tex); Texas A. & 0 io College Station (TA&M); North Texas State Teachers College, Denton (NTSTC); U. S. National Herbarium (US); U. S. National Arboretum (USNA). 28 - WRIGHTIA [VoL. 1, No. 1 to have been applied to more than one collection, however, for at the Missouri Botanical Garden are specimens each bearing the number 450 but collected, respectively, in April 1845, and May 1846. Presumably the latter is the collection of which Vatke had a sheet, for the specimen is ac- companied by Lindheimer’s original handwritten label in addition to the printed label bearing the number 450 and the “‘Fasce. III’; the handwritten label bears the number 65, evidently a field-number, and the date ‘‘Mai 1846,” with the locality “bei Braunfels” and the notation ‘‘30 Exemplare,”’ the latter suggesting wide distribution for the plants collected at this time. It should be pointed out here that if this species be referred to Specularia the valid name for it is Specularia Lindheimeri Vatke, for the transfer of Campanula coloradoense Buckl. to Specularia is prevented by the existence of the name Specularia coloradoensis Buckl. ex Small, which was published legally but probably inadvertently, without reference to Campanula coloradoense or to any previous publication by Buckley. The many features that this species shares with T'riodanis falcata are certainly indicative of some relatively close relationship or of a remarkable parallel development. Were it not for the capsule of 7’. coloradoensis open- ing from apex toward base, I should suggest the establishment of a formal section of 7’riodanis, to include these two species. This anomaly, however, suggests that the two species should not be too closely linked on the basis of what may be fortuitous resemblances. Triodanis coloradoensis and Campanula Reverchoni A. Gray are unique in their respective genera in the direction in which their capsules open; that is, from apex toward base. Except for this feature common to both, neither species shows any marked anomalies. The geographical propinquity of the two (both are confined to south-central Texas) inevitably suggests some phylogenetic connection between them, but they are dissimilar in morphology except for this one character, and the apparent relationship between them is hardly closer than that between 7’. falcata, for example, and Campanula ramosissima. One may argue that this peculiarity of the capsule of these species suggests for them an origin different from that of the remainder of the species of Triodanis and Campanula, or suggests a di- or polyphyletie origin for these genera. On the other hand it emphasizes the fact that the relationship between genera, specifically between Triodanis and Campanula, is probably not to be thought of as a simple linear one, but as a series of links between groups of species. It is unwise to attempt to reason too far from this single capsule-character, however, for it seems likely that it is not a particularly fundamental feature; the direction of opening of the capsule depends upon the relative strengths of the tissues at the two ends of the elastic portion of the capsule, and these may well undergo a change in the majority of any population in comparatively few generations. 2. Triodanis falcata (Ten.) MceVaugh, comb. nov. Prismatocarpus faleatus Ten., Prodr. Fl. Nap. 16. 1811. Campanula syriaca R. & S., Syst. 5: 133. 1810. 1945] McVauaGu: TRIODANIS 29 Campanula falcata R. & 8., Syst. 5: 154. 1810. Specularia falcata A. DC., Monog. Campan. 345. 1830. Prismatocarpus scaber Lowe, Trans. Camb. Phil. Soe. 6: reimp. 16. 1838. Specularia falcata var. scabra (Lowe) A. DC, in DC., Prodr. 7(2): 490. 1830. Specularia castellana Lange, Ind. Sem. Hort. Haun. 1854: 25. 1855. Specularia falcata var. pusilla Boiss., Fl. Or. 3: 960. 1875. Pentagonia falcata Kuntze, Rev. Gen. 381. 1891. Legousia falcata Fritsch, Mitteil. Naturw. Ver. Wien 5: roo. 1907. Erect or reclining, the stems (5—)15-60 em. high and 1-2.5 mm. in diameter at base, smooth and glabrous or minutely scabrous on the angles, or the whole plant scabrous or hispidulous. Foliage leaves nearly glabrous, strigose or hispidulous above and on the veins beneath, finely and densely ciliate with very short broad-based hairs except near base, the decurrent bases and the petiolar bases of the lowermost blades bearing on each margin a row of slender white cilia 1-2 mm. long; bracts mostly glabrous except for the ciliate or scabrous margins; blades of the foliage leaves sessile, elliptic to oblanceolate or obovate, rounded or blunt-pointed at tip, acute at base or the lower attenuate into a margined short petiole, all crenate with 2—6 shallow teeth per em., 2-3(—4) times as long as wide, mostly 7—10(—20) mm. wide, (8-)15~-30(—40) mm. long, the lower leaves usually smaller and rounder, with definite petioles up to 6-10 mm. long (sometimes all leaves petiolate, the lower petioles slender, up to 4 cm. long); bracts sessile, mostly ovate, with broad rounded base and acute tip (blunt and mucronate at extreme tip), the median ones mostly 2—-2.5 times as long as wide, 4-10(-20) mm. wide, 18-27(—40) mm. long, inconspicuously crenate with 6-7 teeth on each side, often appearing subentire. Flowers in a loose spike occupying 6-15 nodes below the terminal flower, at least those of the lower nodes cleistogamous; branches at the lower nodes of the spike sometimes appreciably elongated and bearing 2 or 3 fertile flowers, but usually only the terminal flower maturing; terminal flower of the main axis usually maturing fruit, often open. Corolla of the cleistogamous flowers cylindroid, o.8-1.6 mm. long; opening corollas rose-colored (according to A. DC.), pale purple (Cowgill), or purple (Koelz) 6-10 mm. long, the tube 1-2.5 mm. long, the lobes about 2.5—3.5 mm. wide. Filaments about 1.2—-1.5 mm. long. Anthers (1.7—)2-3.5 mm. long. Capsules in both sorts of flowers trilocular, prominently scabrous at least on the 3 angles of each locule, oblong-linear or clavate (in depauperate plants shorter and obovoid), 2.5-3 mm. in diameter, 8-16(—25) mm. long, often slightly curved, rather deeply sulcate and the pores sunken; pores at the distal extremity of the capsule, 0.71.2 mm. wide, 2-3 mm. long, the central cartilage narrow, with broad scarious margins, divaricate or completely recurved after opening. Style 5-6 mm. long, the branches 1 mm. long. Calyx-lobes 5, all alike or those of the cleistogamous flowers slightly shorter and 1 or 2 narrower than the others, all linear-subulate or often broader, smooth and glabrous or sometimes ciliate or scabrous on the margins, mostly 1-2 mm. wide at base and 9-16 mm. long (sometimes up to 8 mm. wide and 3 em. long), often spreading or recurved-faleate, the margins often revolute. Seeds bright chestnut- 30 WRIGHTIA [Vou. 1, No. brown, smooth, highly polished, lenticular, about o.8-1 mm. long and al- most as wide.—Fields, orchards and cultivated grounds generally, dry rocky woods, shaded rocks, in areas bordering the Mediterranean Sea and on the Mediterranean islands; Canary Islands: east to Iran and the Cau- casus; flowering and fruiting from April to July. Flowers intermediate between the cleistogamous ones and the normal open ones have open but narrowly tubular corollas up to 6 mm. long, style up to 4.5 mm. long and anthers 1.2—1.5 mm. long. The type of Prismatocarpus falcatus came presumably from Naples, Italy, or its vicinity; I have not been able to examine Tenore’s original description of P. falcatus, or any authentic specimens. Specimens examined: CANARY ISLANDS: without locality, Desprieux (ANS); Gran Canaria, Canaria, A. C. Cook 355 (US); Lanzarote, Rib. de Sta. Luzia, J. M. Moniz 795 (GH); Teneriffe, Laguna in arvis, H. de la Perrandiére in 1855 (GH, Mo); Hieris-Las Lapas, C. J. Pitard in 1905 (Mo). BALEARIC ISLANDS: Majorque, Barranco de Soller, EH. Bourgeaw in 1869 (US). FRANCE: Var: LeLuc, Metz (?) in 1868 (US); Cannes, ex hb. Joad, in 1851 (Mo); Alpes Maritimes, Mt. Authion, M. Moggridge in 1868, in part (US). CORSICA: without loc., “Serafino misit Sept. 1832” (GH); stagno de Palo- Bachbett, P. Aellen 2779 (Mo). SARDINIA: Sta. Teresa Gallura, par Tempio, E. Reverchon in 1881 (GH,US). ITALY: Liguria occid., Ventimiglia, C. Bicknell [Fiori & Béguinot 2366] (GH); ‘massif di Pegli Lig[uriae] occid.,” coll. in 1856 (Mo); Corniglia (Spezia) presso le Tre Crosi, O. Mattirolo Fon- tana (?) in 1928 (USNA); Anxur [Terracina], J. Ball in 1841 (GH); Lesina Island, Dalmatia, Roemer (hb. G. Engelmann, Mo). CYPRUS: “in mont. pr. Palaeo Khova,” Sintenis & Rigo 23 in 1880 (US). PALESTINE: Banias, Post 95 in 1877 (ANS). SYRIA: near Brumana, A. Kneucker in 1904 (GH). IRAN: “In umbra rupium m. Kuh-Ajub prope ruinas u. Persepolis,’ Th. Kotschy 407 (isotype of var. pusilla Boiss., Mo); Talimansur, Bakhtiari, W. Koelz 15085 (USNA). FROM CULTIVATION: “FE, semin. H. Parisiensis (1893) in Ho. experim.'' Vallisumbrosae culta,” R. F. Solla in 1894 (US); from seeds collected in Iran (W. Koelz 6036), U. S. Plant Introduction Gar- den, Glenn Dale, Md., W. Cowgill 2435 (USNA). Specularia falcata var. pusilla Boiss. is the name applied to the ecologi- cally dwarfed form of this species. The original specimens collected by Kotschy resemble those taken by Koelz (no. 15085) in the same general region; the plants are mostly 10 em. high or less and open flowers are not developed. Plants grown at Glenn Dale, Maryland, from Iranian seed collected by Koelz, however, grew 30-50 em. high, although the height of the original plants was noted by Koelz as “about 6 inches.” If trans- ferred to Triodanis, Boissier’s name should be used to designate a forma, not a variety. Boissier (1875) cites various specimens that I have not seen, including examples from Mt. Parnes, Attica (coll. Heldreich), the Island of Rhodes (Bourgeau), Cilicia (Balansa), the littoral of Syria and lower Lebanon (Blanche, Gaillardot) Mt. Carmel, Palestine (Boissier), and Mts. Avroman, Schahu, Pir Omar, Gudrun, all of Kurdistan ( Haussknecht). 1945] McVaucu: TRIODANIS 31 Additional locality records for Syria and Palestine may be found in Post’s flora (1933, p. 170), and a single record for the Caucasus (Alagir, on the Ardon, Lagofsky) is given by Fomin (1906). The status of Specularia castellana Lange is unknown to me; perhaps it should be recognized as a species, but I have not seen any material refer- able to it. According to Lange it differs from S. falcata by its shorter calyx- lobes and the scabrosity of the whole plant. Lange supposed also that his plant was the same as S. falcata var. scabra (Lowe) A. DC. Many European workers have recognized both S. falcata and S. castellana. The two species are separated by Coste (Fl. France 2: 491. 1937) as follows: 1. Calyx smooth, the lobes about as long as the tube at anthesis, the corolla about a third as long as the calyx-lobes; plant more or less glabrous. Ae re ean Per brn ware Mae dees cr edie a ERE Ss S. falcata 1. Calyx scabrous, the lobes a third or a half as long as the tube, about equal- ing the corolla; whole plant more or less scabrous......... _S. castellana, 3. TRIODANIS BIFLORA (R. & P.) Greene, Man. Bot. San Francis. Bay 2 30, without citation of name-bringing synonym. 1894; Nieuwl., Am. Mid]. Nat. 3: 192. 1914. Campanula biflora R. & P., Fl. Per. 2: 55. 1790. Campanula montevidensis Spreng., Syst. 1: 738. 1825. Specularia biflora Fisch. & Mey., Ind. Sem. Hort. Petrop. 1: 17. 1836. Dysmicodon californicum Nutt., Trans. Am. Phil. Soc. n. s. 8: 256. 1843. Dysmicodon ovatum Nutt., Trans. Am. Phil. Soc. n. s. 8: 256. 1843. Campanula intermedia Engelm. ex Nutt., Trans. Am. Phil. Soc. n. s. 8: 256, in syn. 1843. Specularia californica Vatke, Linnaea 38: 714. 1874. Specularia ovata Vatke, Linnaea. 38: 713. 1874. Campanula ludoviciana Torr. ex A. Gray, Proc. Am. Acad. 11: 83, in syn. 1876. Pentagonia biflora Kuntze, Rev. Gen. 381. 1891. Legouzia biflora Britton, Mem. Torrey Bot. Club 5: 309. 10 Oct. 1894. Specularia perfoliata f. ramosa Arech., Anal. Mus. Nac. Montevideo 7[FI. Uruguay 4]: 14. 1900. : Specularia perfoliata f. rigida Arech., Anal. Mus. Nac. Montevideo 7[Fl. Uruguay 4]: 14. 1909. Slender, erect or reclining, the stems 15~45(—80) cm. long, I-I. 5(-3) mm. in diameter, densely to rather sparsely hirsute below with pale flaccid hairs, the hairs reduced above and the upper parts of the stem retrorsely hispid or merely scabrous. Leaves and bracts hispid (chiefly on the veins) to completely glabrous beneath; foliage leaves elliptic to ovate, the lower- most often obtuse and short-petiolate, the upper more acute and sessile, all the blades crenate, 6-12 mm. wide, 1o-30 mm. long, usually 1.7—-2.5(—3) times as long as wide; bracts relatively broader, ovate, sessile and some- what clasping, sharply acute and often mucronate at tip, often spreading and usually not much concealing the fruits, the middle ones 2.5~12 mm. wide, 6-16(-20) mm. long, usually 1.3-1.8(-2.5) times as long as wide, the margins crenate on the proximal two-thirds of the blade with 1-3 teeth 43 WRIGHTIA [Vou. 1, No. 1 on each side of the midrib, the basal teeth occasionally prolonged and the bracts then 3—-(s—) lobed. Flowers at most or all the (10o—50) nodes of each axis, the main axis terminated by an open flower; axillary flowers 1—4(—8) at each node, usually consisting of a terminal and 2 lateral ones, but one or two often aborting, one flower in this lateral sub-inflorescence occasionally producing an open corolla in vigorous plants, but only the terminal flower of each main branch regularly of this type. Corollas of the cleistogamous flowers rounded, scarcely 0.5 mm. high; opening corolla ‘‘blue,” ‘‘purple,” “Jilac,”’ “violet,” ‘white with violet lines,’’ ‘“‘pinkish-blue” or “‘rose-red,” glabrous or sparsely hirtellous on the veins without and ciliate at the tips of the lobes, s—o(-11) mm. long, the tube 1-1.5(—2) mm. long, the lobes 2—3(—4.5) mm. wide. Filaments 1.3—2 mm. long, the proximal half gradually or abruptly expanded. Anthers (1.4—)1.6—2.2(—2.6) mm. long. Capsule glabrous or sparsely hispid, at maturity ellipsoid, clavate, obovoid or ovoid, mostly about 1.3-1.7 mm. in diameter (sometimes 2 mm., or when bilocular 1 by 2 mm.) and 4.5~7 mm. long (occasionally 2-9 mm.), the oval or nearly circular pores at the distal extremity or occasionally as much as 0.3-0.5 mm. below it, 0.5—0.8(—1.0) mm. wide, 0.7—1.8(—2.2) mm. long, with nar- row central cartilage and broad scarious margins, tightly uprolled after opening; locules and pores usually 2 only in the cleistogamous flowers. Style 3.5-6.5 mm. long (2 mm. in depauperate plants), the branches 1 mm. long or less. Calyx-lobes triangular or triangular-ovate (then somewhat narrowed at base), acutely pointed and often mucronate, glabrous or sparsely hirtello-scabrous, closely appressed or somewhat spreading or re- curved in fruit, those of the open flowers 0.7—1.5(—3) mm. wide, (3.5—) 4-6(—7) mm. long; those of the cleistogamous flowers 3-5, 0.3—1 mm. wide, 1~3(—4) mm. long, 1 or 2 often much smaller than the others. Seeds chest- nut-brown, smooth and highly polished, biconvex, o.5-0.65 mm. long, a little longer than wide.—Paraguay, Uruguay and southern Brazil (Rio Grande do Sul, Rio de Janeiro); Argentina and Chile south about to lat. 41° and north to Tucuman and Jujuy; cis-Andean Peru and Chile and highlands of trans-Andean Peru and of Ecuador; sandy embankments, grassy slopes, margins of forest, cultivated fields, sandy beds of dry rivers, coastal deserts, at elevations from sea-level to about 3000 meters, flowering from September to February (also collected May to August in the Peruvian highlands). Southeastern Virginia to Florida and westward in the Coastal Plain and Piedmont; west of the Appalachian Mountains from southern Kentucky to southeastern Kansas and southward to southern Texas, Nuevo Leén and San Luis Potosi; central and southeastern Arizona; northern Baja California to southwestern Oregon; in various situations, including waste and cultivated fields, prairies, dry hills, forests, moist river-banks and deep canyons, chiefly at low elevations, flowering and fruiting from April to June or earlier in southern Florida, southern Texas, Arizona and southern California. The South American plants of this species appear to be not even vari- etally distinct from those of North America, although the former are often 1945] McVauacu: TrIODANIS 33 (but by no means always) more densely pubescent. In the material I have examined the only measurable differences between the two series are those which appear to be the result of inadequate sampling of one population or the other. The corollas of most South American specimens, for example, measure between 6 and 7.5 mm. in length, while those of most North Ameri- can plants measure between 7 and 9 mm., but with a fair sprinkling of corollas 5-6 mm. long, and some 9.5 to 11 mm. The anthers of South American plants rarely exceed 2 mm. in length, while many North American specimens have them 2.2 mm. long, and a few have them 2.5 to 2.6 mm. The calyx- lobes of the open flower rarely exceed 1.5 mm. in width, but in one South American specimen they measure 3 mm. Open flowers in addition to the terminal one are occasionally met with both in North American and South American specimens, but the percentage of plants bearing such flowers is higher among the South American, as far as my examination shows. Most South American material of 7. biflora in American herbaria has been identified as Specularia perfoliata, but apparently that species does not occur naturally in South America, except possibly in Ecuador. The type of Campanula biflora R. & P., as represented in Field Museum nega- tive no. 29455, is an average plant of the rather hairy South American form of the species here called Triodanis biflora. John Torrey seems to have been the first to understand clearly that in the southeastern United States there were two distinet species, both of which had before him usually been referred to Campanula perfoliata of Linnaeus. Torrey noted in his flora of New York (1843, vol. 1, p. 428-9) that he had described “the structure of the early flowers of this plant [Specularia perfoliata], and of an allied but very distinct species from Louisiana,’ in a memoir read before the New York Lyceum of Natural History, in 1830. Apparently this allied species, Trzodanis biflora, was called by Torrey Campanula ludoviciana, for this name is found on her- barium labels attached to plants collected as late as 1875, but he seems never to have published it, although on this point Gray (1876) was in doubt. Nuttall likewise recognized that Campanula perfoliata had been too broadly interpreted, and also supposed the Californian plants of what we now call T'riodanis biflora to comprise a species distinct from those of the southern and central states. The type-material of Dysmidocon californicum is apparently not represented in the herbarium of the Philadelphia Acad- emy, but from Nuttall’s description it is evident that his plant must have been 7. biflora. Under Dysmicodon ovatum Nuttall gave Campanula inter- media Engelmann as a synonym, with the locality “Fort Gibson, Arkan- sas,” andthe additional locality “Arkansas and Louisiana.” In the Acad- emy’s collection is a Nuttall collection from “‘Ark.,” labelled with Nuttall s asterisk indicating a new species, and the name “ovata” associated with an unpublished generic name; this may be considered the type collection of Dysmicodon ovatum. The plant is characteristic ts riodanis biflora. Mounted on the same sheet is a plant of the same species designated by 34 WRIGHTIA [Vou. 1, No. 1 Nuttall with the generie and specific names that appear on his Arkansas collection; in another hand is written on the ticket “‘Louis{iana] Trudeau.” This is evidently the basis for Nuttall’s reference to Louisiana. Also in the Academy’s collection is a sheet of Engelmann’s collection of Campanula intermedia, made in ‘‘Arkansas,’”’ June 1835. This collection includes at least 4 plants of Triodanis biflora and 1 which may be of hybrid origin; it bears Engelmann’s original label, but no definite locality. Presumably, however it is the collection from Fort Gibson alluded to by Nuttall. In the southeastern United States, where 7. biflora and T. perfoliata are both common and are often found growing in intimate association, it may be supposed that the two species hybridize; certainly there are occa- sional plants which are not referable with certainty to either species and which must be regarded as intermediates. These will be discussed under T. perfoliata. I do not think there is justification for the view expressed by Holzinger (1892, p. 210) that the two species should be combined into one because of the many puzzling forms. In spite of these puzzling intermedi- ates, Triodanis biflora is apparently biologically distinct from 7’. perfoliata; perhaps the best evidence for this may be found in its habit of producing but a single open flower, even under favorable conditions for growth. This is evidently determined by generic factors in this species, while in 7’. per- foliata and the 3 species segregated from it in this paper individual reduc- tions in the number of open flowers are determined chiefly by environment, and in average healthy individuals open flowers are produced at several of the upper nodes. 4. TRIODANIS PERFOLIATA (L.) Nieuwl., Am. Midl. Nat. 3: 192. tor4. Campanula perfoliata L., Sp. Pl. 760. 1753. Campanula amplexicaulis Michx., Fl. Bor.-Am. 1: 108. 1803. Campanula flagellaris H. B. K., Nov. Gen. & Sp. 3: 301. 1819. Prismatocarpus perfoliatus Sweet, Hort. Brit. ed. 1, 251. 1826. Specularia perfoliata A. DC., Monog. Campan. 351. 1830. Triodanis rupestris Raf., N. Fl. N. Am. 4: 67. 1837. Triodallus rupestris Raf. ex A. DC., in DC., Prodr. 7(2): 491, in syn. 1839. Dysmicodon perfoliatum Nutt., Trans. Am. Phil. Soc. n. s. 8: 256. 1843. Pentagonia perfoliata Kuntze, Rev. Gen. 381. 1801. Legouzia perfoliata Britton, Mem. Torrey Bot. Club 5: 309. 10 Oct. 1804. Exceedingly variable in size, posture, and vestiture, the stems mostly 30-60 em. high and 1~3 mm. in diameter, but sometimes more than 1 m. high or in poor soil only 5-8 em. high and then very slender; stems usually erect (sometimes decumbent or prostrate in woods and wet places), the base pilose to hispid with white hairs up to 1.5 mm. long, the upper parts pilose to hispid like the base, or the hairs shorter and the stems retrorsely hispidulous or scabrous. Leaves and bracts densely ciliate with short broad-based sharp hairs, beneath scabrous, or vestite like the stems (at least near base on the principal veins), or the bracts smooth and glabrous except for the margins; blades of the median foliage leaves sessile, ovate- cordate, the lower varying to broadly elliptic with abruptly contracted eee. 1945] McVauau: TRIODANIS 3 wn base, the lowest (often withered at flowering time) obovate with attenuate base or with distinct elliptic or suborbicular blade and slender petiole up to 1.5 em. long; all blades blunt or rounded at tip, often somewhat un- dulate-crisped, coarsely to obscurely crenate with 3-6 teeth on each side, mostly 1.5-2 times as long as wide, up to about 2.5 em. wide and 3.5 cm. long; bracts relatively much broader and more coarsely toothed, ovate to reniform, sessile and usually strongly cordate-clasping and concealing the fruit, the tip blunt (in shade forms sometimes rounded and the bract as a whole suborbicular) to abruptly acute and mucronate, the median (o. 5) 0.6-0,8(—1.0) times as long as wide, up to about 3 em. wide and 2~2.5 em. long, crenate with about 4—6(—10) teeth on each side (in some forms varying to almost entire or to sharply serrate). Flowers in average plants at the upper 20-35(—60?) nodes, the lowest 6-20(—40) (or usually 14-24 of all fertile nodes) bearing cleistogamous flowers, the upper nodes usually bearing open flowers; terminal flower usually open; axillary flowers 1-3 (or up to about 8) at a node. Corolla of the cleistogamous flowers less than 0.5 mm. high; opening corollas bluish-lavender (in rare individuals white), 8-12(-13) mm. long, the tube (1.5—)2-3.5(-5) mm. long, the lobes (2-) 3~4.5(-5.5) mm. wide. Filaments 1.5s—2.5 mm. long, the proximal half abruptly dilated. Anthers (2—)2.s—3.3 mm. long. Capsule at maturity quite smooth or retrorsely scabrous on the angles, usually obovoid or oblong, I.5-2.5 mm. in diameter and 4-6(-8) mm. long (in the cleistogamous flowers) or up to 3.5 mm. in diameter and ro mm. long (in the open flowers), in the open flowers sometimes somewhat suleate and the pores sunken; pores broadly elliptic or rounded, the uplifted portion (0.6—)o.8-1.5 mm. wide, 1.5—3(—4) mm. long, tightly curled after opening, with narrow cen- tral cartilage and broad rounded scarious margins, its distal extremity 1~1.5(—2.5) mm. below the summit of the capsule (or rarely at the summit !) ; locules and pores 2 or 3 in the cleistogamous flowers. Style 5-7.5 mm. long, the branches 1-r1.5 mm. long or less. Calyx-lobes triangular to lance- olate or ovate, with slender subulate tips, often with 1-2 capillary bristles at tips, glabrous or the margins scabrous-ciliate; lobes in the open flowers (0.7—-)1.3-2.5 mm. wide, (4—)5-8.5 mm. long, those of the cleistogamous flowers 3~5 (usually 3 or 5) in number, variable in size in the same flower, 0.5-2 mm. wide, 14.5 mm. long. Seeds ellipsoid, brown, biconvex or slightly flattened on the sides, about o.4 mm. wide, o.5-o.6 mm. long, somewhat lustrous, either muriculate over the entire surface or the proc- esses reduced in various degrees or apparently wanting, the surface of the seed then smooth and lustrous but not highly polished, broken by tiny irregularities—In various habitats (but avoiding deserts and permanently wet soil) almost throughout the United States; often occurs as a weed in disturbed areas. British Columbia to southern Ontario and Maine, south to northern California and in the Sierra Madre to Chihuahua (and So- nora?); southern Mexico; occurs (perhaps as an introduction) in Jamaica and Hispaniola and in Ecuador; rare or uncommon along the Rieern borders of the United States and in the more arid portions of the Rocky 36 WRIGHTIA fVou. 1, No. 1 Mountain and Great Basin States; common in eastern United States and west to western Nebraska, Kansas and Oklahoma, and to the 1ooth meridian in Texas; not reported from Nevada, southern Florida, or south- ern Texas; flowers from April to July (sometimes earlier in Texas and in Florida); occurs at elevations up to 6000 feet (in Colorado). Although this species, like most of its congeners, readily occupies dis- turbed areas throughout its range and is often regarded as a weed, it is by no means universally distributed. Its principal area of distribution is in eastern United States south of the Great Lakes and, like many other native species of the same area, its western limit is approximately at the tooth Meridian, it is absent from the southern tip of Texas and the south- ern half of peninsular Florida, and it is relatively rare or entirely absent from the glaciated zone across northern New England, northern New York and west to the prairies. In addition to this primary area of distribution, however, the species is apparently native along the mountain chains from Chihuahua, Arizona and western Texas to Montana and British Columbia, its western range thence extending south, chiefly along the Cascade Ranges, to northern California. This species is represented in herbaria by relatively few collections from outside of the United States and Canada, and there is a possibility that it is not native in southern Mexico, South America, and the West Indies; it may have been introduced into these areas from the United States at an early date: Specimens examined: MEXICO: Curauanua, Cafion del Rio Piedras Verdes, C. V. Hartman 685 (GH, US); Mexico, Salto de Agua, C. A. Purpus 1708 (GH, Mo, US); Oaxaca, Chinantla, Liebmann pl. mex. 9230 (Haun); PurBLa, Boca del Monte, G. Arséne 7096 (US), Tezuitlan, C. R. Orcutt 4021, in part (Mo); Sonora, Los Animos, G. Thurber 328 in 1851 (GH); VERACRUZ, Las Vigas, EF. W. Nelson 17 (US), Atoyac, E. Kerber 489 (US), km. 320, be- tween Orizaba and Fortin, J. K. Langman 3443 (USNA); Jalapa, C. L. Smith 1665 (GH). JAMAICA: Abbey Green, Oreutt 5175 (US), Cinchona, Wm. Harris 12078 (GH, Mo, US). HISPANIOLA: Sto. Domrnao, prope Constanza in pineto, von Tuerckheim 3345 (GH, IA, Mo, US). ECUADOR: Pichincha, ad flum. Machangara pr. Quito, J. Holmgren 523 (US), Perucho, W. Jameson 642 (ANS); [the type of Campanula flagellaris H.B.K. (not seen by me) was col- lected ‘‘inter Quito et Puembo’’]. It has been supposed, or at least inferred, by many writers, that plants of this species producing no open flowers were reproducing normally and producing normal viable seeds. Trent has found, however (1942), that al- though sporogenesis and embryogeny are similar in the two types of flowers, the open flowers produce approximately 47.5% viable seed, while the seeds of cleistogamous flowers germinate in but 0.07% of the samples tested. Ungerminated seeds, when sectioned, are usually found to be with- out embryo, although apparently normal in appearance and with fully developed endosperm. Although Triodanis perfoliata is exceedingly variable in vegetative 1945] McVauGu: TRIODANIS 37 characters, most of the variation is probably related to conditions of the habitat; variations in stems, leaves and pubescence appear to be those usually associated with modifications of local conditions, and show no correlation with regional distribution. Often two plants on the same her- barium sheet are superficially very dissimilar, one perhaps strict and markedly pubescent, the other decumbent and weak and more nearly glabrous. The flowers and fruits show very little variation from plant to plant, except as their size may be increased in vigorous individuals. The one exception so far discovered is in the seeds; as this seed-character seems to be correlated with regional distribution, although not with any other morphological character, it may be of some significance. The seeds of Triodanis perfoliata are about o.5 mm. long and a little longer than wide, plump, irregularly lenticular, with thin blunt edges; under a magnification of about 15 diameters they appear smooth, with a dull or slightly lustrous surface. With a magnification of 20 diameters, a good source of light, and a white background, however, it may be seen that the surface is thickly beset with minute erect dark points—one of my colleagues suggests that these resemble a heavy crop of sprouting whiskers—these are the muriculate seeds referred to in the description of this species. Not all plants have muriculate seeds, however. On seeds of some individuals the spinulose processes are so reduced as to be invisible except under higher magnification, and then are sometimes found only on the margins of the seed. On seeds of other plants the surface appears to be entirely smooth and without irregularity except that close inspection will often disclose what appear to be minute flat spots or perhaps depressions on the surface; the mass effect of these seems to be the dispersal of the reflected light from the surface, so that even these smooth seeds are con- siderably less lustrous than the seeds of Triodanis biflora, which they resemble otherwise. Fortunately for the investigation of this character, mature or nearly mature seeds are available in practically all herbarium specimens of this species. On the other hand, ample suites of specimens are not available from many areas, so that it is not always possible to ascertain the relative abundance of smooth- and muriculate-seeded plants from an area. It has been possible, however, to make the following generalizations: — In the eastern Great Plains states, north to Kansas and Missouri and east to the Mississippi River, plants with muriculate seeds predominate in the following approximate ratios: Ratio of Number of State muriculate/smooth seeds plants examined Kansas 15/1 : 5 Missouri 11/1 5 Oklahoma 9/2 04 Arkansas 12/1 25 Texas 14/1 . 5 Louisiana 9/2 38 WRIGHTIA [Vou. 1, No. 1 In the southern Rocky Mountains approximately the same ratios pre- vail, but the evidence is less conclusive because of the fewer specimens available: Ratio of Number of State muriculate/smooth seeds plants examined Utah 3/0 3 Colorado 5/1 20 Arizona 8/1 9 New Mexico 6/1 7 North and east of these areas plants with smooth seeds are much more frequent; throughout most of the states east of the Mississippi River the two types occur in approximately equal numbers, both often at the same locality as indicated by their presence on the same herbarium sheet. Ap- parently the muriculate type preponderates in the inland states south of the Ohio River and the smooth type becomes most abundant on the south- eastern Coastal Plain and in areas most recently glaciated. The smooth type occurs almost to the exclusion of the other along the northern edge of the range of the species, except in the Rocky Mountains, where both occur. The following table gives the ratios for the eastern states and smaller areas discussed above; the ratio was approximately 1/1 for a small number of specimens in those states not included: Ratio of Number of State muriculate/smooth seeds plants examined Georgia (Piedmont except 1 plant) 7/0 7 Tennessee 4/2 9 Kentucky 6/o 6 West Virginia 4/1 5 Pennsylvania 5/6 55 New Jersey (Coastal Plain) 1/4.3 32 Delaware o/2 2 Maryland (eastern) 1/4 10 District of Columbia 1/1 22 Florida 1/3 20 Conditions in the northwestern states and in British Columbia may be tabulated as follows: Ratio of Number of * State muriculate/smooth seeds plants examined British Columbia 0/4 4 Washington o/19 19 Oregon 1/1 ite) California 0/3 3 Idaho 5/7 12 Montana 3/2 5 Wyoming 1/5 II North Dakota ° South Dakota 0/10 10 Minnesota 1/2 3 lowa 1/7. 8 Nebraska a : ty 1945] McVauGu: Triopanis 39 In New England, where the species is relatively less frequent than in the central states, the tabulation is as follows: Ratio of Number of State muriculate/smooth seeds plants examined Maine 1/2 3 New Hampshire 1/2 3 Vermont 0/3 3 Massachusetts 1/2 22 Rhode Island 1/2 3 Connecticut 1/2 9 North of Kansas and Missouri plants with muriculate seeds are rarely found; apparently there is an abrupt termination of range here. The ratios for Missouri and Iowa, for example, show a complete reversal in the relative abundance of the two types in these two states. Presumably the figures for Kansas and Nebraska would be similar, but very few specimens from Nebraska have been available. The above data may be interpreted tentatively as follows: This species originally occupied a part or all of the Appalachian-Ozarkian region of eastern United States. Its seeds were all or principally of the muriculate type. Plants with seeds of this type early became established in suitable situations in the Rocky Mountains as well as in the eastern prairies and forested areas. More recently the smooth-seeded type has developed and has spread onto the southeastern Coastal Plain and northward into glaci- ated territory more extensively than plants of the original type. It is possible that this smooth-seeded type may be a polyploid, since polyploid variants of otherwise diploid species are in many instances known to oc- cupy new areas more rapidly than the original diploids. The accompanying map (fig. 2) shows the distribution of the two forms of Triodanis perfoliata, each dot representing an actual specimen from the locality. The apparently great relative abundance of the species in Kansas, Oklahoma, and Texas is due at least in part to the fact that collectors in these states have been active, while in Nebraska, for example, but little collecting has been done. It seems probable, however, that this is essentially a prairie species, represented by most individuals in most localities in the states from Iowa and Nebraska southward, so that the picture given by the map may not be as misleading as may appear at first glance. ot Mention was made above, in the discussion devoted to Triodanis biflora, of the plants which appear to be intermediate between this species and T. perfoliata. These intermediates are relatively few; as far as can be ascertained from material in herbaria they comprise less than 1 percent of the combined populations of the two species. The supposed intermediate nature of any individual plant is suggested chiefly by some aberrance in one or more of the following: number of open flowers, position of the opening in the capsule, degree and nature of surface roughness of the seeds, and shape and venation of the bracts. Certain plants perhaps appear intermediate because of dwarfing or the reverse, induced by conditions of the habitat: it is difficult to say, at least in herbarium material, whether a 40 WRIGHTIA [Voxu. 1, No. 1 plant is a dwarfed, narrow-bracted and smooth-seeded form of 7’. perfoliata in which the open flowers have been reduced to 1 by adverse conditions, or whether it is an intermediate form having the bracts and inflorescence of T. biflora and the capsules of 7. perfoliata. On the other hand, certain luxuriant plants have the numerous open flowers of perfoliata and rather broad leafy bracts, but the other features of biflora; if these are of mixed Fig. 2. The distribution of Triodanis perfoliata in the United States and adjacent Canada and Mexico. Solid dots indicate the occurrence of the form with muriculate seeds, open circles the form with smooth seeds; half-open circles indicate the occurrence of both forms in the same county or parish. parentage, certainly they owe most of their characteristics to bzflora. Herbaria_ contain very few specimens that can be interpreted as possible intermediates between perfoliata and biflora. The following have been seen: LOUISIANA: Acadia Par., Crowley, D. E. Ellis 20 (LSU; bracts inter- mediate; infl. of perfoliata; caps. of biflora); East Baton Rouge Par., Baton Rouge, collector & date unknown (LSU; dwarfed, bracts narrow, open fl. 1, caps. of perfoliata, seeds rough but not muriculate). TEXAS: Bexar Co., San Antonio, G. Jermy 345, in part (GH; 2 plants are perfoliata; 3rd has bracts like perfoliata but more pointed; infl., caps. and seeds of biflora). Brazos Co., College Station, R. G. Reeves 71, 72 (USNA; infl. of perfoliata; otherwise biflora but bracts rather broad); Denton Co., 3 mi. s. w. of Denton, W. L. McCart 1003 (NTSTC; infl. & bracts of perfoliata; caps. & seeds of biflora). Relatively more frequent, at least in herbaria, than the supposed inter- mediates between 7. perfoliata and T. biflora, are plants which seem to combine the characters of 7. perfoliata on the one hand, and 7’. terana or 1945] McVauau: TRIODANIS 41 T. Holzingeri. This is presumably to be expected, if these intermediate plants are the result of hybridization, since 7’. perfoliata seems less closely related to 7. bzflora than to the others. In eastern Texas, where in some localities all 4 species occur, there are rather numerous individuals of a type that is not referable to any of the species. These plants usually have several or many of the upper nodes bearing open corollas; the bracts are often as broad as long, and 5-nerved, but more narrowly and acutely pointed than in 7. perfoliata, suggesting in this respect 7’. biflora or 7. Holzingeri; the capsule opens near the middle or slightly above it; the seeds are usually obscurely tuberculate, rather than either smooth or muriculate, and are usually prominently umbonate. Specimens of this general type are: TEXAS: Without definite locality, C. Wright (GH); Bexar Co., San An- tonio, V. Havard in 1884 (US); Brazos Co., without loc., R. G. Reeves 1421, south of F. & B. station, Reeves 72A, Bryan, Reeves 18A, near Jones Bridge, Reeves 869, 869B, College Station, U. H. Williams 4/18/30, in part, 4/24/30 (all TA&M); Brooks Co., s. of Encino, C. L. Lundell & Amelia A. Lundell 12767, in part (SMU; 1 plant typical Holzingeri, 5 are intermediates); Burleson Co., Lyons, W. German s. n., in part (Tex); Cameron Co., without loc., Mrs. A. M. Davis in 1942, in part (Tex); Goliad Co., Goliad, Rev. C. B. Williams 117 (ANS, Tex); Gregg Co., without loc., Biltmore herb. 968e, in part (US); Grimes Co., Navasota, collector unknown [Flower Contest] in 1934 (Tex); Harris Co., Buffalo Bayou, Houston, J. F. Joor in 1876 (LSU), Houston, G. L. Fisher 57, in part (US), Hockley, F. W. Thurow 20 (US); Lee Co., 13 mi. n.e. of McDade, R. R. Innes 879 (GH); Llano Co., Granite Mt., B. C. Tharp in 1931 (Tex); Walker Co., Huntsville, M. S. Young in ror4, in part (Tex); Washington Co., without loc., EZ. Brackett s.n., Apr. 30 (Tex); Willacy Co., 1 mi. n. Sebastian, R. Runyon 2515 (Runyon), end of highway, Runyon 1610 (Runyon). Specimens which appear surely to combine the characters of 7’. per- foliata and T. Holzingeri are listed below (see also Lundell & Lundell 12767, above). These have the relatively narrow, pointed, and stiff bracts of the latter species, but the capsule openings are intermediate in width between the two species: TEXAS: Bell Co., Taylor’s Valley, S. E. Wolff 2950, in. part (TA&M; a plant of typical Holzingeri forms part of the same collection); DeWitt Co., without loc., M. Riedel in 1942 (Tex); Erath Co., without loc., L. C. Gough in 1921, in part (Tex; also a plant of Holzingeri); Kleberg Co., Kingsville, Ji F. Sinclair in 1940, in part (Tex, GH, also a plant of Holzingeri on the Gray Herbarium sheet); Maverick or Valverde Co., Del Rio to Eagle Pass, £. Whitehouse in 1934 (Tex). A few specimens appear certainly to combine the characters of 1’. per- foliata and T. texana. Plants of typical texana are mounted on the sheets with each of the supposed intermediates cited below: TEXAS: Brazos Co., near Jones Bridge, R. G. Reeves 869A (TA&M); this collection consists of 1 plant of 7. biflora, 1 of T. texana, and 2 resembling 42 WRIGHTIA [Vou. 1, No. 1 terana but with acute bracts, the seeds nearly smooth, more or less quad- rangular; Van Zandt Co., n.w. of Edom, C. L. Lundell & Amelia A, Lundell 10456 (SMU); this collection consists of 1 plant of 7. perfoliata, 1 of texana, and 1 resembling perfoliata except that the seeds are smooth but longitudinally lined; this latter plant is intermediate in degree of pubescence between per- foliata and texana. 5. Triodanis lamprosperma McVaugh, sp.nov. Habitu 7. perfoliatae; bracteae plerumque serratae, capsula apice dehiscens; semina nitida, o.8-1 mm. longa. Erect, the stems (15—)30-60(—85) em. high and up to 4 mm. in diam- eter, the base pilose to hispid or retrorsely scabrous, the upper parts retrorsely scabrous to entirely glabrous. Leaves and bracts scabrous or hispid beneath at least on the veins near base (or the bracts glabrous and smooth on both surfaces), finely and densely ciliate with very short broad- based hairs except near base, where pilose with slender white hairs up to 1.5 mm. long; blades of the foliage leaves sessile, oval, or the lowest obovate and attenuate into a short petiole, all rounded at tip and crenate with about 6 teeth on each side, 1.3~—2 times as long as wide, mostly 7-12 mm. wide and 8-25 mm. long; bracts relatively and actually broader, ovate to reniform, sessile and often strongly clasping and usually concealing the fruit, bluntly pointed (often abruptly acute and mucronate), the median mostly o.6—-1 times as long as wide, 1o-25 mm. wide, 8-16 mm. long, uni- formly crenate-serrate with 3-7 teeth on each side, the teeth up to 1.5 mm. high. Flowers at the upper 15~s0 nodes, the lowest (6—)12—-25(—40) fertile nodes (or usually 14 to 24 of all fertile nodes) bearing cleistogamous flowers, the upper nodes usually bearing open flowers; terminal flower often aborting; axillary flowers 1-3 at a node, the terminal of the trio usually maturing, the laterals often aborting. Corolla of the cleistogamous flowers less than 0.5 mm. high; opening corollas probably bluish-purple, 9-12 mm. long, the tube 1.5—2.5(—4) mm. long, the lobes 4-5 mm. wide. Filaments 1.8-2 mm. long, the proximal half abruptly dilated. Anthers 2.5-3 mm. long. Capsule at maturity retrorsely scabrous on the angles to nearly or quite smooth, usually obovoid or oblong, 1.5—2 mm. in diameter and 4-6(—7) mm. long (in the cleistogamous flowers) or 2.3—3 mm. in di- ameter and 7-10 mm. long (in the open flowers), in the open flowers often suleate and the pores somewhat sunken; pores broadly elliptic or rounded, at the distal extremity of the capsule, 0.7-1.3 mm. wide, 1.4-2.5 mm. long, with narrow central cartilage and broad rounded scarious margins, usually tightly curled after opening; locules and pores predominately 2 in the cleistogamous flowers. Style 5.7-7.2 (usually about 6.5) mm. long, the branches 1 mm. long or less. Calyx-lobes triangular (or in the corolliferous flowers often truly lanceolate and then broadest 2-3 mm. above base), with slender subulate tips, usually with 1 or 2 capillary bristles at tips, glabrous or the margins scabrous-ciliate, in the open flowers 0.8—1.6(—2.5) mm. wide, 5-5-9 mm. long, those of the cleistogamous flowers 3-5, (0.7) I~1.7 mm. wide, (1.8—)2.5-4 mm. long. Seeds smooth and highly polished, 1945] McVavuau: TRIODANIS 43 biconvex or slightly flattened on the sides, o.6-0.8 mm. wide, 0.8-1.0 mm. long (usually about o.7 mm. wide and 0.8 mm. long).—Dry rocky ridges and ledges, open rocky woods, glades, moist woods and flood-plains, flow- ering and fruiting from early May to mid-June; southern Missouri, western Arkansas and eastern Oklahoma, at elevations up to 2800 ft. Type in the Gray Herbarium, Harvard University, collected May 6, 1940, by Delzie Demaree (no. 21152), on dry rocky ridges of Fourche Mountain, Little Rock P.O., Pulaski Co., Arkansas, at 500 feet elevation; isotypes in the herbaria of the Missouri Botanical Garden and Iowa State College. Additional specimens examined: MISSOURI: Barry Co., Eagle Rock, EF. J. Palmer 30434 (Mo); Iron Co., Iron Mountain Lake, J. H. Kellogg 1216 (Mo); Jasper Co., Carterville, Palmer 2148 (Mo), Neck City, Palmer 3962 (Mo); Shannon Co., Monteer, B. F. Bush 432 (GH, Mo). ARKANSAS: Logan Co., Magazine Mt., D. Demaree 17725, 21309 (Mo); Yell Co., Mt. Nebo State Park, Demaree 21285 (Mo). OKLAHOMA: Haskell Co., without loc., R. Bebb 5471 (Okla); Latimer Co., e. of Gowen, O. Clark 2696, in part (Okla); Leflore Co., Kiamichi Mts., 7. Johnson 96 (Okla), near Stapp, Hop- kins & VanValkenberg 4283 (Okla); MeCurtain Co., Garvin, Demaree 12645 (Okla), swamp of Little Creek, Eagletown, Hopkins & VanValkenberg 4192, in part (Okla). This species has previously been confused with 7’. perfoliata, but it seems undoubtedly distinct, with a well defined range in the Ozark-Ouachita region. Vegetatively it may usually be distinguished from 7. perfoliata by the conspicuously serrate and imbricate bracts and the relatively smooth upper portions of the stems. 6. Triodanis texana McVaugh, sp.nov. T. perfoliatae similis, sed inferiore parte caulis et faciebus inferioribus foliorum inferiorum conspicue hirsutis; semina quadrangularia, rugoso- reticulata. Slender, erect or reclining at base, the stem 10-75 (usually 25~50) em. long, 1~1.5(—4.5) mm. in diameter, hirsute below with colorless flaccid hairs up to 1 mm. long, densely so on the angles and often between them, the hairs much reduced above and the upper parts of the stem retrorsely hispid or merely scabrous. Lower leaves hirtellous to hirsute beneath es- pecially on the veins, the upper foliage leaves and all bracts sparsely hirtellous to (often) perfectly glabrous except the margin, which in all leaves and bracts is ciliate with short stiff broad-based hairs; blades of the foliage leaves sessile or the lower with attenuate bases, the uppermost (immediately below the floriferous nodes) ovate, the median elliptic, the lowermost oblanceolate to obovate, all rounded or obtuse at tip or the uppermost blunt-pointed, all uniformly crenate from tip to near base with 4-6 (9) teeth on each side, mostly 1.5-2.5 times as long as wide, mostly 6-15 mm. wide, (10-)15~30(—40) mm. long; bracts relatively broader, ovate, sessile and somewhat clasping, obtuse to sharply acute and often 44 WRIGHTIA [Von. 1, No. 1 mucronate at tip, often spreading and exposing the fruits, the middle ones mostly o.7—1.0 times as long as wide (rarely twice as long as or only about half as long as wide), 6-17 mm. wide, 4-22 mm. long, the margins rather uniformly crenate with 3-8 teeth on each side (in very small bracts the teeth reduced to glandular callosities). Flowers in the upper 1o—30(—40) axils, the lowest fertile nodes (usually 5~15, or 14-24 of all fertile nodes) bearing cleistogamous flowers only, the upper nodes usually bearing open flowers; terminal flower open or cleistogamous; axillary flowers 1-3 at each node, usually 1 only (the terminal) maturing fruit, the two laterals usually aborting. Corolla of the cleistogamous flowers less than o.5 mm. high; opening corollas probably bluish-purple, glabrous, 7—10(—14) mm. long, the tube 1.5—2.5(-4) mm. long, the lobes 2-5 mm. wide. Filaments 1.5—2.2 mm. long, the proximal half gradually or abruptly dilated. Anthers (1.7—) 2.1-3 mm. long. Capsule at maturity glabrous or sparsely hirtellous, el- lipsoid or ovoid or in the open flowers truncate and clavate, 1.5—-2.5 mm. in diameter, 4-6(—7) mm. long, the oval pores about the middle of the capsule (1-2.5 mm. below the base of the calyx-lobes), 0.7—1(—1.6) mm. wide, (1.5—)2—3 mm. long, with narrow central cartilage and broad scarious margins, tightly curled after opening; locules and pores usually 2 in the cleistogamous flowers. Style 5—7.5 mm. long, the branches 1 mm. long or less. Calyx-lobes triangular or triangular-ovate, acutely pointed and often mucronate, or with a capillary tip, glabrous or sparingly ciliate like the bracts, erect or spreading in fruit, those of the open flowers 0.g—1.5(—2) mm. wide, 3—6(—7) mm. long; those of the cleistogamous flowers 3~-5, 0.5— 1.2 mm. wide, (1—-)1.5-3 mm. long, often all alike, but sometimes 1 or 2 smaller than the others. Seeds dark chestnut-brown, lustrous, o.2—-0.3 mm. in diameter, 0.4-0.6 mm. long, more or less quadrangular with usually concave or flat sides and obtuse angles, the surface roughened with low narrow anastomosing ridges in longitudinal rows.—In woods, pinelands, plains, open banks and depressions, mostly in sandy soils, flowering in April and May; eastern Texas. Type in the herbarium of the Missouri Botanical Garden, no. 828770, collected April 26, 1917 by E. J. Palmer (no. 11697), in sandy open ground at Somerville, Burleson Co., Texas; isotypes in the Rocky Mountain Herbarium of the University of Wyoming and in the herbarium of Iowa State College. Additional specimens examined: TEXAS: Bastrop Co., Bastrop to Elgin, B. C. Tharp in 1940 (Tex); Brazos Co., R. G. Reeves 869A, in part (TA&M); Caldwell Co., without locality, J. B. McBryde in 1931 (Tex); Dallas Co., Dallas, J. Reverchon 2087 (Mo), B. F. Bush 680 in 1900 (Mo, US); Erath Co., without locality, L. C. Gough in 1921 (Tex); Fayette Co., Colony, HE. W. Crawford 38 (US), 41 (Mo); Frio Co., Moore, E. J. Palmer 33880, in part (GH, TA, Mo); Gonzales Co., Ottine, ZH. R. Bogusch [UTex 1435] (Tex), Tharp in 1935 (GH), Gonzales, H. B. Parks & V. L. Cory 8369 (TA&M), without locality, Bogusch (UTex 1433] (Tex); Hardin Co., Fletcher, Palmer 9550 (1A, Mo); Milam Co., Between Milano & Gause, S. E. Wolff 4010 (TA&M); Travis Co., Austin, Tharp in 1935 (GH); Van Zandt Co., Edom, C. L. Lundell & Amelia A. Lundell 10456, in part (SMU). nese tt scciesbieneangemen “inate eet s eaceeenteattebintiasiatip te ke be Aan CLS ha PAL aS OTN NP ila 6 ALON, aap a oe eNO = agri 1945| McVauanu: TRIODANIS 45 Superficially this plant is little different from 7. perfoliata except in the seeds, but it also differs from that species in its more hirsute lower stems and lower leaves, its nearly glabrous bracts and upper stems, and its somewhat shorter calyx-lobes. Apparently it may be associated in nature with 7. perfoliata and sometimes with T. Holzingeri, as herbarium sheets occasionally bear both or all three together. 7. Triodanis Holzingeri McVaugh, sp.nov. Habitu 7. perfoliatae, sed strictior et bractibus plerumque adpressis et acutis; capsula versus mediam partem valvulis linearibus dehiscens; semina tuberculata. Slender, erect, the stem 25—60(—85) em. high, up to 4 mm. in diameter, strict and simple or with few branches, the base hispid or hirsute with stiff or flaccid colorless hairs up to 1 mm. long, the upper parts retrorsely hispid or merely scabrous. Leaves and bracts scabrous beneath with short broad-based hairs chiefly on the veins or the bracts essentially glabrous; blades of the foliage-leaves sessile or the lower with attenuate bases, the uppermost ovate, the median elliptic, the lowermost oblanceolate or ob- ovate, all rather blunt-tipped with a rounded point and a very short glandular mucro (the uppermost acute), all more or less undulate-crisped and uniformly crenate from base to apex with 5-8 teeth on each side, mostly 1.6-2.5(—3.2) times as long as wide, mostly 5—10(~13) mm. wide, (10~)16-24(—30) mm. long; bracts relatively broader, ovate, sessile and somewhat clasping, sharply acute and often mucronate at tip, often more or less appressed in fruit or only the tips spreading, often exceeded by the mature capsules, the median mostly 1-1.5 times as long as wide, (5—) 7-15(-20) mm. wide, 7-20 mm. long, the margins uniformly crenate from base to apex with 3—6 teeth on each side, or the distal third entire, deltoid; margins of the foliage-leaves ciliate with short broad-based hairs; margins of bracts glabrous or similarly vestite, but often revolute except at base and so appearing glabrous. Flowers at the upper 20-50(—85) nodes, the lowest to—so fertile nodes (usually 15-30, or 14 to 7% of all fertile nodes, but often all fertile nodes) bearing cleistogamous flowers, the upper nodes usually bearing open flowers; terminal flower open or cleistogamous; axil- lary flowers 1-3 at a node, the terminal usually maturing fruit, the laterals often aborting. Corolla of the cleistogamous flowers less than o.5 mm. high; opening corollas probably bluish-purple, glabrous or with coarse hairs without on the midveins of the lobes, 7-9(—11) mm. long, the tube 1~2(—2.5) mm. long, the lobes 2—3(—5) mm. wide. Filaments about 1.6~—2 mm. long, the proximal half gradually or abruptly dilated. Anthers 2~2.6 mm. long. Capsule at maturity scaberulous to essentially glabrous, ellipsoid or oblong or in the open flowers truncate and obovoid, often strongly angled, 0.7-1.5 mm. thick and 2—2.5 mm. wide (when bilocular) or 1.5— 2.5 mm. in diameter (when trilocular), (3.5—)6-8(-12) mm. long, often deeply sulcate between the locules and the pores thus sunken; pores linear, about the middle of the capsule (1-2.7 mm. below the base of the calyx- lobes or rarely less than 1 mm.), 0.2—-0.4(—o.5) mm. wide, 1.4-2.5 mm. long, 46 WRIGHTIA [VoL. 1, No. 1 the cartilage filling the entire width of the pore, or with very narrow scarious margins, after opening divaricate or curved into an open ring; locules and pores predominately 2 in the cleistogamous flowers. Style 4.5—6 mm. long, the branches 1 mm. long or less. Calyx-lobes triangular or tri- angular-ovate, acute and often attenuate-faleate, often with 1 or 2 capillary bristles at tip, glabrous, or scabrous on midrib without, or on midrib and margins, often recurved-spreading in fruit, those of the open flowers 1— 1.5(—2.5) mm. wide, 4-6(—7.5) mm. long; those of the cleistogamous flowers 3-5, 0.7-1.5(—2.5) mm. wide, (1.5—)2.5—6 mm. long, often either 3 or 5 and all alike, but sometimes 1 or 2 smaller than the others. Seeds dark chestnut- brown, lustrous, lenticular, about o.25—-0.4 mm. wide, o.4-0.7 mm. long, minutely low-tuberculate in longitudinal lines——Open prairies and plains, river bottoms, canyons, in sandy and gravelly soils, flowering and fruiting from May to July (or earlier in southern Texas and Arizona). South- eastern Wyoming to western Missouri, south in the prairies to southern Texas; with outlying stations in western Tennessee and southeastern Arizona. Type in the herbarium of the National Arboretum, no. 102701, col- lected on open prairies near Washington, McClain Co., Oklahoma, June 12, 1936, by Milton Hopkins and C, T. Eskew (No. 688 of the Plantae Exsiceatae Grayanae). Additional specimens examined: LOCALITY UNKNOWN: Rocky Mts., C. Thomas' in 1860, in part (US). ARIZONA: Graham Co., 8 mi. s.w. of Saf- ford, B. & R. R. Maguire 11766 (GH); Pima Co., Tucson, M. Zuck in 1806 (US), J. W. Touwmey in 1894 (GH, US), “Catalina Mts. near Tueson,” V. L. Cory 3511 (GH), Santa Rita Mts., C. G. Pringle’, “Apriland May, 1881,” in part (Mo). KANSAS: Barber Co., Medicine Lodge, A. S. Hitchcock in 1892 (Kans); Cloud Co., ?Aurora, S. V. Fraser 384 (Kans); Ellsworth Co., Kanopo- lis, M. Becker in 1896 (Kans); Gray Co., without locality, Hitchcock in 1897 (Kans); Seward Co., Liberal, Hitchcock in 1892 (Kans). MISSOURI: Jack- son Co., without locality, B. F. Bush in 1888 (Mo), Courtney, Bush 120 in 1899 (Mo), Bush 6846 in 1912 (GH, Mo, US), Sheffield, Bush 429 in 1895 (Mo). NEBRASKA: Thomas Co., Dismal R., s. of Thedford, P. A. Rydberg 1346, in part (US). OKLAHOMA: Blaine Co., Greenfield, J. Engleman 197 (Okla); Caddo Co., Devil’s Can., near Hinton, G. W. Stevens 946 (GH, Mo, US); Cleveland Co., near Norman, F. L. Hambrick 33, C. Whaley 26, C. C. * The collector’s name is given thus on the label. This is Cyrus Thomas (1825-1910), later an eminent ethnologist. In 1869 he undertook the post of entomologist and botan- ist to the Hayden Survey. A catalogue of the plants collected in 1869 was published by C. C. Parry in the Report of the Hayden Survey for 1870, pp. 484-487, but nothing is known in regard to the localities where they were collected, except as these may be inferred from the route of the Hayden party as given in the report for 1869. The speci- men of T’riodanis Holzingeri cited above was probably collected in June, somewhere along the line of the railroad between St. Joseph, Missouri, and Cheyenne, Wyoming. * The source of Pringle’s material may be questioned, as the sheets of this collection distributed in different herbaria bear not only this species, but also 7’. perfoliata and T. biflora, and the collector evidently assembled the material over a period of some weeks and perhaps from several localities, in the assumption that it was all the same species. 1945| McVauau: TRIODANIS 47 Smith 484, C. T. Eskew 1018, 1813, Hopkins & Cross 117 (all Okla); Custer Co., Cedar Twp., L. Mericle 647 (Okla), Barnitz Twp., Mericle 494 (Okla); Ellis Co., Shattuck, R. L. Clifton [Stevens 3120) (GH); Greer Co., Mangum, Rk. Bull in 1928 (Okla); Kingfisher Co., Lacey, V. Young (Stevens 1565] (GH); Kiowa Co., 4 mi. e. Headrick, R. R. Innes & B. Moon 996 (GH, USNA); Logan Co., n.w. part of county, C. C. Smith 677, 709, 736 (all Okla), without locality, M. A. Carleton 167 in 1891 (US®*), Guthrie, Stevens 3221 (GH); Mur- ray Co., Arbuckle Mts., M. Hopkins 6052 (Okla); Noble Co., without locality, P. J. White in 1900 (R); Pottawatomie Co., St. Louis, M. Faulkner 103, in part (Okla); Woods Co., Alva, Stevens 653 (GH, Mo, Okla, US), Freedom, Stevens 726 1/2 (GH); Woodward Co., Fort Supply, J. P. Kimball in 1891 (Kans); County undetermined, between Ft. Cobb & Ft. Arbuckle, Z. Palmer 168 in 1868 (US). TENNESSEE: Shelby Co., President’s Island, P. O. Memphis, D. Demaree 21373 (Mo). TEXAS: Bell Co., Taylor’s Valley, S. FE. Wolff 2950, in part (TA&M); Bexar Co., San Antonio, FE. D. Schulz in 1921 (Tex); Brewster Co., Alpine, H. J. Cottle in 1926 (Sul Ross), F. J. Palmer 30525 (Mo, Tex), E. L. Reed 1835 (US); Brooks Co., Encinal, C. L. Lundell & Amelia A. Lundell 12767, in part (SMU); Childress Co., Childress, Childress H. S. s. n. (Tex); Comanche Co., Comanche, H. Eggert in 1900 (Mo); Erath Co., Stephenville, J. C. Brown 2 (Tex), H. B. Parks & V. L. Cory 13171 (TA&M), without locality, Z. C. Gough in 1921, in part (Tex); Frio Co., Moore, E. J. Palmer 33880, in part (GH, IA, Mo); Kleberg Co., Kingsville, J. F. Sinclair in 1940 (GH); Maverick Co., Eagle Pass, C. G. Pringle 9184, in part (GH, US); Midland Co., Midland, FE. Whitehouse in 1932 (Tex); Mitchell Co., Colorado, H. Eggert in 1900 (Mo); Nueces Co., Corpus Christi, C. R. Orcutt 5870 (Mo); Parker Co., Millsap, EZ. Wadsworth in 1927 (Tex); San Patricio Co., Mathis, 8S. D. McKelvey 1720 (GH); Travis Co., Austin, A. M. Ferguson in 1901 (Tex), Lake Austin silt, Whitehouse in 1933 (Tex); Webb Co., sands, Laredo, J. Reverchon in 1903 (Mo); Wheeler Co., Mobeetie, B. C. Tharp |U Tex 6298] (Tex, US, USNA); County undetermined, Valley of the Rio Grande below Donana, C. C. Parry et al. [Mex. Bound. Surv. 697, in part] (US), Johnson City to Fredericksburg, Whitehouse in 1929 (Tex). WYOMING: Platte Co., Whalen Canyon, A. Nelson 514 in 1894 (GH, R, US). This species is named for John M. Holzinger (1853-1929), who seems to have been the first to suspect that plants of it were unlike those of typical 7. perfoliata. In his account of Carleton’s plants from the Indian Territory (1892), Holzinger associated Carleton’s no. 167 with Palmer’s no. 168, C. Thomas’ collection from the Rocky Mountains, and no. 679 lie. 697] of the Mexican Boundary Survey; he considered all these the same peculiar “form of S. perfoliata;” all are now to be referred to Trio- danis Holzingeri. 8. TRIODANIS LEPTOCARPA (Nutt.) Nieuwl., Am. Midl. Nat. 3: 192. 1914. ?Triodanis scabra Raf., N. Fl. N. Am. 4: 67. 1837. 2 Campylocera leptocarpa Nutt., Trans. Am. Phil. Soe. n. s. 8: 257. 1843. Campanula leptocarpa Engelm. ex Nutt., Trans. Am. Phil. Soc. n. s. 8: 257, in Syn. 1843. _ ®No locality is given on the label, but in Holzinger’s account of Carleton’s collec- tions (1892, p. 210), the locality for this number is given as “Guthrie, Cimarron Valley.” 48 WRIGHTIA [VoL. 1, No. 1 Campylocera leptocarpa var. glabella Nutt., Trans. Am. Phil. Soc. n. s. 8: 257. 1843. Specularia Linsecomia Buckl., Proc. Acad. Phila. 13[1861]: 460. 1862. Specularia leptocarpa A. Gray, Proc. Am. Acad. 11: 82, 1876. Pentagonia leptocarpa Kuntze, Rev. Gen. 381. 1891. Legouzia leptocarpa Britton, Mem. Torrey Bot. Club. 5: 309. 10 Oct. 1894. Slender, erect or reclining, the stems 1o—so0(—75) em. high and 1-2 (—3.5) mm. in diameter, the base spreading-hispid or retrorsely scabrous, the upper parts (occasionally) hispid or (usually) retrorsely scabrous to entirely glabrous. Leaves and bracts scabrous or prickly-hispid beneath at least on the veins, or the bracts smooth; leaves and bracts finely and more or less antrorsely ciliate with short broad-based hairs, these somewhat longer near the base of the blade; blades of the foliage-leaves sessile, el- liptic to oblanceolate, or the lowest oblanceolate to obovate and attenuate into short petioles; blades acute and blunt-pointed, callose-tipped, or the lowest rounded, all inconspicuously callose-toothed or shallowly crenate with 3-6 teeth on each side, mostly 3-6 times as long as wide, 3—5(—8) mm. wide, 15-35 mm. long, the lower smaller and relatively broader; bracts lanceolate to linear, sessile, acute and callose-tipped, the median 3.5-10 (usually 6-8) times as long as wide, 2-6 mm. wide, 15-25 mm. long, callose-denticulate or crenate like the leaves, the lower bracts o ten con- spicuous and longer than the sterile leaves. Inflorescence often very many- flowered with many and crowded flowering nodes but not appearing dense because of the spreading, slender capsules; flowers at the uppermost 15- 50(—65) nodes, the lowest 1o-35(—55) fertile nodes (or usually 14 to *4 of all fertile nodes) bearing cleistogamous flowers, the upper nodes usually bearing open flowers alone, or both open and cleistogamous; terminal flower open, or often aborting; lateral flowers 1-7 in each axil, the terminal flower strongest; bracteoles linear, present at the bases of the weaker lateral flowers and usually even when these are abortive, but often com- pletely wanting at 1-flowered nodes. Corolla of the cleistogamous flowers less than o.5 mm. high; opening corollas “violet”” (Pennell), mostly 7-10 mm. long, the tube 1.5—2 mm. long, the lobes 3-4 mm. wide. Filaments 1.5-2.3 mm. long, the proximal half abruptly dilated. Anthers (1.8—)2.2- 2.8(—3.2) mm. long. Capsules prickly-hispid to retrorsely scabrous or quite smooth and glabrous, markedly dimorphic, those of the cleistogamous flowers terete, usually subulate, more or less faleate with divaricate tips and often spirally twisted, or straight and appressed, 0.7—1.5 mm. in di- ameter, 8-12(—20) mm. long, unilocular with parietal placentation, at length dehiscent in the distal half by longitudinal slits alternating with the calyx lobes, or less frequently by a single pore up to 2.5 mm. from the apex, 0.5~o.7 mm. wide and 1.7~2.6 mm. long, the relatively broad carti- laginous process broadly fusiform, occupying most or essentially all the opening, opening from base toward apex but ultimately free at both ends, curved into a half-circle but not tightly rolled; capsules of the open flowers usually straight and more erect, thicker and longer, 15-25 mm. long, linear, ae 1945] McVaucu: TRIODANIS 49 terete and unilocular (and then usually dehiscent by slits only) and 1.3~ 1.8 mm. in diameter, or flattened and suleate and imperfectly or perfectly bilocular (with one or 2 apical pores, respectively) and then 1-1.5 mm, thick, 2-2.5 mm. wide. Style (3.5—)5.5—6.5(—7.5) mm. long, the branches 1 mm. long or usually less. Calyx-lobes subulate, faleate or widely divari- cate or ascending in fruit, glabrous or often somewhat ciliate like the leaves; lobes variable in length and width in both types of flowers, those of the open flowers o.5—1 mm. wide, 6-10(—15) mm. long, those of the cleisto- gamous flowers 3~5 in number, 0.3—0.7 wide, 2—-6(—10) mm. long; seeds light brown, smooth and lustrous, biconvex and somewhat flattened, with rounded ends, about o.4-0.6 mm. wide, o.7—1 mm. long.—Prairies, barrens, rocky open slopes and ridges, dry or moist woods, bottoms and draws, creek banks; mostly in limestone outcrops and glades, apparently begin- ning to flower in late April (in southern Texas) or mid-May (Oklahoma) and continuing throughout the summer; prairies from southern Texas to Montana, North Dakota and Minnesota. This species is known from relatively few localities from the northern tiers of states: MONTANA: Cascade Co., Sand Coulee, R. S. Williams 310 (US); Gal- latin Co., 16 Mile Creek, Scribner 131 in 1883 (ANS, US), near Bridger Peak (Fort Ellis), W. B. Platt [Hayden Survey, 1872] (ANS, US); Park Co., Liv- ingston, HE. W. Scheuber in 1901 (IA). NORTH DAKOTA: Fort Berthold Indian Reservation, collector unknown, no. 88, in 1935 (USNA); Morton Co., Glen Ullin, G. A. Holzinger 29 (US). MINNESOTA: Hennepin Co., Fort Snelling, W. H. Forwood in 1888 (US). WYOMING: Sheridan Co., Upper Little Goose, V. Willits 228 (R). SOUTH DAKOTA: Meade Co., near Ft. Meade, W. H. Forwood 245 1/4 (US), Bear Butte, A. C. McIntosh 699 (R); Pennington Co., Cedar Pass near Interior, E. J. Palmer 37646 (Mo); Tripp Co., twp. 99, H. J. Rehorst in 1938 (USNA), Winner, W. H. Over 15838 (US). NEBRASKA: “Prairies, Mauvaises Terres,” F. V. Hayden in 1853 (Mo; this probably came from the “Bad Lands” of what is now South Dakota, but the label is headed “Flora Nebraskana’”’). IOWA: Winnebago Co., without loc., D. R. Porter in 1923 (IA). COLORADO: Boulder Cafion, W. A. Henry 191 (ANS); without loc., C. C. Parry in 1862 (IA), and 1864 (Mo). In Kansas this species is abundant and common except in the western fifth of the state, and is equally so in Oklahoma except possibly in the extreme northeast and in the counties immediately east of the Texas Pan- handle. In Texas it is known to occur in Bexar, Collin, Dallas, Llano, Palo Pinto, Parker, Tarrant, and Travis counties. Along the eastern edge of its range it occurs in southwestern Missouri (Benton, Cass, Cedar, Dade, Jackson, Jasper, Polk, St. Clair, and W ash- ington counties) and is said to be introduced in Macon Co. (Ethel, Bf. Bush 7607 in 1915, Mo). It isknown to me from a single locality in Arkansas (Logan Co., Magazine Mt., Demaree 17724, 21331, Mo), and it has been reported by Deam (1940, p. 1094) as a weed along the railroad in Indiana. No type-material of Campylocera leptocarpa has been found at Phila- 50 WRIGHTIA [Vou. 1, No. 1 delphia, unless a collection made by George Engelmann at Fort Gibson, Arkansas (now Oklahoma), in June, 1835, be taken for the type. This collection was designated by Engelmann as his new Campanula leptocarpa, and is evidently the same one cited by Nuttall under Campylocera lepto- carpa var. glabella. In Nuttall’s original description of C. leptocarpa he cited neither any definite collection nor any locality except ‘‘Arkansas”’ for the species proper, while under the variety he cited not only the lo- cality, Fort Gibson, but also the collector and the manuscript name pro- posed by him. Engelmann’s collection is therefore certainly the type of var. glabella; at least one large plant of the 5 on the sheet fits the descrip- tion (‘‘stem and leaves nearly smooth’’). It is of course possible that Nuttall had no collection of his own to refer to C. leptocarpa; if so the same Engel- mann collection may be regarded as the type of his species, for at least two of the plants on the sheet are noticeably shaggy, fitting his description (“stem hirsute at the angles, leaves ciliate’). The type of Specularia Linsecomia Buckley, an over-mature fruiting specimen, is likewise at the Academy of Natural Sciences of Philadelphia, mounted on the same sheet with the type of Campylocera leptocarpa var. glabella. Buckley’s collection, according to the label, was made in June, 1861 at Brady’s Creek north of Fort Mason, Texas. The identity of Triodanis scabra Raf. cannot be determined with certainty, but on the basis of his description it seems most probable that it is identical with Campylocera leptocarpa Nutt. Rafinesque’s description was as follows: “905. Triodanis scabra Raf. erect rough humble, leaves sessile oblong acute subentire, upper linear; capsules axillary solitary terete curved crowned by 3 subulate teeth—annual, 2 to 4 inches high, seldom with one or two branches, lower leaves broader subcrenate. Found by me 1823 in the glades near the mouth of the Tennessee R. and by Nuttal at Cedar prairies in Arkanzas; but out of 7 specimens not one is in bloom, all are in seeds, probably very early vernal.” The plant found by Rafinesque near the mouth of the Tennessee River hardly can have been the same as Nuttall’s Campylocera leptocarpa, but on the basis of a collection sent by Nuttall to Schweinitz and now in the herbarium at Philadelphia, I suppose Rafinesque’s description to have been drawn up mostly, if not entirely, from Nuttall’s Arkansas material, which is leptocarpa. Nuttall’s personal material of this collection seems not be at Philadelphia and may be at the British Museum; if it can be located, and agrees with the description of Triodanis scabra, I think there can be no doubt that Rafinesque had it in mind (even if not before him!) when writing the description. Schweinitz’ part of Nuttall’s collection is labelled “Red River Nuttall,’ and bears Nuttall’s own ticket with the locality “Ark” and the name ‘“‘Specularia *secaber” in Nuttall’s hand crossed out and another unpublished name substituted for it. The collection consists of a portion of a plant with open flowers, and 6 or 7 plants 7-12 em. high, in fruit, with cleistogamous flowers only; 1 plant is branched slightly. If 7 1945] McVauau: TRIODANIS 51 Nuttall divided his collection into approximately equal parts, as he may well have done, then Rafinesque’s description may well apply to the part kept at Philadelphia, for the reader may note how well it applies at critical points to the plants kept by Schweinitz: upper leaves linear, capsules terete curved crowned by 3 subulate teeth, plants 2-4 inches high, seldom with t or 2 branches, out of 7 specimens not one is in bloom, all are in seeds. Even the name, scabra, seems to have been taken from Nuttall. This species well illustrates a point made earlier in this paper (Recom- mendation 3, p. 15). The mature capsules of the cleistogamous flowers, as first pointed out by Nuttall, are for the most part without definite valves, and are in addition unilocular, with parietal placentation, opening by longitudinal slits in the manner of Downingia, of the LoBELIOIDEAE. Nuttall considered this character of generic value, since no comparable condition is found elsewhere in Specularia (including Triodanis). Certainly the character is unique, but all other characters of 7. leptocarpa are so unquestionably those typical of Triodanis, that it seems clear that T. leptocarpa is biologically a member of the genus but with one strikingly anomalous feature. It is probable that suppression of parts in the cleisto- gamous flowers has gone further in this species than in others, with ac- companying reduction not only in the corolla and androecium, but also in the ovary and accessory parts. DOUBTFUL SPECIES Campanula angulata Raf., Fl. Ludov. 55. 1817. This is evidently either Triodanis biflora or T. perfoliata, but it is impossible from Rafinesque’s description to decide which. Specularia Juliani Batt. in Batt. & Trab., Fl. Alger. Tunis. 222. 1902; Legousia Juliani Briq., Candollea 4: 332. 1931. This is known only from the type collection, which is said to differ from Specularia ( Triodanis) falcata chiefly in having the flowers long-pedunculate. No mention is made, either in Battandier’s original description or in Briquet’s fuller one, of cleistogamous flowers. The plant may be an aberrant form of 7’. falcata, as similarly pedunculate forms of normally sessile-flowered species are found occasionally both in Triodanis and in Campanula. LITERATURE CITED 1. Ascherson, P. [no titles] Bul. Mens. Soc. Linn. Paris 1: 250-251. 21880; Sitzungsber. Ges. Naturforsch. fr. Berl. 1880: 97-108. ?1880. 2. Boissier, E. Flora Orientalis 3: 960. 1875. : 3- Bromfield, W. A. Catalogue of the plants growing wild in Hampshire, ete. Phytologist 3: 529-530. 1840. : 4. Buser, R. Genus Trachelium L. revisum. Bul. Herb. Boiss. 2: 501-532. pl. 15-19. Au 1894. : 5. Darwin, C. The effects of cross and self fertilisation in the vegetable king- dom. pp. 174-176. ed. of 1895. 6. Deam, C. C. Flora of Indiana. pp. 1236. 1940. 52 29. WRIGHTIA [Vou. 1, No. 1 . DeCandolle, A. Monographie des Campanulées. pp. viii, 384. 20 pl. 1830° . Feer, H. Beitrage zur Systematik und Morphologie der Campanulaceen. Engl. Bot. Jahrb. 12: 608-621. 1890. . Fomin, A. Flora Caucasica Critica 4(6): 128. 1906. . Gray, A. Specularia. Proc. Am. Acad. 11: 81-83. 1876. . Syn. Fl. N. Am. 2(1): 9-10. 1878. . Holzinger, J. M. List of plants collected by C. 8. Sheldon and M. A. Carleton in Indian Territory in 1891. Contr. U. 8. Nat. Herb. 1: 189- 219. 12D 1802. . Hooker, J. D., and T. Thomson. Praecursores ad floram indicam. Journ. Linn. Soe. 2: 1-29. 1858. . Knuth, P. Handbuch der Bliitenbiologie 1: 65-66. 1898. . Linnaeus, C. Hortus Upsaliensis 40. 1748. . Species Plantarum 163-171. 1753. . ——. Genera Plantarum ed. 5. 77, 78. 1754. . ——. Amoenitates Academicae 3: 396. 1756. . Praelectiones in Ordines Naturales Plantarum. P. D. Giseke, ed. 399. 1792. . MeVaugh, R. A new name for Heterocodon rariflorum Nutt. Leafl. West. Bot. 3: 48. 1941. . Nuttall, T. Descriptions and notices of new or rare plants in the natural orders Lobeliaceae, Campanulaceae . . . etc. Trans. Am. Phil. Soc. n. s. 8: 251-272. 1843. . (Oliver, D.] [In review of paper by C. Darwin] Nat. Hist. Rev. n. s. 2: 240-241. 1862. . Post, G. E. Flora of Syria, Palestine and Sinai. ed. 2, v. 2, pp. xviii, 928. 1933- . Ritzerow, H. Uber Bau und Befruchtung kleistogamer Bliiten. Flora 98: 163-212. 1907. . Torrey, J. A flora of the State of New-York. 2 vols., 1843. . Trent, J. A. Floral variations in Specularia perfoliata (L.) A. DC. Am. Midl. Nat. 23: 448-454. 19408. . Flowering behavior of Specularia perfoliata in relation to light in- tensity and light duration. Trans. Kans. Acad. 43: 199-205. 1940b. . Studies pertaining to the life history of Specularia perfoliata (L.) A. DC., with special reference to cleistogamy. Trans. Kans. Acad. 45: 152-164. 1942. VonMohl, H. Einige Beobachtungen iiber dimorphe Bliithen. Bot. Zeit. 21: 309-315, 321-328. 1863. Division of Plant Exploration and Introduction U. 8. Department of Agriculture Plant Industry Station, Beltsville, Maryland dg enna 13 Sean aGlO caacaSeh aca Sircid ah he ERO MPA MSR or Ee NEW PHANEROGAMS FROM TEXAS, MEXICO AND CENTRAL AMERICA Cyrus LonGwortH LUNDELL The wartime search for new sources of plant products to substitute for materials formerly obtained from Asiatic and other areas no longer ac- cessible has brought about a long overdue examination of the plant re- sources of the Western Hemisphere. In spite of the accelerated program of exploration, only the surface has been scratched. One of the most fertile fields open to scientists is the study of our rich native flora with a view to discovering new plant products for industrial and other uses. In cooperation with the Foreign Economie Administration of the U. 8S. Government in its search for new sources of rotenone in Mexico and Central America, studies of Lonchocarpus and related genera have been undertaken. In this paper, three new species of Lonchocarpus are described. Obviously the genus has been neglected by field botanists, for at least a third of the collections received from Mexico and Central America repre- sent rare or unrecognized species. Whenever a genus assumes economic importance, these same deficiencies in exploration become apparent, a clear indication of our imperfect knowledge of the American flora. In continuation of taxonomic studies, seven other new species are pro- posed in the genera Cryosophila, Abronia, Bernardia, Zuelania, Parathesis, Brazoria and Scutellaria. Hypericum aphyllum Lundell is transferred to the genus Sarothra. Cryosophila bifurcata Lundell, sp. nov.—Arbor gracilis, caudice ca. to em. diam., spinis rhizoideis pungentibus armato. Folia petiolata; ligula crassa triangula; lamina 80 em. longa, segmentis ad 3.5 cm. latis, apice breviter bifidis, subfaleatis, subtus argenteo-pubescentibus. In- florescentia laxe ramosa. Flores glabri, ca. 3 mm. longi. Calycis segmenta carnosa, basi connata, petalis vix aequilonga. Petala imbricata, libera. Trunk slender, ro em. in diameter, about 8 m. high, armed with spines. Leaves palmate, the petioles slender; ligule produced, triangular; blades up to 80 em. long, multicostate, the segments parted to or below the middle, up to 3.5 em. wide, tapering to a bifurcate apex, margin entire, midrib strong and paralleled by numerous fine veins, silvery-sericeous on the undersurface, the hairs closely appressed. Inflorescence about 60 cm. long, downward-curved, sheaths white tomentose at first, 15-23 cm. long; rachis stout, tomentose at first, side branches up to 12 cm. long, bearing the floriferous branches 3-6 em. long, the flowers sessile, dense. Flowers at anthesis about 3 mm. long, ellipsoid or nearly globular, bearing 6 de- flexed anthers at apex. Perianth of 6 parts in two series, the calyx lobes 53 54 WRIGHTIA [Vou. 1, No. 1 connate at base with conspicuous obtuse sinuses between them, petals equaling calyx lobes, not connate at base, imbricate. Filaments connate. Ovaries 3, each with long exserted style recurved at apex. Immature fruits depressed-globose. vBRITISH HONDURAS: Toledo District, Punta Gorda-San Antonio Road, near Jacinto Creek, in high ridge, November 14, 1944, Percy H. Gentle 4972 (type in the herbarium of Southern Methodist University), a tall palm about 25 feet high and 4 inches in diameter, trunk covered with spines; San Antonio, Moffredge Creek, in wild coffee ridge, December 9, 1944, Gentle 5063; vernacular name, ‘‘give-and-take.”’ The leaf segments with bifurcate apex suggest an affinity to Cryosophila Cookii Bartlett, but the inflorescence of C. bifurcata is altogether different, being paniculiform. In C. Cook7i the inflorescence consists of short simple branchlets fastigiate in a densely crowded interrupted spiral. A. bifurcata is near C. Warscewiczii as interpreted by Bailey in Gentes Herbarum 3: rog-112 (1933). The narrower leaf segments, flowers only 3 mm. long at anthesis, and imbricate petals free to base are differences by which C. bifurcata may be recognized. The very immature fruits from the British Honduran tree are depressed-globose, while fruits of C. War- scewiczii are oblong-pyriform. Bartlett in Carnegie Inst. Wash. Publ. 461: 39 (1935) described C. Warscewiczii as having leaves with the segments obliquely and coarsely cross-veined, a condition not evident in C. bifurcata. , Abronia Ameliae Lundell, sp. nov.—Herba, ad 60 em. alta, glanduloso- villosa. Folia petiolata, petiolo 1-8.5 em. longo, lamina ovata, elliptica vel rotundata, 3-8 em. longa, 2-6 em. lata, apice rotundata, basi subcordata vel subtruncata, margine sinuata, viscida. Bracteae 5-7, ad 16 mm. longae, to mm. latae. Flores 18-25 mm. longi, glanduloso-villosi. Fructus turbinatus, 5-alatus, 7-9 mm. longus, 3~4.5 mm. latus. Coarse spreading perennial herb up to 60 em. high, viscid throughout, the stems thick, villous, the hairs variable in length and gland-tipped. Leaves petiolate, the petioles 1-8.5 em. long, the lower often exceeding the blades; leaf blades fleshy, drying chartaceous and brittle, concolor, ovate, elliptic, suborbicular or depressed-orbicular, 3-8 em. long, 2-6 cm. wide, apex rounded, base subcordate or subtruncate and decurrent, margiD conspicuously sinuate, rather sparsely viscid pubescent on both surfaces with short hairs, costa and veins nearly plane, the primary lateral veins usually 3 on each side. Peduncles rather slender, viscid-puberulent, 2.5—12 em. long. Bracts 5~7, pale greenish to orchid, thin, oblong, elliptic or obo- vate, 10-16 mm. long, 5-10 mm. wide, apex broadly rounded and acutish, viscid-villous externally. Flowers numerous, forming head 5 em. in diameter. Perianth orchid (tube and limb), 18-25 mm. long, viscid-villous, the tube slender, the limb 1o mm. wide. Stamens 5s, included. Fruits narrowly turbinate, 7-9 mm. long, 3—4.5 mm. wide at apex, s-winged, not narrowed at apex, attenuate to the base, apex rounded or truncate, reticulate-veined, glabrous or with a few scattered hairs at apex only. Seeds oblanceolate- oblong, about 2.5 mm. long. Bed a RTs BR Ee al RE EE ae 1945] LUNDELL: NEw PHANEROGAMS 55 TEXAS: Brooks County, north of Encino, in sandy live oak belt, May ro, 1940, C. L. Lundell & Amelia A. Lundell 8832; 14 miles south of Falfurrias, in sandy live oak belt, March 19, 1942, Lundell & Lundell 10816 (type in the herbarium of Southern Methodist University); north of Encino, along sandy roadside, April 9, 1944, Lundell & Lundell 12807. Although long confused with A. fragrans Nutt., to which it has close affinity, A. Amelzae differs in its glabrous narrowly turbinate fruits widest at the rounded apex, orchid colored perianth, and bracts rounded at apex. The fruits of A. fragrans are short-villous or puberulent, usually biturbin- ate, and narrowed at apex. This fine plant is worthy of widespread cultivation for ornamental pur- poses. In Brooks County it is planted in gardens. Lonchocarpus belizensis Lundell, sp. nov.—Arbor, 15 em. diam.; ramulis striatis, minute subadpresse puberulis. Folia 5—o-foliolata. Foliola membranacea, oblanceolato-oblonga vel anguste oblongo-elliptica, 4~-9.5 em. longa, 1.5—3.3 em. lata, apice acuminata, basi rotundata, supra minute subadpresse puberula, subtus subglauca, strigillosa. Paniculae terminales. Pedicelli ad 2 mm. longi. Calyx cupulatus, 3 mm. longus, 5-dentatus. Petala atro-rubra. Ovarium strigillosum, 4-ovulatum. Tree, 15 em. in diameter, twigs striate, pubescent with minute sub- appressed reddish-brown hairs. Leaves 5—9-foliolate, the rachis subterete, up to 7 em. long, pubescent with minute appressed golden-brown hairs. Stipules deciduous early. Leaflets membranaceous; petiolules pubescent, 3-4 mm. long; blades oblanceolate-oblong or narrowly oblong-elliptic, 4-9.5 em. long, 1.5—3.3 em. wide, apex short acuminate, base rounded, the sides subequal, paler beneath, the upper surface pubescent with minute subappressed golden-brown hairs, the undersurface subglaucous, strigil- lose, primary veins 7—9 on each side, plane on upper surface, prominent on undersurface. Racemes forming a large open terminal panicle, the lower ones axillary, up to 18 em. long, pubescent with minute subappressed golden-brown hairs. Peduncles pubescent, 2-4-flowered, sometimes branched, 1.5—3 mm. long. Pedicels up to 2 mm. long. Calyx pubescent with minute appressed golden-brown hairs, cupulate, 3 mm. long, 5-den- tate. Petals dark red; standard densely strigillose, suborbicular, the blade 9.5 mm. wide, ro mm. long, base subcordate, the claw 1.5 mm. long; wings glabrous, the blade rounded-auriculate at base, 9 mm. long, 3 mm. wide, the claw 2 mm. long; carinal petals cohering above middle, ciliate at base. Vexillar stamen free at base. Ovary silvery-strigillose, 4-ovulate; style arcuate, hirsute. BRITISH HONDURAS: Toledo District, upper reach of Golden Stream in cohune ridge, May 11, 1944, Percy H. Gentle 4600 (type in the herbarium of Southern Methodist University), tree, 6 inches in diameter, flowers dark red. The species is allied to L. latifolius (Willd.) H.B.K. Lonchocarpus cruentus Lundell, sp. nov.—Arbor, 10-12 m. alta, 28 em. diam.; ramulis parce strigillosis. Folia 9—13-foliolata. Foliola chartacea, 56 WRIGHTIA [Von. 1, No.2 oblonga, oblongo-elliptica vel oblanceolata, 3-7.5 cm. longa, 1.6-3 cm. lata, apice abrupte acuminata, basi acutiuscula, subtus parce strigillosa. Inflorescentiae racemosae, axillares. Pedicelli 1-1.5 mm. longi. Calyx strigosus, cupulatus, 3 mm. longus. Petala atro-purpurea. Ovarium sericeum, 5—6-ovulatum. Tree, 10-12 m. tall, 28 em. diam., branches dark brown, conspicuously lenticellate, sparsely strigillose with short brown hairs. Leaves 9~13- foliolate, the rachis strigillose, up to 9 em. long, canaliculate. Stipules deciduous early. Leaflets chartaceous; petiolules strigillose, 3.5—4.5 mm. long; blades oblong, oblong-elliptic or oblanceolate, 3—7.5 cm. long, 1.6~3 em. wide, apex abruptly acuminate, the acumen obtusish, base acutish, persistently hirtellous along the veins on upper surface, the hairs sub- appressed except along the costa, entire undersurface sparsely strigillose, primary veins 7~11 on each side, slightly impressed above, conspicuous on undersurface. Racemes solitary in the axils of the upper leaves, stout, 5.5— 11.5 em. long, pubescent with short subappressed ferruginous hairs. Peduneles and pedicels ferruginous-pubescent, the peduncles solitary, 2- flowered, 2—2.5 mm. long, the pedicels 1-1.5 mm. long. Calyx strigose with brownish hairs, cupulate, 3 mm. long, the margin subentire. Petals dark reddish-purple; standard sericeous, suborbicular, biauriculate and callous at base, emarginate at apex, the claw about 2.3 mm. long, the blade 10o~11 mm. long, 12-13 mm. wide; wings silky-pubescent without, auriculate at base, the claw about 4 mm. long, the blade about 9 mm. long, 4.3 mm. wide; carinal petals rounded at base, silky-pubescent. Vexillar stamen free at base. Ovary sericeous, 5—6-ovulate; style arcuate, hairy at base only. MEXICO: Tazasco, southeast of Macuspana, between Macuspana and Cerro del Tortugero, September 30, 1944, C. L. Gilly, Sr. & Efraim Hernandez Xolocotzi 411 (type in the herbarium of Southern Methodist University); vernacular name, ‘‘palo de sangre.” The collectors state that the bark yields a reddish fluid when cut, and that this fluid coagulates into gummy resin which becomes glossy hard within a few hours. In the absence of fruits, the relationship of the species is doubtful, but it appears to have affinity to L. sericeus (Poir.) H.B.K. The latter has leaves with veins strongly impressed on the upper surface. Lonchocarpus Gillyi Lundell, sp. nov—Arbor, 4-5 m. alta, 20-25 cm. diam.; ramulis striatis. Folia 8—r1o-foliolata. Stipulae s—8 mm. longae. Foliola subcoriacea, oblonga vel oblanceolata, 1.8—4 em. longa, o.6—-1.4 cm. lata, apice rotundata, basi obtusa, costa et nervis impressis. Inflorescentiae racemosae, axillares. Pedicelli fructiferi 2-3 mm. longi. Calyx campanula- tus, ca. 2.5 mm. longus, dentatus. Legumen 1—8-spermum. Tree 4-5 m. tall, 20-25 em. in diameter; twigs rather stout, striate, sparsely pubescent with spreading hairs. Leaves 8—10-foliolate, the rachis pubescent with brown spreading hairs, the rachis and petiole together 8-11 cm. long. Stipules attenuate, 5-8 mm. long. Leaflets firm, subcoriace- ous; petiolules pubescent, about 2 mm. long; blades of lateral leaflets 1945] LuNDELL: New PHANEROGAMS 57 oblong, the terminal oblanceolate, 1.8-4 em. long, 0.6-1.4 em. wide, apex rounded or inconspicuously emarginate, base obtuse, the sides inequi- lateral, pubescent on undersurface with loose spreading reddish-brown hairs, the hairs fewer and shorter on upper surface, lateral veins 9-12 on each side, not strongly ascending, impressed on upper surface, conspicuous on lower surface, reticulate. Racemes axillary, borne on older wood, 3~9 em. long, rather densely pubescent with shaggy reddish-brown hairs. Pedicels of fruits 2-3 mm. long. Persistent calyx campanulate, about 2.5 mm. long, toothed, densely strigose with reddish-brown hairs. Legume thin, elongate, narrowed to base, pubescent with appressed reddish-brown hairs, 1-8-seeded, the immature legumes up to 12 em. long, 1.8 em. wide. * MEXICO: Cutapas, hills south of Salto de Agua, about 2 kilometers from Salto de Agua on trail to Yajalon, October 4, 1944, C. L. Gilly, Sr. & Efraim Hernandez Xolocotzi 430 (type in the herbarium of Southern Methodist Uni- versity); vernacular name, ‘“‘biche rayo.” Although closely resembling L. apricus Lundell, it is separable at once from that species by its smaller leaves and spreading reddish-brown pubescence. The affinity of L. Gillyi to L. rugosus Benth. is evident, but the latter has larger leaves usually elliptic-oblong rather than strictly oblong. In the material of L. Gillyi at hand, the racemes are borne on old wood, a noteworthy characteristic. Bernardia aurantiaca Lundell, sp. nov.—Arbor parva, 10 m. alta, ramis ramulisque parce stellato-puberulis. Folia petiolata, petiolis ad 3.5 cm. longis, stellato-puberulis; lamina lanceolata, 4-12 em. longa, 1-4 em. lata, apice acuminata, basi obtusa vel acutiuscula, crenulato-serrulata, utrinque parce stellato-puberula. Flores dioeci. Inflorescentiae co racemosae, ad 14 em. longae, stellato-puberulae; bracteis late cordato-ovatis, ca. 2 mm. longis, 17~20-floris; pedicellis 2-4 mm. longis. Sepala 1.2-1.5 mm. longa. Stamina ro vel rr. A slender tree, 10 m. high, 10 em. in diameter, the cambium bright orange; branchlets and twigs obscurely angled, rather sparsely puberulent with stellate hairs. Petioles slender, up to 3.5 em. long, stellate-puberulent. Leaf blades slightly paler beneath, lanceolate, 4-12 em. long, 14 em. wide, apex acuminate, base narrowed, obtuse or acutish, margin remotely crenulate-serrulate, sparsely stellate-puberulent on both surfaces, the in- dument persistent only on the veins at maturity, costa and primary veins conspicuous on lower surface, the primary veins 5 or 6 on each side. Stam- inate inflorescence slender, unbranched, up to 14 em. long, stellate-pu- berulent; bracts broadly cordate-ovate, about 2 mm. long, obtusish, puberulent; flowers 17-20 in basal bracts, pedicels slender, 2~4 mm. long, jointed near or at the base. Sepals 3, elliptic, 1.2-1.5 mm. long, acute, with a few hairs on the outer surface. Stamens 10 or 11, radiating; filaments about 1 mm. long, glabrous. Pistillate flowers and fruits unknown. “BRITISH HONDURAS: El Cayo District, Chalillo Crossing, in ad- vanced forest on hillside, July 15, 1936, C. L. Lundell 6509 (type in the her- barium of Southern Methodist University). 58 WRIGHTIA [Vou. 1, No. 1 This collection, originally distributed as a new species of Acalypha by P. C. Standley, was referred by the writer to B. interrupta (Schl.) Muell. Arg., Contrib. Univ. Mich. Herb. 4: 12 (1940). Subsequent field studies in 1943 of B. interrupta in Hidalgo and San Luis Potosf have convinced the writer that the British Honduran tree should be segregated as a distinct species. B. aurantiaca, notable for its orange colored cambium, has smaller acuminate leaves, and smaller flowers twice as numerous in each bract. The indument of B. interrupta is distinctly coarser. Sarothra aphylla (Lundell) Lundell, comb. nov. Hypericum aphyllum Lundell, Am. Midl. Nat. 29: 477. 1943. Zuelania belizensis Lundell, sp. nov.—Arbor, ramulis glabris. Folia glabra, petiolata, petiolis o.9—2 em. longis; lamina subchartacea, oblanceo- lata, oblongo-oblanceolata vel oblongo-elliptica, 9-25 em. longa, 4-9 cm. lata, apice acuminata, basi rotundata. Inflorescentiae terminales, multi- florae, pedunculatae, ad 3 em. longae. Pedicelli glabri, ad 1.5 cm. longi. Calyx 5-6 mm. longus, lobis ciliatis. Stamina 14, filamentis et antheris pubescentibus. A small tree or woody vine (?); twigs elongate, glabrous, rather stout. Leaves glabrous, large; petioles canaliculate, o.g—-2 em. long; leaf blades subchartaceous, oblanceolate, oblong-oblanceolate or oblong-elliptic, 9-25 em. long, 4-9 em. wide, apex abruptly acuminate, base rounded, the sides unequal or subequal, glabrous on both surfaces, often drying reddish, shining above, pellucid-punctate, margin inconspicuously and remotely serrulate-crenulate, reticulate-veined, costa impressed above, prominent on undersurface, the primary veins 8-12 on each side, slender but conspicu- ous. Inflorescences borne at the apical nodes, many-flowered, pedunculate, up to 3 cm. long, conspicuously bracteate; peduncles short, up to 6 mm. long, glabrous; bracts and bractlets ciliate. Pedicels slender, glabrous, up to 1.5 em. long. Calyx 5—6 mm. long, the 5 lobes ciliate, elliptic, unequal. Stamens 14, filaments and anthers pubescent; appendages of the disk alternating with the stamens, a third shorter than the filaments, pubescent, Ovary pubescent; stigma capitate. Fruits fleshy, about 3 em. in diameter , BRITISH HONDURAS: Toledo District, Bolo Camp, upper reach of Golden Stream, in high ridge, April 15, 1944, Percy H. Gentle 4537 (type in the herbarium of Southern Methodist University), woody vine, flowers whitish; Rio Grande, in wild coffee ridge, July 7, ro44, Gentle 4696, tree, 6 in. diam., bark grayish, wood creamish. The common species of Mexico and Central America, Z. Guidonia (Swartz) Britt. & Millsp., has altogether different leaves densely soft- pubescent on the undersurface. Parathesis membranacea Lundell, sp. noy.—Arbor parva, ramis ramulisque brunneo-lepidotis. Folia petiolata, petiolis 1-2 em. longis, lepidotis; lamina membranacea, elliptica vel obovato-elliptica, 8.5-15 em. longa, 4.5-7.5 em. lata, apice abrupte acuminata, supra glabra, subtus stellato-lepidota, crenulato-dentata. Inflorescentiae terminales, anguste paniculatae, brunneo-lepidotae, 8-10 em. longae. Flores corymbosi, brun- 1945] LuNDELL: New PHANEROGAMS 59 neo-lepidoti. Pedicelli ad 4 mm. longi. Sepala 1.3~1.5 mm. longa, acuta. Petala anguste lanceolata, ca. 5.5 mm. longa. Antherae 2.2 mm. longae, punctatae. A small tree; branches rather stout, persistently lepidote; twigs brown lepidote. Petioles 1-2 em. long, canaliculate, lepidote. Leaf blades mem- branaceous, elliptic or obovate-elliptic, 8.5—15 em. long, 4.5—7.5 em. wide, apex abruptly acuminate, base decurrent on the petiole, stellate-lepidote on the undersurface at first, essentially glabrous with age, glabrous on upper surface, black punctate, finely reticulate-veined, primary veins 12— 15 on each side, conspicuous on undersurface, plane or slightly impressed on upper surface, margin conspicuously crenulate-dentate almost to base. Inflorescence leafy, terminal, narrowly paniculate, brown lepidote, 8-10 em. long. Flowers corymbose, brown lepidote, borne on pedicels up to 4 mm. long; buds 4 mm. long at anthesis. Sepals narrowly triangular, 1.3-1.5 mm. long, acute. Petals about 5.5 mm. long, coherent at base only, narrowly lanceolate, attenuate to the apex, punctate. Filaments stout, about 1.3 mm. long. Anthers conspicuously black punctate, ovate-lanceo- late, 2.2 mm. long, apiculate. Ovary with a few hairs at apex, glabrous otherwise. Style 3.5 mm. long, glabrous. ° BRITISH HONDURAS: Tea Kettle, May 12, 1931, H. H. Bartlett 13140 (type in the herbarium of Southern Methodist University), a small tree, flowers dull pink. P. membranacea is separable from P. serrulata (Swartz) Mez, to which it is closely allied, by the broadly elliptic or obovate-elliptic membranaceous leaves with conspicuously dentate margin. Further, P. membranacea has smaller inflorescences and flowers. In superficial aspects, the species re- sembles P. pleurobotryosa Donn. Sm. ‘ Brazoria arenaria Lundell, sp. nov.—Herba, 40 cm. alta, ramulis pu- bescentibus. Folia membranacea, oblongo-elliptica, oblanceolato-elliptica, vel lineari-oblonga, usque ad 12 em. longa, 3.5 em. lata, apice rotundata, basi attenuata, denticulata. Inflorescentiae racemosae, ad 30 cm. longae. Calyx campanulatus, 5.5; mm. longus. Corolla 1.3-1.8 em. longa. Annual, up to 4o em. high, branching at the base, stems erect or ascend- ing, pubescent with short hairs. Leaves membranaceous, the basal petiolate, oblong-elliptic or oblanceolate-elliptic, up to 12 em. long, 3.5 em. wide, rounded at apex, attenuate at base, denticulate, undersurface glabrous, upper surface with a few short hairs along midrib at base of blade, the petiole often subequaling the blade; cauline leaves few, remote, linear- oblong, oblanceolate-oblong or obovate-oblong, sessile, reduced, those be- low inflorescence bract-like, often partly clasping, denticulate. Racemes up to 30 em. long, interrupted, the flowers rather remote, pubescent, the hairs short with glandular ones intermixed. Bracts broadly elliptic or ovate- elliptic, about 3.5~s; mm. long, acute. Calyx campanulate, 5.5 mm. long, becoming 9 mm. long in fruit, the lips subequal; tube pubescent with short hairs, some glandular, not bearded, the upper lip of 3 broad rounded lobes, the lateral ones apiculate, the middle one rounded, the lower lip slightly 60 WRIGHTIA [Vou. 1, News longer, shallowly 2-lobed, the lobes sharply dentate with 3-5 short subulate teeth. Corolla lavender with maculate throat, sparsely puberulent along back, 1.3-1.8 em. long, tube dilated upward, the upper lip with 2 oblong subentire (not toothed) lobes, the lower lip with 3 smaller lobes which are emarginate at apex. Nutlets pubescent, 2 mm. in diameter. ” TEXAS: Brooks County, off U. S. highway 281, south of Encino, on sandy plains, April 6, 1944, C. L. Lundell & Amelia A. Lundell 12766 (type in the herbarium of Southern Methodist University). In characteristics of the calyx and corolla, B. arenaria approaches B. truncata (Benth.) Engelm. & Gray. The calyx of B. arenaria differs in not having a viscid-villous beard, and in having unequal lips with the middle lobe of the upper lip rounded, not apiculate. The upper lip of the corolla in B. arenaria has subentire lobes, while the lobes in B. truncata are distinctly toothed. In habit there is a broad divergence between the two species. In B. arenaria the cauline leaves are reduced and bract-like below the inflores- cence; in the other species the cauline leaves are large and well developed to the base of the raceme. In B. truncata the racemes are short and densely flowered while in B. arenaria they are interrupted their entire length. Scutellaria petenensis Lundell, sp. nov.—Herba, 20-40 cm. alta, ramulis gracilibus, pubescentibus. Folia petiolata, petiolis ad 5 mm. longis; lamina late ovata vel rhomboideo-ovata, parva, raro ad 1.7 em. longa, 1.6 em. lata, apice rotundata, basi acuta, crenata. Flores axillares vel breviter racemosi. Pedicelli ca. 4 mm. longi. Calyx pubescentibus, 2-3 mm. longus. Corolla 10-11 mm. longa. A low spreading herb, 20-40 em. high, the stems slender and weak, pubescent with short recurved hairs. Petioles short, slender, up to 5 mm. long, pubescent. Leaf blades broadly ovate or rhomboid-ovate, usually very small, sometimes up to 1.7 em. long, 1.6 em. wide, apex rounded, base abruptly acute, margin conspicuously crenate with 3 or 4 teeth on each side, basal third entire, sparsely bairy on both surfaces, the hairs coarser and denser along the veins on the undersurface. Flowers axillary, or borne in short racemes about 2 em. long. Pedicels slender, pubescent, about 4 mm. long, bearing two filiform bracts near the base. Calyx pubescent, 2~3 mm. long at anthesis. 3.5-4 mm. long at maturity. Corolla blue, the tube and galea 1o-11 mm. long. Lower stamens seated about 5 mm. above base of tube. * GUATEMALA: Department of Petén, Chimah Savanna near La Libertad, in open grassland, June 4, 1933, C. L. Lundell 3614 (type in the herbarium of Southern Methodist University), flowers blue. This herb has the habit suggesting S. havanensis Jacq. and S. Gaumert Leonard. In its smaller flowers with galea and tube of corolla ro to 11 mm. long, and lower stamens seated only 5 mm. above the base of the tube, it differs from both of these species. The relationships of S. petenensis have been pointed out by Epling in Carnegie Inst. Wash. Publ. 522: 230 (1949); and in Univ. Calif. Publ. Bot. 20: 109 (1942). NOTES Charles Wright, immortalized in the plant names of the state, was the foremost botanical explorer of early Texas. His portrait of unknown date, reproduced as the frontispiece, was supplied by the Gray Herbarium of Harvard University through the courtesy of Mr. C. A. Weatherby. NEWS On February 1, 1945, Dr. Lloyd H. Shinners was appointed Research Fellow in the Institute of Technology and Plant Industry to undertake studies upon the Composirak of Texas. Dr. Shinners received his Ph. D. degree from the University of Wisconsin under Dr. N. C. Fassett in 1943, his thesis being ‘““The Grasses of Wisconsin.” In 1943-1944 he was Assitant in the Department of Soils of the University of Wisconsin, and later Bot- anist of the Whitnall Arboretum in Milwaukee. Since 1944 he has been a Research Associate in Botany of the Milwaukee Public Museum. In addi- tion to his work upon the grasses of Wisconsin, Dr. Shinners has published various papers on the genera Aster and Liatris in North America. Dr. Rogers MeVaugh of the Division of Plant Exploration and Intro- duction of the U. S. Department of Agriculture is spending three months, May through July, in Texas carrying out field studies upon the RosacEak of the state. The project is sponsored jointly by the Institute of Technology and Plant Industry and the U. 8. Department of Agriculture. Dr. C. H. Muller, formerly Associate Botanist of the Bureau of Plant Industry of the U. 8. Department of Agriculture, and recently engaged in wartime research upon guayule under Dr. A. C. Hildreth, has been ap- pointed Research Fellow in the Institute of Technology and Plant Industry for three months, June through August, 1945. Dr. Muller has resumed his field work upon the oaks of Texas, and in addition is carrying out ecological studies on the vegetation. 61 I , j , WRIGHTIA ok BoraNicaL JOURNAL : WRIGHTIA : EDITOR Cyrus Loneworta LunpEL ao ae Each westanie will contait ssuec at Sregular intervals, each number to WRIGHTIA : VOLUME 1 APRIL, 1946 NUMBER 2 REVISION OF THE GENUS CHAETOPAPPA DC.1 Luoyp H. SHINNERS INTRODUCTION The taxonomy of the Compositae has suffered greatly from excessive reliance on the nature of the pappus, or its presence or absence, as a character of generic value. On this sole basis, species obviously very similar, and differing in no other significant respect, have been regarded as con- stituting separate genera. Serinia has been thus arbitrarily separated from Krigia, Brachychaeta from Solidago, Amphiachyris from Gutierrezia, Eremiastrum from Monoptilon, Achaetogeron from Erigeron. Conversely, some species have been detached from their near congeners and placed with species to which they have much less resemblance, on little more grounds than similarity of pappus. Aster has been made to include Leuce- lene, which in structure of florets and phyllaries (involucral bracts), and vegetative parts, is much closer to Chaetopappa, and Brachyactis, which is hearer to Conyza; Bellis has been made to include Astranthium, which is closer to Dichaetophora; Solidago to include Euthamia, which is nearer to Gutierrezia, The previously recognized species here included in Chaetopappa have been placed not only in separate genera, but in separate subtribes, largely on the basis of presence or absence of pappus. In the type species of Chaeto- pappa (ASTEREAE-ASTERINAB), C. asteroides (Nutt.) DC., the pappus nor- mally consists of five well-developed hyaline scales and as many bristle-like awns. But var. imberbis A. Gray was described as lacking the awns. In addition, the seales vary greatly in length, being sometimes hardly more than vestigial in the species proper; and the variations may be similar or may differ greatly between disk and ray florets. Bourdonia bellidzfolia (Gray & Engelm.) Greene ( Keerlia bellidifolia Gray & Engelm.), which in the field looks exactly like a robust form of Chaetopappa asteroides, was placed in another genus and subtribe (ASTEREAE-BELLIDINAE) because of the absence of a pappus. But the summit of the achene bears a thickened ‘ The five revisions which follow have benefited from the work of many more hands and minds than the author’s. I am grateful to Dr. Eula Whitehouse and Mr. V. L. ry, upon whose wide and intimate knowledge of Texas plants and Texas places I have drawn many times; to Dr. S. W. Geiser for great and constant help in matters relating to botanical collectors in the Southwest and Mexico; to Mr. H. B. Parks, 64 WRIGHTIA [Vou. 1, No. 2 ring, and may rarely show a vestigial scaly crown. The second species of Bourdonia, B. effusa (A. Gray) Greene, regularly has a crown of very short hyaline scales, like that sometimes found in Chaetopappa asteroides. The genus Diplostelma, differing from Chaetopappa chiefly in the more numerous florets and 2-nerved achenes, was described by Asa Gray with the observation that its single species, D. bellioides, might better be trans- ferred to Chaetopappa. This was later done by him, but he adopted the name of De Candolle’s Distasis modesta, which he mistook for the same plant. Fragmentary and immature specimens of the two are very similar in appearance, but De Candolle’s plant was a closer congener of Chaeto- pappa asteroides, differing from Gray’s in the less numerous florets and several-nerved achenes. De Candolle was doubtful as to the affinities of his genus Distasis, having been unable to determine that the anthers were not caudate. He curiously fails to compare it with his own Chaetopappa, though his descriptions of the two genera are remarkably similar, and give no marked differences between them. The plants of these three groups (Bouwrdonia, Chaetopappa, and, by error, Distasis) closely resemble each other in details of floral structure, and in form and texture of phyllaries. Ignoring the fickle and wholly unreliable character of the pappus, the eight species now known fall nat- urally into four series on the basis of shape of involucre, number of florets, nature of the achenes, duration of plant, foliage, and habit and general appearance. These series do not correspond to any of the genera in which the species were described, being based on similarities in a number of characters instead of the single, arbitrary, inconstant one of the pappus. They are here adopted as subdivisions of a single amplified genus. curator, and Mrs. J. J. Taubenhaus, of the Tracy Herbarium, to Dr. B. C. Tharp, director, Dr. Fred Barkley, curator, and Mr. Barton H. Warnock, of the University of Texas Herbarium, all of whom have borne cheerfully the double. imposition of de- mands in personal visits and in correspondence; to those in charge of the herbaria given in the list of abbreviations following, who responded readily and generously to requests for large loans, including many types and isotypes; and to the librarians of F ondren Library of Southern Methodist University, the Gray Herbarium, and the Missour! Botanical Garden, in particular to Miss Nell C. Horner and Miss E. Mepham of the last institution, for their part in furthering the large and critical portion of my work which was almost exclusively bibliographical. Especial thanks are due Dr. E. D. Mer- rill, of the Arnold Arboretum, who responded promptly and generously to a request for assistance with Rafinesque references by sending his personal notes, abstracts, tracings, and parts of the manuscript of his forthcoming Index Rafinesquianus. : The location of specimens cited in the following revisions is indicated by abbrevl- ations of herbaria, as follows: University of California, Berkeley (C); Herbarium ° I. W. Clokey, University of California, Berkeley (C-Cl); Dudley Herbarium, Stanford University (D); Gray Herbarium, Harvard University (G); University of Illinois, Urbana (Ill); Missouri Botanical Garden, St. Louis (Mo); New York Botanical Garden, New York (NY); Academy of Natural Sciences, Philadelphia (P); Rocky Mountain Herbarium, University of Wyoming (RM); Southern Methodist University, Dallas (SMU); University of Texas, Austin (T); Tracy Herbarium, Texas A. & M. College, College Station (TAM); United States National Herbarium (US); United States National Aboretum (USNA); West Virginia University, Morgantown (WV). 1946| SHINNERS: CHAETOPAPPA 65 SYSTEMATIC TREATMENT CuHaEToPAPPA DC., Prodromus 5: 301. 1836. Chaetophora Nutt., Journ. Phila. Acad. 7: 111. 1834. (Printed as Chaetanthera, corrected in erratum slip to Chaelophora.) Not Chaetophora Schrank, 1780. Distasis DC., Prodr. 5: 279. 1836. (This genus first merged with Chaetopappa, under the latter name, by Gray, Proc. Amer. Acad. 16: 82. 1880.) Diplostelma Raf., New Fl. N. A. 2: 44. 1837 (dated 1836). Diplostelma A. Gray, Plantae Fendlerianae (Mem. Amer. Acad. n.s. 4): 72-73. 1849. (Rafinesque’s name used for a professedly new genus, on the grounds that fewer synonyms would be created if it also proved to be assignable to Chaetopappa.) Keerlia A. Gray, Plantae Wrightianae (Smithsonian Contr. 3): 92. 1852. Not Keerlia DC., Prodr. 5: 309-310. 1836. (Gray transmogrified De Candolle’s genus, and after all the original species had been placed elsewhere, took it over as his own. In Plantae Lindheimerianae II (Boston Journ. Nat. Hist. 6): 221, 1850, where Keerlia is used essentially in its new form, it is still attributed to De Candolle.) Bourdonia Greene, Erythea 1: 207. 1893. ComposirTak, tribe ASTEREAR, subtribe ASTERINAE. Plants annual with taproots, or perennial and either tufted from a vertically forking rootstock or caudex, or forming loose tufts from super- ficial slender rhizomes. Leaves alternate, simple, entire, sessile or with narrowed petiole-like bases. Inflorescence (when of more than a single head) eymose, the first head to flower terminating the main axis, the later ones (often much overtopping it) on lateral branches, solitary and terminal on the branches or branchlets. Involucre hemispherical to subeylindrical, of elliptic to linear-lanceolate, stiff, chartaceous, green (sometimes purple to red brown), scarious-margined phyllaries, appressed and imbricated in about 2-6 series, the outermost not more than half as long as the inner- most, all often concave, especially in drying, and appearing broadly keeled. Receptacle flat or slightly convex, naked, roughened by the points of at- tachment of the achenes. Heads heterogamous, radiate, heterochromous, few to many flowered. Ray flowers uniseriate, pistillate, fertile, ligulate; the ligules obtuse, entire or minutely denticulate, 4-nerved, white to rosy, lavender, or violet blue; style branches linear, more or less flattened, obtuse or subacute. Disk flowers hermaphrodite, fertile, but the central ones often abortive; corollas yellow, tubular-funnelform, with a very short basal tube, or limb and tube scarcely differentiated, the summit regularly 5-lobed; style branches flattened, tipped with a broadly to narrowly acute, triangular appendage; anthers obtuse at base, the acute appendage at the summit somewhat thickened, at least down the center. Achenes 2-, 3-, 4-, 5-, or ro-nerved, compressed to subterete, those of disk and ray either similar or dissimilar. Pappus of 5 equal, erose, hyaline scales, free or united up to half way into a crown; or of scales (sometimes very short) and 1 or 5 alternating, longer, setiform, scabrous awns; or pappus vestigial or apparently wanting, represented by a minute erose crown or merely by 66 WRIGHTIA [Vou. 1, No. 2 a thickened ring on the summit of the achene. Pappus of disk and ray flowers usually similar when present, but that of the ray flowers often shorter, rarely longer; in species normally having both awns and scales, the awns may be wanting in either or both types of flower. Type species: Chaetophora asteroides Nutt. |= Chaetopappa asteroides (Nutt.) DC.] KEY TO THE SPECIES OF CHAETOPAPPA 1. Involucres narrowly cylindrical to broadly conical in flower (broader in fruit), more or less acute at base, with 5-18 rays and 6-26 disk florets; disks 1.5—5.0 mm. across in flower (as pressed).............-0-+00 03 2. a. davolicres. 5:0-6.5 tum, or more high, ¢... 0... 6.8... eee Cee 3 3. Stem leaves broadest near the base or nearly the same width throughout, the upper more or less clasping....2. C. effusd. 3. Stem leaves broadest near the middle or above, not at all ao eas os a ee 4 4. Stem leaves broadly spatulate to elliptic, those just below the inflorescence 8-12 mm. wide, 16-25 mm. long. . ee Ce Pare ete a ek rete cd pew ¢ 3. C. keerlioides. » 4. Stem leaves narrowly spatulate, those just below the in- florescence 1-6 mm, wide, 6-20 mm. long... .4. C. Parryt. ~ Bi AVOUT OR 50-8. 0D. Dig a ag a ee 5. 5. Middle and upper stem leaves broadest near the base or about the same width throughout, the uppermost more or less clasp- ae Se te ay ee ee Ce ew aS 2. C. effusa. 5. Middle and upper stem leaves broadest above the middle, not at WV Olanniiy oO er ea a tr ee ee 6. 6. Plant a low, matted, almost moss-like perennial from rather woody, much-divided rootstocks; stem leaves linear-ob- lanceolate, coriaceous, persistent, of nearly uniform size; species of the mountains of west Texas and New Mexico. Deas ew er ak oe ea ee bea 1. C. Hershey. 6. Plant annual from a simple taproot; stem leaves spatulate to linear, the lower soon withering, the upper much MISAIOE ee a ee te th 7. Inner phyllaries glabrous and entire or with very minutely erose margins, abruptly pointed and often awn-tipped; pappus represented merely by a thick- ened ring or minute erose crown. . .5. C. bellidifolia. 7. Inner phyllaries with prominently lacerate-ciliate margins, especially near the apex, the excurrent midvein made inconspicuous by the ciliation; papPus of well developed scales in either the ray or disk florets, or both, with or without longer awns in sdditighe i: A a ee 8. 1946] SHINNERS: CHAETOPAPPA 67 8. Pubescence of stem and branches loosely to closely appressed, rarely spreading, in plants of the central Texas granite area (a single ex- ceptional collection also seen from Oklahoma); pappus various, but awns usually present in either the ray or disk florets if not both... .. 9. g. Rays 5-13, disk florets 6-15; younger in- volucres narrowly cylindrical; middle phyllaries about o.5-o.8 mm. wide; wide- spread plants of Texas and states north and Gant oc ei 6a. C. asteroides. go. Rays 12-18, disk florets 15-26; even the younger involucres conical; middle phyl- laries about 0.7-1.0 mm. wide, plants of the lower Rio Grande Valley, Texas. ieee Sea Oe 6b. C. asteroides var. grandis. 8. Pubescence of stem and main branches rather coarse and widely spreading, that of the branch- lets and peduncles spreading to appressed, the plant rather shaggy in appearance; pappus of well developed scales only, without awns in either ray or disk florets; plants of south central LOeeB Se in eos 6c. C. asteroides var. imberbis. 1. Involuecres campanulate or hemispherical in flower, rather broad and rounded at base, with 12-28 rays and 32-106 disk florets; disk 4-12 mm. BOrOGl ie etl bie ere ene es Be ass ig OR ee ae aS Io. 10. Pubescence of peduncles appressed, or long, irregularly spreading, and matted; phyllaries imbricated in about 4-6 series.......... 5%; 11. Pubescence of peduncles appressed or closely ascending, the hairs mostly less than 1 mm. long; involucres 4.0-5.1 mm. Wit ee ae ee ee za. C. bellioides... t1. Pubescence of peduncles dense, spreading, more or less matted, the hairs mostly about 1.0-1.5 mm. long; involucres 5.0- G.c. nom. high): 14.35. 7b. C. bellioides var. hirticaulis. 10. Pubescence of peduncles stiffly spreading at right angles, the hairs mostly o.5-0.75 mm. long; phyllaries imbricated in about 3-4 BRPING SS ere ea a ok ee ee ee 8. C. pulchella. — Series 1. HeRSHEYANAE. Perennes exiguae quasimuscoideae, basi sub- lignosae, capitulis paucifloris (disci 5-6); achaenia florum radii 2- vel 3- costata, disci 5-costata. Very small, matted, almost moss-like perennials from more or less woody bases; heads few-flowered (disk florets 5-6); achenes of ray florets 2- or 3-nerved, those of the disk florets 5-nerved. 1. CHAETOPAPPA HersHeYI Blake, Proce. Biol. Soc. Wash. 59: 47-48. 1946, Tufted or matted perennial from slightly woody, vertically or obliquely much forked rootstocks. Stems up to 2 em. long (or according to the col- 68 WRIGHTIA |Vou. 1, No. 2 lector of the type, plants up to 1 or 2 inches, or as much as 5 cm., high), simple, flexuously ascending, sparsely pubescent with ascending or loosely appressed hispid hairs; terminated by a solitary head on a peduncle naked for 1-5 mm., or a few by a tuft of small leaves. Basal rosette-like leaf clusters (beginning of next season’s stems?) with spatulate or oblanceolate, subacute, apiculate leaves 1.5-6.0 mm. long, 0.3-1.0 mm. wide. Stem leaves 4-7, linear-oblanceolate or linear, 3.0-7.5 mm. long, 0.4~1.0 mm. wide, acute and with a translucent spiny tip, coriaceous, r-ribbed (rib almost keel-like beneath), sparsely hispid-strigose or glabrous, persistent; lower leaves with narrowed, somewhat petiole-like bases, the upper slightly smaller. Involucres 4.0-4.6 mm. high; phyllaries lanceolate, acute or acuminate and slenderly spine-pointed, sparsely appressed pubescent on the midrib or glabrous, green or rose-purple to brownish, with white- scarious, ciliate margins, imbricated in about 4 series; middle phyllaries about o.5-0.8 mm. wide. Receptacle flat or in age convex (but extremely small, less than half a millimeter broad, and its true form scarcely disting- uishable), glabrous, the points of attachment of the achenes marked by minute knobs. Ray florets 6-10; ligules when fresh ‘‘bluish, fading to white in age,” when dry reddish purple, elliptic, about 5 mm. long, 2 mm. wide; tube about 2.5 mm. long. Disk florets 5-6, corollas 3.3-4.3 mm. long, yellow. Achenes of ray florets 2- or 3-nerved, 1.3 mm. long (not fully mature), sparsely pubescent. Achenes of disk florets 5-nerved, 0.8~1.° mm. long (not fully mature), pubescent. Pappus similar in disk and ray florets, of 5 (rarely 4 or 6) awns 3.6-4.2 mm. long, and as many very minute, ciliate or erose scales about o.1 mm. long. The type was collected on barren, rocky ledges in canyons of the Guadalupe Mountains, 5000 ft., in Eddy Co., New Mexico, by A. L. Hershey, no. 3532, May 23, 1944 (in herb. U. 8. National Arboretum). Known only from the type and the following collections: TEXAS: Culberson Co., McKittrick Canyon, Guadalupe Mts., Paul C. Standley 40551 (US; collected Aug. 15-17, 1924, showing only the remnants of a few disintegrated heads); Guadalupe Mts., V. Havard in 1882 (G). Series 2. Errusar. Perennes serovernales, aestivales, vel autumnales longicaules, capitulis paucifloris (disci 4-16); achaenia fl. radii bi- vel tri- costata, disci bicostata compressa. Perennials flowering in late spring, summer, or fall; stems rather tall, simple for some distance above the base; heads few-flowered (disk florets 4-16); achenes of ray florets 2- or 3-nerved, those of the disk florets 2- nerved, compressed. 2. Chaetopappa effusa (A. Gray) Shinners, comb. nov. Keerlia effusa A. Gray, Plantae Lindheimerianae II (Boston Journ. Nat. Hist. 6): 222. 1850. Bourdonia effusa (A. Gray) Greene, Erythea 1: 207. 1893. Perennial from a somewhat woody, irregular crown. Stems one ao several, 20-70 em. tall, loosely pubescent with rather long, whitish, spre? 1946| SHINNERS: CHAETOPAPPA 69 ing to appressed hairs, simple up to the diffusely branched inflorescence which occupies the terminal 14—4 of the plant, with about 15—35 leaves below the inflorescence. Leaves rather coriaceous, dark olive green, sparsely strigose on both surfaces and ciliate on the margin with hispid hairs; the lowest oblong to oblong-lanceolate, with more or less narrowed bases, 1.8-5.5 cm. long, o.5~1.3 em. wide, the narrowed base making up 4-14 the total length; middle and upper leaves gradually and slightly reduced, lanceolate-oblong to deltoid-oblong, the upper broadest at base, sessile and more or less clasping, 1-2 em. long, o.5-1.0 em. wide. Leaves of the branches abruptly much reduced to narrowly oblong or (on the branch- lets and peduncles) linear-subulate bracts, the larger ones 0.6-1.5 em. long, 0.3-0.6 em. wide, the smaller as little as o.3 mm. long, the width hardly measurable without a strong lens. Involucres conical or broadly cylindric, 4.0-5.2 mm. tall. Phyllaries imbricated in about 3~s series, oval to oblong-lanceolate, mucronate or abruptly aristate-acuminate, glabrous, the broad scarious margins entire or irregularly erose; middle phyllaries 0.8-1.2 mm. wide. Ray florets 6-9, ligules elliptic-oblong, 3.0—4.5 mm. long, white. Disk florets 4-7, corollas 2.5-3.3 mm. long, yellow. Achenes of ray florets compressed, 2-ribbed, pubescent, obtusely lanceolate in outline, 2.0-2.2 mm. long. Achenes of disk florets irregularly prismatic, 3-, 4-, or 5-ribbed, pubescent on the ribs, 1.6-2.0 mm. long. Pappus a thickened ring or cup-like erose crown, similar in disk and ray florets. Thickets and open ground, dry hillsides, chiefly in rocky limestone soil, in and near the southeastern margin of the Edwards Plateau, south central Texas; flowering from mid July to late September (a single collection dated May 29). The type was collected in shady declivities on the banks of the upper Guadalupe River near Comanche Spring (in northern Bexar Co.), by F. Lindheimer, Fl. Tex. Exs. no. 629, ‘August, September,”’ 1847 (in Gray Herb., not seen; isotypes in herb. Missouri Botanical Garden, herb. New York Botanical Garden, and U. 8. National Herb.). Some specimens examined: TEXAS: Bexar Co., Comanche Spring (? no data given, but specimens probably from type locality), Lindheimer Exs. 933 (C, Mo, NY, T, US). Edwards Co., Leakey, E. J. Palmer 10165 (Mo). Gil- lespie Co., White Oak Creek, G. Jermy 815 (Mo, US). Kendall Co., 5 miles n.e. of Boerne, F. W. Pennell 5483 (NY); 6 miles s. of Boerne, H. B. Parks & V. L. Cory 20709 (TAM); Spanish Pass, Parks & Cory 19389 (TAM); E. J. Palmer 1083 (Mo, US). Kerr Co., Hunt, Hunt School, fall, 1926 (T; apparently the same, without collector, US); near Hunt, Eula Whitehouse, Sept. 1, 1930 (T); Ingram, Parks 27688 (TAM), Parks & Cory 27706 (TAM). Medina Co.?, Amanda Evans, May 20, 1939 (NY, T; label reads merely “Medina *); San Geronimo Creek, J. Reverchon 1534 (Mo; location unknown, but quite probably in Medina Co.). Real Co., Thousand Springs, Parks & Cory 19100-01 (TAM). 3. Chaetopappa keerlioides Shinners, sp. nov. C. effusae et C. Parryi affinis. Perennis? ad 45 em. alta; caulis ad in- florescentiam simplex, infra crebre foliosus; folia inferiora oblongo-cuneata 70 WRIGHTIA [Vou. 1, No. 2 subpetiolata 2.5-3.5 em. longa, o.7-1.1 em. lata, superiora elliptica vel elliptico-oblanceolata sessilia vel subsessilia nec amplectentia. Involucra subeylindrica ca. 5.3 mm. alta. Flores radii 5~9, ligulis oblongis ca. 3.5 mm. longis albis demum roseo-violaceis. Flores disci 12-14, corollis ea. 2.8 mm. longis. Achaenia 5-costata; pappus paleaceus, paleis hyalinis 0.4 mm. (radii) ad 0.7 mm. (disci) longis. Perennial? up to 45 em. tall; root unknown. Stem simple to the diffuse inflorescence which occupies about the terminal 4 or 14 of the plant, loosely villous with widely spreading crisped hairs, with over 50 leaves below the inflorescence, mostly crowded in the basal half of the stem proper. Leaves loosely villous over the whole undersurface and on the veins above, the lower cuneate-oblong, about 2.5-3.5 em. long, o.7—-1.1 em. wide, broadest near the apex, tapered to a narrow base. Middle and upper leaves elliptic or elliptic-oblanceolate, subsessile or sessile but not clasping, the upper- most 16-25 mm. long, 8-12 mm. wide. Leaves of the branches abruptly smaller, lanceolate, up to 18 mm. long, 6 mm. wide. Involucres sub- cylindrical, about 5.3 mm. tall. Phyllaries imbricated in about 4-6 series, lanceolate, glabrous, the scarious margins sparingly erose or erose-ciliolate toward the apex; middle phyllaries about 1.1 mm. wide. Ray florets 5—9, ligules oblong, 3.5 mm. long, white, turning rose violet in withering. Disk florets 12-14, corollas 2.8 mm. long. Achenes of ray and disk florets pris- matie or compressed, 5-ribbed, glabrous. Pappus a crown of 5 partially united hyaline fimbriate scales, o.4 mm. long in the ray florets, 0.7 mm. in the disk florets. The type was collected in a moist wooded canyon on the eastern slope of the Sierra de San Manuel, at Rancho Agua Dulce, Municipio de Muz- quiz, Coahuila, Mexico, by F. Lyle Wynd and C. H. Mueller, no. 369, June 30, 1936, in early flower (in herb. Missouri Botanical Garden; isotypes in Gray Herb., herb. University of California, herb. New York Botanical Garden, and U. S. National Herb.). Known only from the type collection. This plant stands between C. effusa and C. Parryi. It has the large stature and diffuse inflorescence of the former, but the more numerous disk florets of the latter, while the broad but narrow-based leaves rather crowded in the lower part of the stem are intermediate between the two. It differs from both in the pubescence and shape of the leaves. 4. CHAETOPAPPA Parryl A. Gray, Proc. Amer. Acad. 16: 82. 1880. Distasis Parryi (A. Gray) Kuntze, Rev. Gen. Pl. 1: 334. 1801. Perennial, forming loose tufts from superficial rhizomes or stolons (resembling those of Antennaria plantaginifolia and its relatives, but much smaller); rhizomes producing a few leafy rosettes (these perhaps merely an early stage in the development of flowering stems). Stems erect, ro~28 em. tall, normally simple for 4-18 em. above the base (the terminal 14-24 of the plant with few, elongate, spreading-ascending branches), sparsely to rather densely pubescent with appressed to loose or spreading somewhat hispid hairs, with about 8-25 leaves below the inflorescence 1946] SHINNERS: CHAETOPAPPA 71 (including the basal ones). Basal leaves crowded in a persistent rosette- like cluster, narrowly oblanceolate to broadly spatulate, 10-30 mm. long (the petiolar base 4-16 mm.), 1.5-9.0 mm. wide, apiculate, sparsely strigose and ciliate, with a single prominent vein beneath. Proper stem leaves few, like the basal ones but rather abruptly smaller, the upper gradually reduced, the middle ones about 8-20 mm. long, 1.0-3.7 mm. wide, those of the branches reduced to narrowly oblanceolate or linear bracts. Involucres conical or broadly subeylindric, 5.2~-6.5 mm. high. Phyllaries imbricated in about 3-5 series, lanceolate, glabrous, rather minutely serrulate-ciliolate toward the apex; middle phyllaries o.9—1.1 mm. wide. Ray florets 7-9, ligules oval or oblong-oval, 2.5—3.5 mm. long, pink in bud, white in flower. Disk florets 10-15, corollas about 2.5 mm. long, yellow. Achenes 2.2.—2.8 mm. long, sparingly hispidulous, gradually tapered in the basal two thirds, abruptly contracted near the summit; those of the ray florets 3-ribbed, those of the disk florets 3-, 4-, or 5-ribbed (the varia- tions often to be found in the same head). Pappus of 5 hyaline, fimbriate- ciliate scales united up to half way into a cup-like crown 0.25-0.85 mm. long; awns absent, or one (most commonly), or five, o.8-2.5 mm. long. Pappus similar in disk and ray florets, but that of the latter usually shorter. The type was collected at “Mt. Carmel on the Rio Grande,” by C. C. Parry, Nov. 8, 1852 (in Gray Herb.; isotype, as Mexican Boundary Survey No. 492, year omitted, in herb. New York Botanical Garden). From the letter of M. T. W. Chandler in the report of the Mexican Boundary Survey, dated at Ft. Duncan, Dee. 1, 1852, it appears that the type locality was in extreme northwestern Coahuila, in the Sierra del Carmen, opposite the Chisos Mountains of Brewster Co., Texas, and within a few miles of the Rio Grande. As yet the plant is known only from northern Mexico. Additional specimens examined: COAHUILA: Municipio de Cuatro Cienegas, Sierra de la Madera, Cafion del Pajarito, C. H. Muller 3182 (SMU). Municipio de Villa Acufa, Sierra del Carmen, Cafionde Sentenela on Hacienda de Piedra Blanca, Wynd & Mueller 531 (G, Mo, NY, US), 437(US). NUEVO LEON: Sierra Madre Oriental, San Francisco Cafion, 15 miles s.w. of Pueblo Galeana, C. H. Mueller & M. T. Mueller 269 (G, T). TAMAULIPAS: 4 km. w. of Miquihuana, lat. 23°42’ N. long. 99°45’ W., L. R. Stanford, K. L. Retherford, & R. D. Northcraft 604 (G, Mo, NY). Series 3. ASTEROIDEAE. Annuae vernales humiles ramosissimae capitulis paucifloris (disci 6-26); achaenia fl. radii et disci subcompressa vel sub- teretia quinque- vel decemcostata. Low, spring-flowering, much branched annuals; heads few-flowered (disk florets 6-26); achenes of ray and disk slightly compressed to nearly terete, 5- or 10-ribbed. 5. Chaetopappa bellidifolia (Gray & Engelm.) Shinners, comb, nov. Keerlia bellidifolia Gray & Engelm. ex A. Gray, Proce, Amer. Acad. a: 47- 1846 (dated 1848). Republished in Pl. Lindh. II (Bost. Journ. Nat. Hist. 6): 220~221. 1850. Bourdonia bellidifolia (Gray & Engelm.) Greene, Erythea 1: 207. 1893. 72 WRIGHTIA [Vou. 1, No. 2 Annual from a taproot which is usually bent or twisted at the base of the stem. Stems single or few, 8-40 em. tall, loosely and rather widely bushy-branched from the base, up to 50 em. across when in flower, the branches ascending or spreading-ascending, the branching markedly cymose, the early-flowering tip of the main axis soon overtopped and hardly discernible on full grown plants. Stem and main branches rather coarsely and densely pubescent with widely spreading hairs; pubescence of branch- lets and peduncles spreading to loosely appressed. Basal leaves (soon drying and disappearing) oblanceolate to oblong-spatulate, hispid pubes- cent on both surfaces and ciliate on the margin, especially in the narrowed basal portion, subacute or obtuse, o.7-3.5 em. long (petiolar base about half as long), 3.0-8.5 mm. wide. Stem leaves similar, the upper gradually smaller, the petiolar base shorter and less distinct. Involucre narrowly to broadly conical, 3.8-4.5 mm. tall. Phyllaries imbricated in about 3-4 series, broadly lanceolate or lanceolate-oval, often awn-tipped by the excurrent midrib, glabrous, the broad scarious margins entire or very minutely erose; middle phyllaries o.8-1.2 mm. wide. Ray florets 6~15, ligules oblong, 2.6-4.0 mm. long, 1.1.-2.0 mm. wide, violet or lavender to violet blue. Disk florets 8-22, corollas 2.5-3.0 mm. long, yellow. Achenes subterete, somewhat clavate, 10-ribbed, glabrous or sparsely pubescent, 1.7~2.2 mm, long. Pappus usually none, represented by a thickened ring on the summit of the achene, or rarely by a minute erose crown. Late flowering form: spring leaves withered, plants nearly leafless, intricately much branched, the many delicate secondary branchlets ter- minated by the numerous, solitary, narrowly conical or cylindrical in- volucres, the heads much narrower than in the spring form, resembling those of young C. asteroides. This develops in summer or fall in favorably moist seasons, the spring plants persisting and flowering a second time. An abundant and rather showy little spring-flowering plant of open rocky limestone soil in the eastern and southeastern sections of the Ed- wards Plateau and adjacent areas, central Texas. The type was collected ‘in margin of woods and thickets, in sterile soil, Comale Creek and near New Braunfels, April to June,” 1846, by Lindheimer, Fl. Tex. Exs. 415 (in Gray Herb., not seen; isotypes in herb. University of California, herb. Missouri Botanical Garden, herb. New York Botanical Garden, and U.S. National Herb.). Some additional specimens examined: TEXAS: Bandera Co., ro miles s.e. of Bandera, Cory 1029 (G). Bell Co., 3 miles s. of Salado, Simon E. Wolff 2934 (TAM); near Sparta, Wolff 3784 (TAM). Bexar Co., San Antonio, B. C. Tharp, April 1, 1921 (T). Blanco Co., Blanco, Whitehouse, April 6, 193° (T). Caldwell Co., without locality, Lucille Barber, April 5, 1931 (T). Comal Co., New Braunfels, Lindheimer Exs. 628 (Mo, US), 932 (C, Mo, NY, T, US). Edwards Co., Rocksprings, Geo. L. Fisher 3694, July 19, 1936 (US); 19 miles s.e. of Rocksprings, Lloyd H. Shinners 7335, 7355 (SMU). Gillespie Co., Threadgill, G. Jermy 615 (US; same number, without locality, Mo). Hays Co., San Marcos and vicinity, S. W. Stanfield, June, 1896 (NY). Kerr Co., 2- of Hunt, Whitehouse 10419 (SMU); Kerrville, Mrs. J. M. Milligan, May, 1897 1946| SHINNERS: CHAETOPAPPA 73 (US), EB. J. Palmer 9946 (Mo, US). Kimble Co., Junction, Whitehouse 10282 (SMU; apparently the same, without number, T). Kinney Co., Laguna to Brackettville, Whitehouse 10419 (SMU). Mason Co., Loyal Valley, LE. Dapp- rich, in 1881 (SMU). Real Co., 1 mile s. of Camp Wood, Parks & Cory 18295 (TAM); 1 mile n. Leakey, Shinners 7323 (SMU); 16 miles n. of Leakey, Cory 8571 (USNA; apparently the same, with same number but locality given as “below Thousand Springs,’ TAM). Sutton Co., Sonora, M. E. Jones 28028, 28025 (C), Richard W. Pohl 4756 (SMU). Travis Co., Austin, J. E. Bodin 30 (US; apparently the same, without number, C), Palmer 13665 (Mo, US), Tharp, April 28, 1933 (C-Cl, T, TAM), April 28, 1940 (C-Cl, T), June 25, 1920 (C-Cl, T, TAM); Bull Creek, HZ. R. Bogusch, May 2, 1926 (US); near Marshall Ford Dam, C. L. Lundell & Amelia A. Lundell 9091 (SMU). Uvalde Co., Sabinal, Whitehouse, March 30, 1933 (T); 12 miles s.w. of Uvalde, White- house 10402 (SMU); 20 miles n. of Uvalde, Shinners 7309 (SMU). Val Verde Co., n. of Del Rio, M. E. Jones 28024 (C); Devils River, C. R. Orcutt 6013 (Mo); 12} miles s. of Loma Alta, Cory 39484 (T). Heller 1436 from Nueces Co., cited as Keerlia bellidifolia by Miss Larsen (1933, p. 38), is Chaetopappa asteroides var. imberbis. 6a. CHAETOPAPPA ASTEROTDES (Nutt.) DC., Prodr. 5: 301. 1836. Chaetophora (misprinted Chaetanthera) asteroides Nutt., Journ. Phila. Acad. FELIT LOZA. Diplostelma filiformis Raf., New Fl. N. A. 2: 45. 1837 (dated 1836). Diplostelma pumila Raf., New Fl. N. A. 2: 45. 1837 (dated 1836). Diplostelma radians Raf., New Fl. N. A. 2: 45. 1837 (dated 1836). Distasis asteroides (Nutt.) Kuntze, Rev. Gen. Pl. 1: 334. 1891. (As D. asterodes.) Slender annual from a taproot. Stems single or less often several, 3.5—30 em. tall when in flower, simple or usually divaricately branched, the branches spreading-ascending to horizontal, rather distant and elongate, the ultimate branchlets or peduncles terminated by a solitary small head. Branching markedly cymose: the main axis terminated by the first head to flower, the remaining heads flowering later on branches and branchlets which soon overtop and obscure the first one. Stem and branches strigosely and somewhat hispidly pubescent. Plants at first with a basal rosette of oblanceolate to spatulate-orbicular, obtuse, apiculate, loosely strigose, hispid-ciliate leaves 4-25 mm. long (petiolar base 14-4 the total length), I~5 mm. wide. Stem leaves similar but narrower, the upper gradually smaller, those of the branches reduced to linear or subulate bracts. In- volucres 3.5—4.5 mm. tall, narrowly cylindric to conical. Phyllaries imbri- cated in about 3—4 series, lanceolate, more or less hispid-pubescent on the back, fimbriate-ciliate toward the apex, the inner prominently so; middle phyllaries about o. 5-0.8 mm. wide. Ray florets 5-13, ligules oblong, 2.5- 4.0 mm. long, 0.8-1.8 mm. wide, white, sometimes turning lilac or rosy violet in withering, very rarely colored from the first. Disk florets 6-15, corollas 2.5~3.0 mm. long, yellow. Achenes 1.6-2.0 mm. long, prismatic, S-nerved or -ribbed, pubescent. Central florets commonly retarded in development, often not maturing achenes. Pappus of 5 hyaline scales °.1-0.8 mm. long, alternating with as many awns 1-3 mm. long, or awns 74 WRIGHTIA (Vox. 1, No. 2 rarely wanting. Pappus of ray like that of the disk florets or shorter, rarely the reverse. Pappus of abortive central florets like the rest, or much reduced, or sometimes awnless while the rest have awns. Late flowering form: plants nearly leafless, intricately branched, the branchlets very slender, naked or with few minute subulate bracts, the numerous involuecres very narrowly cylindrical. As in C. bellidifolia, this form may develop in summer or fall in favorably moist years. Rafinesque’s Diplostelma filiformis probably represented this form. Llano form: stem pubescence loosely appressed to widely spreading, but plants not as coarse as in var. imberbis, and pappus awns usually present. Collections from the granite region of Llano and Burnet Counties, south central Texas, and a single specimen from the Arbuckle Mountains, Murray Co., Oklahoma, differ from typical C. asteroides in having spread- ing stem pubescence, but are otherwise like the species. A delicate little spring-flowering annual of sandy or rocky open ground or open woods, from south central Texas north and east to Kansas, Mis- souri, and Louisiana. Uusally abundant where found, growing by itself or with C. bellidifolia, Astranthium integrifolium (Michx.) Nutt., Aphano- stephus ramosissimus DC,, or A. skirrhobasis (DC.) Trel. Dwarf, 1-headed plants of all these species can look surprisingly alike, in spite of marked differences in detail. In the field, C. asteroides and C. bellidifolia can best be distinguished by the latter’s more robust stature, colored rays, and broader, glabrous involucres with broad phyllaries. It is possible that the two hybridize, the pale rays and rather intermediate appearance of occasional plants where the two grow together suggesting hybridization, but evidence from detailed examination is not convincing. Astranthium and Aphano- stephus are best distinguished from both species of Chaetopappa in the field by the broad, very shallow involucres, with outer phyllaries more than half as long as the inner, and by the fact that the rays do not coil, as in Chaetopappa, but are either deciduous before shrivelling (in Astranthium) or remain extended in drying (in A phanostephus). The type was collected by Thomas Nuttall in ‘“Arkanzas”’; it is probably in the British Museum. There are isotypes in the Gray Herbarium and in the herbarium of the New York Botanical Garden. Some specimens examined: ARKANSAS: Crawford Co., Alma, Geo. C. Haas & Flora A. Haas 230 (US); Van Buren, Delzie Demaree 15309 (Mo). Western Arkansas, Herb. F. L. Harvey, June, 1899 (US) (Harvey collected chiefly in the vicinity of Fayetteville, Washington Co.). KANSAS: Chautauqua Co., 1 mile s. of Sedan, L. H. Shinners, Grant Cottam & H. A. Stephens 3553 (C-Cl). LOUISIANA: Caleasieu Parish, Lake Charles, E. N. Plank 42 (US). MISSOURI: Cedar Co., without locality, B. F. Bush 14908 (Mo). Jasper Co., 3 miles n.w. of Joplin, E. J. Palmer 32359 (Mo). Newton Co., 4 miles 5. of Joplin, Z. J. Palmer 29929 (Mo); Grand Falls, Bush 10456 (Mo, NY, US). Vernon Co., without locality, G. C. Broadhead, June 25, 1873 (G). OKLA- HOMA: Atoka Co., Limestone Gap, Gee. D. Buller 104 (Mo). Carter Co., near Ardmore, G. W. Stevens 75 (G, Mo, NY, US). Cleveland Co., 6 miles €. 1946] SHINNERS: CHAETOPAPPA 75 of Norman, Demaree 12798 (Mo, NY). Comanche Co., 7 miles w. of Cache, Reginald Rose-Innes & Brunelle Moon 989 (G); Ft. Sill, Mrs. J. Clemens 11814 (G, Mo.). Creek Co., Sapulpa, Bush 928 (C, Mo). Johnston Co., near Manns- ville, Florence Griffith, May 20, 1916 (Mo, NY); near Tishomingo, H. W. Houghton, April 8, 1916 (G, NY). Le Flore Co., Poteau, E. J. Palmer 8255 (Mo, NY, US). Lincoln Co., Fonts, J. W. Blankinship, Aug. 27, 1895 (G, Mo, US). Murray Co., Arbuckle Mts., G. J. Goodman 2476 (G, Mo, NY; exceptional form with spreading stem pubescence); Davis, Arbuckle Mts., Demaree 12289 (NY), W. H. Emig 552 (Mo); near Crusher Spur, Stevens 51.1 (G). Muskogee Co., Muskogee, M. A. Carleton, April, t891 (US). Ottawa Co., near Lincoln- ville, E. J. Palmer 29929 (Mo, NY). Payne Co., Stillwater, F. A. Waugh, June 10, 1893 (US; apparently the same, as no. 10, Mo). Seminole Co., Semi- nole, Demaree 12713 (Mo, NY); 6 miles s.e. of Seminole, U. T. Waterfall 584 (NY). Sequoyah Co., near Sallisaw, 2. J. Palmer 33265 (Mo, NY). TEXAS: Anderson Co., near Palestine, Whitehouse 10577 (SMU). Angelina Co., s. of Lufkin, Whitehouse 10206 (SMU). Austin Co., Catspring, Lindheimer, April- July, 1844 (Mo, SMU); Kenney, Pennell 10309 (NY). Bastrop Co., 2 miles e. of Bastrop, in Bastrop State Park, Shinners 7271 (SMU). Bell Co., 3 miles n. of Holland, Wolff 808 (TAM). Bexar Co., 14 miles n. of San Antonio, Sister Mary Clare 511 (C). Brazos Co., Bryan, E. J. Palmer 7786 (Mo, NY, US); near Fish Lake, College Station, D. C. Bain, April 29, r940 (TAM). Burnet Co., 5 miles s. of Burnet, Wolff 3819 (TAM); Inks Lake State Park, w. of Burnet, Shinners 7221 (SMU; form with spreading stem pubescence). Calhoun Co., Port Lavaca, Tharp, May 22, 1930 (T). Clay Co., Henrietta, Bush 5431 (G, Mo). Comanche Co., Comyn, Biology Students, April 30, 1930 (T). Coryell Co., near Holvert, Wolff 1407 (US). Dallas Co., Dallas, Reverchon, various dates and numbers (C, G, Mo, US); Bush 593 (Mo, US). DeWitt Co., Cuero, Bray 135a (US; apparently the same, without number, T). Eastland Co., Carbon, Center Point School, May 7, 1930 (T). Frio Co., 5 miles n.w. of Derby, Cory 12780 (USNA). Gillespie Co., Bear Mt., Parks & Cory 12974-76 (TAM; same, as Cory 12977, USNA). Gonzales Co., Ottine, E. R. Bogusch 986 (T; not typical). Grimes Co., Anderson, Fisher, April 9, 1941 (US). Hamilton Co., 2} miles s. of Hico, Shinners 7182 (SMU). Hardin Co., Fletcher, E. J. Palmer 9542 (Mo, US). Harris Co., Hockley, F. W. Thurow, in 1889 (US). Hood Co., Granbury, Eggert, May 6, 1900 (Mo, NY). Houston Co., 7 miles s. of Crockett, K. M. Wiegand & M. C. Wiegand 2420 (G). Jack Co., 9 miles n.w. of Jacks- boro, Shinners 7949 (SMU). Jackson Co., Edna to Victoria on Highway 509, Whitehouse 10401 (SMU). Kaufman Co., vicinity of Terrell, F. J. Tyler, April 23, 1904 (US). Lee Co., without locality, V. H. Williams, March < 1939 (TAM). Leon Co., 13 miles s.w. of Buffalo, Shinners 7132 (SMU); 23 miles s.w. of Buffalo, Shinners 7137 (SMU). Live Oak Co., southern part, Tharp, March 22, 1931 (NY, T). Llano Co. (the following 7 collections are all of the form with spreading stem pubescence), Bauman’s, Parks & Cory 15137 (TAM); Enchanted Rock, R. R. Innes 899 (G), Whitehouse 10288 (SMU); near Inks Dam, C. L. Lundell & Amelia A. Lundell 9048 (SMU); Llano, Bray 135 (US); along Colorado River 1 mile s. of bridge e. of Llano, Shinners 7195 (SMU); Granite Mt., near Llano, Wolff 3845 (TAM). Mills Co., Mullers Tp., Mamie Egg, April 14, 1931 (T). Montgomery Co., 30 miles n. of Houston, Herb. Charles Mohr, April 24, 1839 (US); Willis, M. B. G. Herb. no. 900588, in March, year not given (Mo). Navarro Co., Raleigh, Reverchon, April 1s, 1903 (Mo). Nueces Co., Corpus Christi, H. W. Ravenel 76 (Mo, 76 WRIGHTIA (VoL. 1, No. 2 NY); Nueces Bay, A. Arthur Heller 1436 (NY; duplicates are mostly var. imberbis); Viola, H. C. Benke 5444 (G). Parker Co., Weatherford, S. M. Tracy 7897 (G, Mo, NY, T, US). Polk Co., 14 miles e. of Corrigan, Cory 22156 (USNA). Tarrant Co., Fort Worth, O. L. Killian, in 1927 (T); Lake Worth, A. Ruth 1593, July 5, 19290 (SMU), 704, April 14, 1925 (NY); same number, without locality, April 14, 1923 (G), April 14, 1919 (US). Taylor Co., n. of Abilene, Eggert, May 6, t900 (Mo). Titus Co., Talco School, July, 1927 (T). Travis Co., Austin, M.S. Young, May 13, 1918 (C-Cl, Mo, USNA); F. Rugel, in 1868 (Mo); Bray 52 (NY); 28 miles n.w. of Austin on road from Beecaves to Spicewood, Shinners 7245 (SMU). Trinity Co., Trinity, Plank, May 17, 1892 (NY). Van Zandt Co., 3 miles e. of Wills Point, Shinners 7540 (SMU). Viec- toria Co., Victoria, EZ. J. Palmer 9100 (Mo). Walker Co., vicinity of Hunts- ville, Royal A. Dixon 409 (G, NY). Waller Co., Hempstead, Elihu Hall 307 (G, Mo, NY). Washington Co., without locality, Eunice Brackett, June 20, 1938 (T). Wilson Co., Kicaster School, Parks & Cory 15137 (TAM). Wise Co., 3 miles w. of Decatur, Shinners 7925 (SMU). 6b. CHAETOPAPPA ASTEROIDES var. grandis Shinners, var. nov. Distasis modesta DC., Prodr. 5: 279. 1836. Chaetopappa modesta (DC.) A. Gray, Proc. Amer. Acad. 16: 82. 1880. (As to name on which based, but the new combination was incorrectly used for Chaetopappa bellioides.) Robustior, plerumque multicaulis; capitula majora, fl. radii 12-18, disei 15~25; phyllaria mediocria ad 0.7—1.0 mm. lata. More robust than the species, when well developed usually with several or many stems and bushy-branched. Involucres narrowly to broadly conical, even when young; middle phyllaries about 0.7-1.0 mm. wide. Ray florets 12-18, ligules 3-4 mm. long; disk florets 15-25. Pappus ap- parently always of both scales and awns. Confined to the lower Rio Grande Valley, as yet known only from southern Texas, but to be expected in adjacent Tamaulipas. Flowering slightly earlier than the species, beginning in February. The type was collected on a gravelly hill off U. S. Highway 83, 3 miles east of Sullivan City, Hidalgo Co., Texas, by C. L. Lundell and Amelia A. Lundell, no. 9982, April 5, 1941 (in herb. Southern Methodist University). Rather than perpetuate a name based on the inadequate scraps which constituted the type and isotypes, and which were largely responsible for the long misunderstanding as to the proper identity of the plant, I have deliberately chosen a new name based on a new and better type specimen. Additional collections examined: TEXAS: Hidalgo Co., La Joya (Sam Fordyce), Mrs. E. J. Walker 26 (G, T). Webb Co., Laredo, Berlandier 1416 G, Mo, NY, US; isotypes of Distasis modesta DC.); Reverchon 4007 (Mo, NY). 6c. CHAETOPAPPA ASTEROIDES var. IMBERBIS A. Gray, Proc. Amer. Acad. 16: 82. 1880. Coarser and stouter than the species; pubescence of stem and main 1946| SHINNERS: CHAETOPAPPA 74 branches longer and widely spreading, appearing rather shaggy. Ray florets 6-14, disk florets 11-18. Pappus of scales only, without awns. The type was collected in Texas by Charles Wright (in Gray Herb.). Gray speaks of it as having come from ‘‘east Texas,’ which would suggest that it was collected in 1844 in either Angelina, Jasper, or Tyler County, far out of the range of the variety as now known. An apparent duplicate in the U. S. National Herbarium, consisting of a mixture of the species, var. imberbis, and Krigia virginica (L.) Willd., bears a label with the information that it was collected between the Colorado and the Guadalupe, in post oak woods, in April. Dr. Geiser tells me that this collection probably was made in 18409 in Fayette Co., or possibly in Gonzales Co., while Wright was en route to San Antonio, to begin his 1849 expedition to western Texas. Fayette and Gonzales counties are in the area to which the variety is apparently confined. As far as known at present, it occurs in a limited area in south central Texas, centering on the northern part of the Rio Grande Plain, either by itself or with the species. Additional specimens examined: TEXAS: Atascosa Co., Campbellton to Pleasanton, Whitehouse 10243 (SMU); n. of Pleasanton on Highway 281, Whitehouse 10346 (SMU). Bexar Co., 16 miles s. of San Antonio, Ellen D. Schulz 445 (US); 18 miles s. of San Antonio, Sister Mary Clare Metz 2172 (NY). Caldwell Co., without locality, J. B. McBryde, spring-summer, 1931 (T). Gonzales Co., 101 miles w. of Monthalia, Parks & Cory 7804 (TAM); Ottine, Bogusch 1735 (T); Palmetto State Park, Innes 618 (G); without locality, Bogusch 1438 (C-Cl). Guadalupe Co., Seguin, Whitehouse 10260 (SMU). Nueces Co., Nueces Bay, Heller 1436 (G). Wilson Co., Sutherland Springs, Parks & Cory 7804 (TAM); near Sutherland Springs, Cory 8571 (USNA). Series 4. BreLitiomear. Annuae vel perennes caespitosae, capitulis multifloris (disci 32-106); achaenia fl. radii tricostata (rarius bicostata), disci bicostata compressa. Annuals or tufted perennials; heads many-flowered (disk florets 32- 106); achenes of ray florets 3-ribbed, rarely 2-ribbed, achenes of disk florets 2-ribbed, compressed. 7. Chaetopappa bellioides (A. Gray) Shinners, comb. nov. Diplostelma bellioides A. Gray, Plantae Fendlerianae (Mem. Amer. Acad. N.S. 4): 72-73. 1840. This has generally been known as Chaetopappa modesta (DC.) A. Gray, but the type collection of Distasis modesta, on which this name was based, belongs to C. asteroides var. grandis. Perennial, but flowering the first year and appearing annual, with a taproot (ultimately a forked caudex) producing several to many simple or sparingly branched, ascending or spreading, strigose-pubescent stems 3-5~28 em. tall when in flower; these usually at first short and compact, elongating in age, or early elongate in shady places. Plants at first with a basal rosette-like cluster of oblanceolate or spatulate, subacute to obtuse and apiculate, hispidly strigose and ciliate leaves 0.5-4.7 em. long (petio- 78 WRIGHTIA [VoL. 1, No. 2 lar base 0.3—2.7 em.), 2-8 mm. wide; these soon withering and disappearing. Proper stem leaves similar but narrower, the upper gradually smaller. Heads solitary and terminal on the stems or branches, the later ones on peduncles naked for as much as 2.5 em. below the head. Involucre hemis- pherical or broadly campanulate, 4.0-5.1 mm. high, the disk 4-12 mm. across in flower (as pressed). Phyllaries imbricated in about 4-5 series, broadly lanceolate or oblong, appressed pubescent on the back, the broad white scarious margin lacerate-ciliate, especially towards the apex, the central green portion commonly tinged red or purple toward the tip; middle phyllaries 1.0-1.3 mm. wide. Ray florets 12-18, ligules linear- oblong, 3.5~5.2 mm. long, white to lavender or violet blue. Disk florets 33-105, corollas 2.8-3.5 mm. long, yellow. Achenes about 2 mm. long, pubescent; those of the ray florets irregularly prismatic, 2- or 3-ribbed, those of the disk florets compressed, obtusely lanceolate in outline, 2- ribbed. Pappus of 5 narrowly oblong, hyaline, erose scales o.6—-1.8 mm. long, often united at base, and as many scabrous, setiform awns 2.2-3.0 mm. long. Dry, rocky, open ground, southwestern Texas and north-central Mexico, as far south as Aguascalientes. Plants flowering in almost any month of the year, depending on rainfall. The type was collected at Buena Vista battle-field, about 10 miles north-east of Saltillo, Coahuila, Mexico, by Dr. Josiah Gregg, March 19, 1847 (in Gray Herb., without exact date; isotypes in herb. Missouri Botanical Garden and herb. New York Botani- cal Garden). Additional specimens examined: AGUASCALIENTES: near Aguascalien- tes, J. N. Rose & Joseph H. Painter 4777 (US). CHIHUAHUA: Chihuahua, Harde LeSueur, Aug. 20, 1935 (Mo, T); near Chihuahua, C. G. Pringle 972 (Mo, NY, US); vicinity of Chihuahua, Dr. E. Palmer 107 in 1908 (Mo, US); Santa Eulalia Mts., Pringle 205 (C, G, NY), E. Wilkinson, in 1885 (NY, US). COAHUILA: near Diaz, Pringle 205 (C), 8295 (G, Mo, NY, US). La Rosa, w. of Saltillo, Forrest Shreve & E. R. Tinkham 9581 (C). Municipio de Ramos Arizpe, e. of Hacienda La Rosa, Wynd & Mueller 38 (G, Mo, US). Saltillo, Dr. E, Palmer 507 in 1880 (Mo, US); Geo. L. Fisher 188 (US), ‘‘Fr. Adole L., Pl. de Mexique 6506, misit Fr. G. Arsene” (US); Saltillo and vicinity, Dr. FE. Palmer 47 in 1898 (C, G, Mo, NY, US); 24 miles w. of Saltillo, Juan M. John- ston 7680 (G), Shreve 8739 (USNA). Sierra de Parras, C. A. Purpus 1022 (C, G, Mo, NY; collections dated Feb., 1905, and March, 190s, all under same number). Soledad, 25 miles s.w. from Monclova, Dr. E. Palmer 508 in 1880 (US). NUEVO LEON: Cerralbo, Gregg, May 29, 1847 (Mo). Near Rio Ramos, 20 km. n.w. of Montemorelos, J. N. Weaver 1030 (G, TAM). Monterey, Dr. Edwards & Maj. Eaton, in 1846 (NY); Bishop’s Hill, Monterey, Gregg, Feb. 6, 1847 (Mo); Sierra Madre Mts., Monterey, Mueller & Mueller 255 (G; same, without number, T). SAN LUIS POTOSI: Charcas, C. L. Lundell 5748 (US, USNA), 5067 (US). Minas de San Rafael, Purpus 4967, 5150, 5164 (C), 5019 (C, G, Mo, NY, US; all with narrower and more acute leaves than usual, and rather smaller heads, with as few as 13 rays, 32 disk florets). Near Salitre, 15 miles n.w. of Salado, Shreve 9359 (G, USNA). ZACATECAS: Cedros, Francis E. Lloyd 115 (US). Hacienda de Cedros, Lloyd 188 (C, US; 1946] SHINNERS: CHAETOPAPPA 79 locality further indicated as ‘Zacatecas, norte’). NEW MEXICO: without locality, Charles Wright 1173, in 1852 (G, NY, US). TEXAS: Brewster Co., Marathon, Cory 2585 (G); 25 miles s. of Marathon, L. C. Hinckley, Oct., 1936 (G, NY); without locality, Tharp, Oct. 9, 1936 (C-Cl, Mo, SMU, T). Hidalgo Co., 5 miles n. of Mission, Parks & Cory 18036 (TAM; same, as Parks 18036, C). Kinney Co., Ft. Clark, Edgar A. Mearns 1264, 1309 (US). Maverick Co., Eagle Pass, V. Havard, Sept. 1882 (US); Coal Mine, Eagle Pass, Bray, May 21, 1898 (T). Presidio Co., foothills of the Chinati Mts., Young, Sept. 11, 1914 (Mo, T). Terrell Co.?, Langtry-Dryden, Tharp, June 14, 1931 (Mo, T). Uvalde Co., Uvalde to Cline, Whitehouse 10585 (SMU). Val Verde Co., near Comstock, Whitehouse 10533 (SMU); Del Rio, E. 0. Wooton, Nov. 5, 1913 (US); vicinity of Del Rio, Rose & Fitch 17985 (Mo, US); to miles n. of Del Rio, Parks & Cory 20851 (TAM; same, as Cory 20852, USNA); 13 miles n.w. of Del Rio, Cory 43867 (SMU); w. of Del Rio, White- house 10583 (SMU); Langtry, F. S. Earle & Esther S. Earle 445 (Mo, NY, US), Orcutt 6149 (Mo); 2 miles e. of Langtry, R. R. Innes & Barton H. War- nock 597 (G). 7b. CHAETOPAPPA BELLIOIDES var. hirticaulis Shinners, var. nov. Differt pubescentia longiore patula vel hispido-lanosa, capitulis major- ibus (involucris ca. 5.0-6.5 mm. altis). Stems and branches densely pubescent with more or less spreading or matted whitish hairs about 1.o-1.5 mm. long. Heads larger, the involucres 5.0-6.5 mm. high, The type was collected on the Cerro Tres Vetas, vicinity of San Jose, alt. 2700 ft., Sierra de San Carlos, Tamaulipas, Mexico, by H. H. Bartlett, no, 10359, July 15, 1930 (in herb. Southern Methodist University; isotype in U. 8. National Herb.). Known only from the region of the type locality; both collections seen were made in July. Additional specimen examined: TAMAULIPAS: Sierra de San Carlos, vicinity of San Jose, Cerro de los Armadillos, Bartlett 10171 (US). 8. Chaetopappa pulchella Shinners, sp. nov. Annua (adeo perennis?) parva 3-7 em. alta ramosa, omnino hispido- pubescens pilis albidis patentissimis brevibus. Folia infima spathulata vel anguste oblanceolata, 6-12 mm. longa, 1.5-4.0 mm. lata, decidua, superiora similia minora (nisi surculis tardis). Involucra campanulata ca. 5-0-6.0 mm. alta, phyllariis 3—4-seriatis lanceolatis. Flores radii 15-20, ligulis lavandulis 5 mm. longis, 1.5 mm. latis. Flores disci 38, corollis 3.5 mm. longis. Achaenia radii 2-, 3-, vel 4-costata; disci bicostata compressa. Pappus duplex paleis 5 scariosis r.r mm. longis et aristis 5 setiformibus 3-3 mm. longis. Low annual with a taproot, perhaps persisting and becoming perennial ; stems 3-7 em. tall, branched from near the base, densely and uniformly pubescent with short whitish hairs about o.5 mm. long, stiffly spreading at right angles. Leaves hispid pubescent and densely ciliate with short hairs like those of the stem, the lower spatulate to narrowly oblanceolate, 6-12 mm. long (petiolar base 2-5; mm.), 1.5-4.0 mm. wide, soon turning 80 WRIGHTIA [VoL. 1, No. 2 yellow and falling; middle and upper stem leaves similar but smaller. Involucre broadly campanulate, 5.0-6.1 mm. tall, the disk 7-8 mm. across (as pressed). Phyllaries imbricated in about 3-4 series, lanceolate, hispid-pubescent on the back, the scarious margins prominently fimbriate- ciliate toward the apex, the green central portion rosy or purple tinged at tip, with a keel-like midrib prominent in the lower 14-24; middle phyllaries r.1-1.2 mm. wide. Ray florets 15—20, ligules elliptic-oblong, 5 mm. long, 1.5 mm. wide, lavender. Disk florets (one count) 38, corollas 3.5 mm. long, yellow. Achenes about 2 mm. long, sparsely pubescent, those of the ray florets compressed or irregularly prismatic, 2-, 3-, or 4-ribbed; those of the disk florets compressed, 2-ribbed. Pappus of 5 hyaline scales about 1.1 mm. long, and as many setiform, scabrous awns 3.3 mm. long. The type was collected among small stones in a bajada (swampy place), on the road from Monclova westward to beyond Cuatro Cienegas, at 4 miles west of Cuatro Cienegas, Coahuila, by Ivan M. Johnston, no. 7151, August 24-26, 1938 (in Gray Herb.). The type collection consists of a number of diminutive, divaricately branched, yellow green annuals in flower and fruit. The second collection, cited below, consists of a single small plant from a slender taproot and apparently three new stems arising from the dead or broken base of an old one. It is not clear whether this represents a perennial, or merely a revival in growth due to rains, in the fashion of C. bellidifolia and C. aster- oides, of a plant which is normally very short-lived. This second collection shows much more resemblance to C. bellioides than does the type, and it may later prove desirable to consider it simply as a variety of that species. Additional specimen examined: COAHUILA: Sierra Mojada Mts., M. £. Jones 263, April 19, 1892 (US; mounted on same sheet with C. bellioides, Dr. E. Palmer 505). EXCLUDED SPECIES Distasis? concinna Hook. & Arn., Bot. Capt. Beechey’s Voyage suppl. Pp. 350. = Hrigeron concinnus (H. & A.) T. & G., Fl. N. A. 2: 174. 1841. Distasis? heterophylla Hemsley, Biol. Cent.-Amer. Bot. 2: 119. 1881. The type was collected at Jalapa, Vera Cruz, Mexico, by Thomas Coulter, no. 406 (in Kew Herb.; fragment and photograph in U. 8S. National Herb.). Described as having trifid or pinnatifid leaves, phyllaries nearly equal in 2 or 3 series, receptacle conical, and ray florets in 2—3 series, with very short ligules. All the species of Chaetopappa have simple, entire leaves, unequal phyllaries, flat or nearly flat receptacle, and ray florets in one series, with well developed ligules. The involucre of the Coulter specimen, consisting of subequal, lance-linear, hispid-pubes- cent phyllaries, would place it in Erigeron, a genus of which there are several Texano-Mexican species with convex to conical receptacle, while those species segregated under Achaetogeron but better retained in Erigeron have a paleaceous pappus. In any case, Distasis hetero- phylla cannot be retained in Chaetopappa as here defined. 1946| SHINNERS: CHAETOPAPPA 81 Keerlia mexicana A. Gray, Proc. Amer. Acad. 22: 422. 1887. = Astran- thium xylopodum Larsen, Ann. Mo. Bot. Gard. 20: 31. 1933. REFERENCES Blake, S. F. Notes on certain type specimens of American Asteraceae in European herbaria. Contr. U.S. Nat. Herb. 26: 227-263. 1930. (Distasis, PP. 233-234.) Chandler, M. T. W. Letter to Major William H. Emory, from Ft. Duncan, Dee. 1, 1852. Report on the United States and Mexican Boundary Survey, by William H. Emory. Vol. 1, pp. 80~85. 1859. Gregg, Josiah. Diary and letters of Josiah Gregg. Excursions in Mexico and California, 1847-1850. Edited by Maurice Garland Fulton. 306 pp. 1044. Larsen, Esther Louise. Astranthium and related genera. Ann. Mo. Bot. Gard. 20: 23-44. 1933. REVISION OF THE GENUS LEUCELENE GREENE Lioyp H. SHINNERS INTRODUCTION There is remarkable similarity in general appearance, and in details of the involucre, between Chaetopappa bellioides (A. Gray) Shinners and an- other small, asteraceous, dry ground plant of the southwestern United States and adjacent Mexico, which has been known variously as Aster ericaefolius Rothrock, Aster Leucelene Blake, Leucelene ericoides (Torr.) Greene or under the names of segregates from the original species. It hardly differs from Chaetopappa except in the entirely capillary pappus, and char- acteristic creeping root and often partially underground stem. Its assign- ment to the genus Aster is but the last of a series of misapprehensions as to its affinities. It was first placed in Inula, and then in Chrysopsis, on the erroneous supposition that the rays were yellow. Later it was transferred to the artificial genus Diplopappus, in the mistaken belief that the pappus was double. It differs from Aster (considering now only the subgenus Euaster, and postponing for later consideration the remaining miscellane- ous elements which make of Aster not a genus but a hodge-podge) in hav- ing an underground creeping root and often partly underground stem, in- stead of superficial rhizomes or caudex; the phyllaries are not herbaceous- tipped, the pappus bristles are relatively few and definite in number (usu- ally 25); and the plant regularly flowers in spring and again later in the season, instead of once and finally in summer or fall. It does not greatly resemble any of the true Asters, but instead, as already mentioned, shows some resemblance to Chaetopappa bellioides. SYSTEMATIC TREATMENT LEUCELENE Greene, Pittonia 3: 147-149. 1896. Normal spring form: Plants perennial, low (about 6-12 em. above ground at flowering time), forming beds of small to large dense separate tufts which are connected by horizontal or oblique creeping rootstocks usually 3-7 em. or more below ground (shallower in eroding soils). Root- stocks producing vertical or oblique and branching, slightly woody, sub- terranean stems, or short naked branches which give rise to subterranean stems; stems again dividing to form numerous above-ground branches, which in turn bear slender erect or ascending branchlets or peduncles; leafy branchlets continuing to appear and elongate during and after flower- ing (eventually producing secondary flower heads, except in very dry or otherwise unfavorable seasons or localities, when persisting as sterile 82 1946] SHINNERS: LEUCELENE 83 shoots). Subterranean stems at first covered with numerous, alternate, acute, triangular scales, these passing into leaves above ground; scales disappearing in age. Above-ground branches and leaves sparsely to densely strigose and granular. Leaves alternate, simple and entire, obtuse and apiculate, prominently hispid-ciliate. Lower primary leaves linear-oblanceo- late to elongate-spatulate, 7-20 mm. long by 0.6—2.0 mm. wide, loose and more or less widely spreading; leaves of younger shoots linear-oblanceolate or linear, from appressed and imbricated to loose and merely ascending or occasionally even spreading, the upper gradually smaller, the uppermost scarcely larger than the outermost phyllaries. Heads solitary and terminal on the slender branchlets. Involucres s.7— 7-0 mm. high, at first broadly turbinate or campanulate, ultimately opening out flat. Phyllaries loosely appressed and imbricated in about 4—7 series, the outermost 1.5—2.5 mm. long, less than half as long as the inner- most, the middle ones about 1 mm. wide; concave, lanceolate or oblong- lanceolate, aristate-acute, more or less appressed-pubescent, and at least toward the apex serrulate- or lacerate-ciliate; all chartaceous, green or toward the apex red or purple, with broad scarious margins and a slender yellowish or brownish midrib extending nearly to the apex and thickened and bluntly keel-like toward base. Receptacle flat or slightly convex, naked and glabrous, slightly roughened by the points of attachment of the achenes. Heads heterogamous and heterochromous. Ray florets 12~24, pistillate, not developing achenes of quite the size and fullness of those of the disk, ligulate; ligules about 4-6 mm. long, 1.0-1.5 mm. wide, oblong or oblong-lanceolate, 4-nerved, white (occasionally rosy or rosy violet in withering, very rarely colored from the first); tube about 3 mm. long, nar- rowly cylindric; styles 2-parted for about a fourth their length, the branches linear, obtuse or subacute, obscurely granulose in the terminal third but not appendaged. Disk florets 12-24 (as many as or fewer than the rays, rarely more), hermaphrodite, fertile; corollas yellow, about 4.5—5.0 mm. long, tubular, subeylindric, slightly expanded upward, sometimes with a discernible short (0.2-0.3 mm.) basal tube, but usually limb and tube not recognizably differentiated, the summit regularly s-lobed, the lobes del- toid- or lanceolate-oval, o:3-0.5 mm. long, erect or slightly spreading ; styles 2-parted for about a fifth their length, the branches flat and with thickened margins, tipped with a triangular, dorsally granular-scabrous appendage about a fourth as long, in all about 0.8 mm. long, slightly or not at all exserted in flower; anthers included, obtuse at base, tipped with a prominent, lanceolate, hyaline appendage about 0.3 mm. long. Achenes about 1.7-3.0 mm. long, s-nerved, subcylindric but often becoming 1r- regularly prismatic or compressed in maturing, contractd at summit and rather abruptly narrowed to a whitish carunculate base, the long hairs toward the summit appearing like a secondary short pappus. Pappus of disk and ray florets similar, of 25 (less often 20 or 30) scabrous, approxi- mately equal bristles (rarely one or two as little as one fourth as long as the longest), 4.5-5.5 mm. long, united at base in a ring on the summit of the achene, those of the disk florets equalling or slightly exceeding the corollas. 84 WRIGHTIA [Vou. 1, No. 2 Late season and other growth forms: As the season progresses, the re- latively long and broad lower leaves wither, the secondary very narrow- leaved branches elongate, and under favorable conditions there is a second flowering. The plants are taller, much looser, and greener, the leaves narrower and less pubescent, the involucres narrower and fewer-flowered (narrowly conical or turbinate-cylindric, with 7-16 disk florets and 8-16 rays), and the rays are slightly shorter (3.5-5.0 mm. long). In general aspect and to some extent in detailed technical characters, the plant thus becomes markedly unlike the spring form, and might readily be taken for a distinct species. Injured plants and those on eroding sites (a habitat in which the plant occurs very commonly) may send up few, much elongate, straggling branches with widely spreading, very narrow leaves, and sometimes with widely spreading instead of ascending branchlets. Some of the outer primary above-ground stems may be prostrate, with numerous side branches, and the creeping root may be shallower or even become partially exposed. If spring is late, the retarded plants may first appear in a tran- sitional form and pass almost directly into the summer form. Plants of transitional, summer, or erosion forms were the basis for the original Inula? ericoides Torr., for Aster ericaefolius var. tenuis A. Gray, Leucelene ericoides serotina Greene, and Leucelene arenosa Heller, and their direct synonyms. Plants of the compact, low, broad-leaved, very pubescent and granular spring form were the basis for Diplopappus ericoides var. hirtella A. Gray, Leucelene alsinoides Greene, Aster bellus Blake, and their direct synonyms. Type species: Inula? ericoides Torr. [=Leucelene ericoides (Torr.) Greene]. The only variant of this highly polymorphic plant which possibly is to be considered a distinct variety or separate species is represented by a few specimens from northern and central Mexico, in which the leaves are fewer and more widely spaced than in the general run of specimens, and the lower have prominently long-attenuate petiolar bases. The scanty material available all belongs to the summer form; only a few separate stems are shown, with no complete tufts, and little or none of the root system. Their resemblance to the fragmentary type specimen of Aster ericaefolius var. tenuis is considerable, and at present I am unwilling to propose a new name for them. LEUCELENE ERICOIDES (Torr.) Greene, Pittonia 3: 148. 1806. Inula? ericoides Torr., Ann. Lyceum of Nat. Hist. N. Y. 2: 212. 1828. Chrysopsis ericoides “‘T\orrey] & J[ames]’’; Eaton, Man. ed. 5, p. 174. 1829. Eucephalus ericoides (Torr.) Nutt., Trans. Amer. Philos. Soc. n. s. 7:299- 1840 (dated 1841). Diplopappus ericoides (Torr.) T. & G., Fl. N. A. 2: 182. 1841. Diplopappus ericoides var. hirtella A. Gray, Pl. Fendl. (Mem. Amer. Acad. N. S. 4): 69. 1840. 1946] SHINNERS: LEUCELENE 85 Aster ericaefolius Rothrock, Bot. Gaz. 2: 70. 1877. Aster ericaefolius var. tenuis A. Gray, Syn. Fl. N. A. 1 pt. 2: 198. 1884. Leucelene ericoides ‘“‘variety or subspecies’’ serotina Greene, Pittonia 3: 140. 18096. Leucelene ericoides var.? tenuis (A. Gray) Greene, Pittonia 3: 149. 1896. Leucelene arenosa Heller, Cat. N. Amer. Pl. ed. 1, p. 143. 1898. (Nomen nudum.) Validated by Rydberg, Fl. Colo. (Colo. Agr. Exp. Sta. Bull. roo) p. 358, 1906, by two lines of key and citation of ‘Aster ericaefolius tenuis A. Gray”’ as synonym. Leucelene alsinoides Greene, Pittonia 4: 99. 1890. Leucelene hirtella (A. Gray) Rydb., Bull. Torr. Bot. Club 33: 153. 1906. Leucelene serotina (Greene) Rydb., Bull. Torr. Bot. Club 33: 153. 1906. Aster bellus Blake, Proc. Biol. Soe. Wash. 35: 174-175. 1922. Aster hirtifolius Blake in Tidestrom, Fl. Utah & Nevada (Contr. U. S. Nat. Herb. 25): 562. 1925. Aster Leucelene Blake in Tidestrom, Fl. Utah & Nevada (Contr. U. S. Nat. Herb. 25): 562. 1925. Aster arenosus (Heller) Blake, Journ. Wash. Acad. Sci. 30: 471. 1940. A common and rather weedy little plant of the Southwest and northern Mexico, from California, Utah, Wyoming, Nebraska, and Texas to Coa- huila, San Luis Potosi, Durango, and Sonora. Found especially on eroding slopes, roadsides, and open ground,.in sand, silt, or rocky soil. The type, a fragment consisting of part of a branch with a single head, representing the transitional or summer form, was collected by Dr. Edwin James in Long’s expedition to the Rocky Mountains in the summer of 1820, The locality is given by Torrey as “on the Canadian?” The plant may well have come from the Texas Panhandle, where it is very common. The type specimen is in the herbarium of the New York Botanical Garden. Some additional specimens examined: CHIHUAHUA: Chihuahua, Harde LeSueur, Oct. 10-15, 1935 (T). Majalea, LeSueur, Aug. 20, 1935 (T). District of Guerrero south of Basuchil, about ro miles n.w. of Mifiaca, Ynes Mexia 2520 (C; densely strigose form, matching the type of Aster bellus Blake). COAHUILA: Parras, C. A. Purpus, March, 1905 (C). Saltillo and vicinity, Dr. E. Palmer 48 in 1808 (C; doubtfully this). DURANGO: Tepehuanes, Dr. E. Palmer 41 in 1906 (C; densely strigose form). SAN LUIS POTOSI: Chareas, C. L. Lundell 5066 (USNA; doubtfully this). SONORA: s. of Nogales on road to Hermosillo, T. C. Frye & E. M. Frye 2261 (C). ARIZONA: Apache Co., near St. Johns, C. Hope 9335 (C). Cochise Co., 5 miles w. of Benson, R. C. Foster & J. F. Arnold 185 (G); Douglas, Leslie N. Goodding 2262 (C); near Douglas, Goodding 2235 (G); Ft. Huachuca, Dr. T. E. WwW ilcox, April, May, and August, 1894 (C; 3 collections), Dr. E. Palmer 436 in 1890 (G); Lowell, W. F. Parish 100 (G). Coconino Co., Cameron, Herbert C. Hanson A54 (T); vicinity of Flagstaff, Dr. D. T. MacDougal 43 (C, G); 14 miles w. of Navajo Bridge, K. M. Wiegand & M. C. Wiegand 2363 (G). Gila Co., near Rock and Rye Creeks, Mrs. Rose E. Collom 78 (G) apparently the same with- out number, dated May 1, 1938, USNA); near Rock Creek, Collom 485 (C). Mohave Co., between Hackberry and Peach Springs, Susan Delano Mc- Kelvey 2264 (G); Peach Springs, Norman C. Wilson, May 4, 1893 (C). Navajo Co., Holbrook, Ivar Tidestrom 12788 (C, USNA). Pima Co., Santa Catalina 86 WRIGHTIA [VoL. 1, No. 2 Mts., Mr. & Mrs. J. G. Lemmon 117 (C). Pinal Co., Oracle Junction, R. H. Peebles 6840 (USNA). Santa Cruz Co., Crittenden, T. S. Brandegee, May 25, 1892 (C); Nogales, Brandegee, May, 1892 (C); n. of Sonoito, Foster & Arnold 149 (G); s. of Tubac, J. H. Ehlers & Lois S. Ehlers 7246 (USNA). Yavapai Co., Ash Fork, Henry H. Rusby 688 (C); Clemenceau, Wyatt W. Jones, April 7, 1921 (C); n. of Humboldt, Tidestrom 12701 (USNA); Prescott, Aven Nelson 10260 (G). CALIFORNIA: San Bernardino Co., Barnwell, Katherine Brande- gee, May, ro11 (C); 5 miles s. of Barnwell, P. A. Munz 13709 (C); New York Mts. near Leastalk, S. B. Parish 10321 (G); Fourth of July Canyon, New York Mts., Annie M. Alexander & Louise Kellogg 1310 (C, G). COLORADO: Archuleta Co., Arboles, C. F. Baker 693 (C, G); Pagosa Springs, H. Bethel, E.S. Willey & I. W. Clokey, June 29, 1921 (C-Cl). Chaffee Co., Salida, C. F. Baker, F. S. Earle & S. M. Tracy 17 (C, C-Cl, G). Fremont Co., Canon City, T. S. Brandegee 552 (C). Garfield Co., Glenwood Springs, Mrs. Wislizenus, Aug., 1892 (G). Jefferson Co., 1 mile w. of Leyden, A. A. Beetle 2083 (G). Mesa Co., Grand Junction, C. S. Crandall, May 28, 1894 (C). Montrose Co., Cimarron, Baker 273 (C, G); Naturita, E. Payson 283 (G). Ouray Co., Ridg- way, E. B. Payson & Lois B. Payson 3848 (G). Pueblo Co., Pueblo, R. W. Woodward, Oct., 1882, and May 25, 1883 (G). Saguache Co., w. of Villagrove, Francis Ramaley & K. Richard Johnson, July 9, 1935 (T). KANSAS: Ellis Co., Ellis, Dr. L. Watson, in 1874 (G). Lane Co., 4 miles n. of Healy, Earl Bondy 552 (G). Logan Co., without locality, A. S. Hitchcock, Pl. of Kans. 240 (G). Norton Co., Almena, Rev. J. M. Bates 4550 (G). Osborne Co., within 5 miles of Osborne City, C. L. Shear 24 (C, G). NEBRASKA: Franklin Co., Franklin, H. Hapeman, May 27, 1938 (USNA). NEVADA: Clark Co., Kyle Canyon, Charleston Mts., Clokey 7738 (Pl. Exs. Gray. 785, as Aster hirtifolvus Blake) (C, C-Cl, G, SMU, USNA); Kyle Canyon at junction with Deer Creek Road, Clokey 7347 (C, G); below Wheeler Wells, Clokey 7348 (USNA); Hidden Forest, Sheep Mts., Alexander & Kellogg 1555 (C, G); Mormon Wells, Sheep Mts., S. G. Jewett 160 (USNA); Timber Mt., 15 miles nw. of Search- light, Percy Train 1475 (USNA); Searchlight, J. Stirting (C); Trout Canyon, 20 miles n.n.e. of Stump Springs, Pahrump Valley, Clark Co.-Nye Co., / LaRivers & N.F. Hancock 416 (USNA). Esmeralda Co., Gold Mts., Purpus 5945 (C; densely strigose form); Palmetto Range, Purpus 5906 (type of Aster bellus Blake, US; isotype, C; densely strigose form). Eureka Co., Italian Creek, Eureka, Train, June 14, 1936 (USNA). White Pine Co., 2 miles e. of Lehman Cave, Bassett Maguire & R. J. Becraft 2838 (C; same, labeled “Baker to Lehman Cave,” G); 3 miles s. of Ruth, B. O. Moore & G. E. Franklin 347 (USNA). NEW MEXICO: Bernalillo Co., near Albuquerque, H. J. Palmer 31174 (G); 2 miles e. of Albuquerque, Dr. Alfred L. Kammerer 50 (T). Chaves Co., 5 miles e. of Roswell, N. A. Palmer 25 (USNA). DeBaca Co., 103 miles n.e. of Dunlap, V. L. Cory 37591 (G). Dona Ana Co., Organ Mt. Water- shed, w. slope, L. D. Love, April 7, 1936 (USNA). Eddy Co., Carlsbad, Cory, April, 1924, and April 25, 1925 (TAM); Carlsbad Caverns National Park, Perry Convis 6 (USNA). Grant Co., Bear Mt., 5 miles from Silver City, Carl B. Wolf 2626 (G); Mangas Springs, 18 miles n.w. of Silver City, 0. B. Metcalfe 1468 (G); Silver City, W. W. Eggleston 16432 (G); Santa Rita Mt., s. end of the Black Range, Metcalfe 1468 (G); 2 miles e. of Separ, Simon E. Wolff 1717 (TAM). Guadalupe Co., Santa Rosa, Eula Whitehouse, Sept. 3, 1929 (T); Vaughn, Lewis S. Rose, July 28, 1919 (C). Hidalgo Co., near Antelope Station, M. W. Talbot 1210 (USNA). Lincoln Co., Gray, Josephine Skehan 4 (C, @), 1946| SHINNERS: LEUCELENE 87 S.F. Earle & Esther 8. Earle 156 (C). McKinley Co., 2 miles w. of Mexican Springs, H. J. Helm 2 (USNA); near Klag-e-toh, Navajo Reservation, long. 109°15' lat. 35°45’, C. P. Starr 15 (USNA). Rio Arriba Co., 13 miles n.w. of Conjilon, Wolf 2937 (G); vicinity of Lake Burford, A. Wetmore 493 (G). San Miguel Co., 2 miles w. of Las Vegas, Francis Drouet & Donald Richards 3228 (G); Las Vegas-Hot Springs, W. C. Sturgis, May 5, 1902 (G); near Pecos, P. C. Standley 4898 (G). Santa Fe Co., Santa Fe, J. T. Rothrock 14 (G). Socorro Co., Mogollon Mts., on Mogollon Creek, Metcalfe 324 (G). Torrance Co., without locality, F. C. Werkenthin, June 7, 1916 (T). Valencia Co., 7 miles n. of Trechado, Hugh C. Cutler 2089 (G). OKLAHOMA: Beaver Co., 15 miles s.w. of Beaver City, G. W. Stevens 361 (G); 5 miles w. of Gate, G. J. Goodman 2399 (G); near Knowles, Stevens 346.1 (G). Beckham Co., Sayre, Delzie Demaree 12447 (G). Blaine Co., 8 miles s. of Watonga, Goodman 2383 (G). Cimarron Co., 17 miles w. of Boise City, U. T. Waterfall 3142 (G); near Kenton, Stevens 480 (G). Ellis Co., near Shattuck, R. L. Clifton, 3125, 31663 (G). Harmon Co., near Hollis, Stevens 1053, 11653 (G). Harper Co., near Hornbeck’s, Stevens 257 (G). Major Co., near Waynoka, Slevens 607 (G). Texas Co., near Camp, Stevens 398, 40634 (G); n. of Guymon, Quentin Williams 13 (T). Woods Co., near Cora, Stevens 222 (G). TEXAS: Armstrong Co., 33 miles s. of Claude, L. H. Shinners 8116 (SMU). Brewster Co., Alpine, Barton H. Warnock, March 3, 1938, and Aug. 18, 1938 (T); 55.4 miles s. of Alpine, H. B. Parks & V. L. Cory 18633 (TAM); Green Valley, Glass Mts., Warnock 270 (T); Chisos Mts., C. H. Mueller 8196 (G; apparently the same, without number, T); 19 miles e. of Marathon, C. L. Lundell & Amelia A. Lundell 14207 (SMU). Briscoe Co., Quitaque, B. C. Tharp, April 28, 1934 (T). Carson Co., 5 miles n. of Panhandle, Shinners 8112 (SMU). Concho Co., Concho, J. Reverchon, Curtiss’s N. Amer. Pl. 1241 (G; isotype of Leucelene alsinoides Greene). Crane Co., 6 miles s. of Crane, Tharp, July 11, 1941 (T). Culberson Co., foothills of the Guadalupes, 5 miles n.e. of Pine Springs, Waterfall 5273 (G); 1 mile n. of Kent, W. B. Davis, Lint Robertson, & L. A. Smith 14 (TAM); 9 miles e. of Van Horn, Waterfall 4014, 5336, (G), 8 miles n. of Van Horn, Waterfall 4074 (G); 183 miles s.w. of Van Horn, Cory 44077 (T). Dallam Co., 7 miles n.w. of Dalhart, Shinners 8174 (SMU). Donley Co., 8 miles s. of Clarendon, F. A. McArty, April 27, 1944 (T); Jericho, Demaree 12435 (T). Edwards Co., Texas A. & M. Substation 14 (s. of Sonora, Sutton Co.) Cory 37072, 39026 (G), 48753 (SMU). El Paso Co., El Paso, Geo. Thurber 189 (G), Whitehouse, April 13, 1932 (T); Frontera near El Paso, Charles Wright, April 1, 1852 (SMU); Hueco Mts., Whitehouse, March 27, 1932 (T). Hansford Co., 5 miles s.e. of Gruver, Shinners 8228 (SMU). Hartley Co., 15 miles s.e. of Dalhart, Shinners 8166 (SMU). Hemphill Co., 5 miles s. of Canadian, Shinners 8277 (SMU). Hudspeth Co., e. side of Eagle Mts. 63 miles s. and 1} miles w. of Hot Wells, Waterfall 4953 (G); Eagle Springs near Sierra Blanca, Whitehouse, April 22, 1932 (T); 3 miles e. of Sierra Blanca, Waterfall 4953 (G). HutchinsonCo., 2 miless. of Borger, Shinner 8096 (SMU). Jeff Davis Co., Davis Mts., M. S. Young, Aug. 20, 1914 (T); Goat Canyon, Mt. Livermore, Davis Mts., L. C. Hinckley, Aug. 22, 1935 (T); Ft. Davis, Whitehouse, April 20, 1932 (T); White Rose Pass, Davis Mts., Cory 18906 (USNA; the local name for this place is Wild Rose Pass). Lubbock Co., 5 miles e. of Idalou, Shinners 8359 (SMU); Lubbock, Demaree 7514 (G), Tharp, Sept. 2, 1920 (C-Cl, T); Landrew’s Ranch, Lubbock, Demaree 7683 (G); Slaton, Whitehouse 10523 (SMU). Midland Co., Midland, Parks & Cory 12494 (TAM). Mitchell 88 WRIGHTIA [Vou. 1, No. 2 Co., 2 miles n.w. of Colorado City, Shinners 8415 (SMU); Renderbrook Pasture, Spade Ranch (sw. } sec. 28, Lavaca Nav. Co. Block 19), Richard W. Pohl 4482 (SMU). Moore Co., 13 miles s. of Dumas, Shinners 8144 (SMU). Nolan Co., Sweetwater, Mrs. V. E. Stanfeld, March 22, 1928 (T). Ochiltree Co., 8 miles s.s.e. of Perryton, Shinners 8254 (SMU). Oldham Co., w. of Adrian, Lundell & Lundell 11464 (SMU); 4 miles n.w. of Magenta, Shinners 8162 (SMU); 3 miles n. of Vega, M. W. Howard 15 (USNA). Pecos Co., 2.2 miles w. of Longfellow, Parks & Cory 18633 (TAM, USNA); 9 miles w. of Ft. Stock- ton, Lundell & Lundell 10195 (SMU); 21 miles e. of Ft. Stockton, Cory 40308 (G, T); Pecos to Ft. Stockton on Highway 285, about 15 miles from Ft. Stockton, Whitehouse 10531 (SMU). Potter Co., 16 miles n. of Amarillo, Goodman 2445 (C, G.); 15 miles n.e. of Amarillo, Shinners 8049 (SMU). Presidio Co., 113 miles s.e. of Casa Piedras, Parks & Cory 26408 (TAM, USNA); between head of McComb’s and Musgrave Canyon, Tierra Vieja Mts., Hinckley 2066 (SMU; apparently the same, without number, T); Shafter, Hanson 629 (G). Randall Co., branch of Palo Duro Canyon, Young, Sept. 4, 19017 (T); 103 miles e. of Canyon, Parks & Cory 13541 (TAM); 123 miles e. of Canyon, Shinners 8042 (SMU). Reagan Co., Big Lake, Parks & Cory 18943 (TAM). Reeves Co., w. of Pecos, Tracy & Earle 103 (G, T). Scurry Co., Roscoe to Snyder, Whitehouse 10220 (SMU). Sherman Co., 25 miles e. of Stratford, Shinners 8209 (SMU). Sterling Co., Gasconade Creek, sec. 32, SPRR Block 15, Pohl 4792 (SMU). Taylor Co., Camp Barkeley, W. L. Tolstead 6963 (C; without number, G), March 20, 1943 (G). Upton Co., 8 miles e. of Rankin, Tharp, July 11, 1941 (G, SMU, T). Ward Co., 10} miles s. of Monahans, Cory 27333 (USNA). Wichita Co., 7} miles s.w. of Iowa Park, Parks & Cory 18251 (TAM, USNA); 6.7 miles w. of Kadane Corners, under mesquite, Whitehouse 9692 (SMU), same locality, along roadway, ‘looked slightly different,’ Whitehouse 9695 (SMU). UTAH: Carbon Co., Price, Susan G. Stokes, June 20, 1898 (C); 8 miles n. of Wellington, Maguire 18569 (G). Daggett Co., Grouse (Summit) Creek Canyon (Jackson Draw), Uinta Basin, Edward H. Graham 8114 (C-Cl, G, USNA). Emery Co., San Raphael River, B. F. Harrison 8065 (C). Garfield Co., Bryce Canyon, F. A. Thackeray 559 (C-Cl, USNA); Panguitch, Eggleston 8125 (USNA). Grand Co., above Mill Creek, La Sal Mts., Bassett Maguire & B. L. Richards Jr. 15722 (G). Iron Co., 1 mile n. of Bear Creek, Dixie National Forest, Grazing Reconnaissance Crew, June 24, 1936 (C); Modena, Goodding 999 (G). Juab Co., base of mountains e. of Nephi, J. Arthur Harris 261 (G). Kane Co., near Willow Tank, Harrison 9115 (USNA). Salt Lake Co., near Bingham Canyon, A. O. Garrett 2275 (G); Ft. Douglas, Mrs. J. Clemens, July 21, 1911 (C-Cl); Salt Lake City, Garrett 8591 (SMU). San Juan Co., 9 miles nw. of Blanding, Bassett Maguire & J. D. Redd 2164 (C); 1 mile w. of Lockerby, Cutler 2739 (G); e. of Monticello, P. A. Rydberg & A. O. Garrett 9197 (C). Uinta Co., Red Wash just n.w. of mouth of Split Canyon, above Island Park, Graham 9163 (G, USNA). Washington Co., St. George, Goodding 782 (G); Springdale, M. E. Jones, May 14, 18094 (C).- Wayne Co., w. of Fruita, Capitol Reef National Monument, Annetta Carter 1543 (C); Natural Bridge near Fruita, Harrison 7379 (USNA). Also reported from Wyoming by Aven Nelson in Coulter and Nelson’s New Manual of Rocky Mountain Botany, p. 522, r909, with the observation that the genus is “essentially monotypic.” The following two type col- 1946| SHINNERS: LEUCELENE 890 lections have also been examined, but for want of adequate or reliable data have not been cited above: NEW MEXICO: without locality, A. Fendler 348 (type of Diplopappus ertcotdes var. hiretella A. Gray, G; isotype, C; in the original description only one number is cited, but the plant is said to grow “‘on sides of ravines in arid places, Santa Fe; also on the Rio del Norte; May, June’’). Without locality, Wright 1171 (type of Aster ericaefolius var. tenuis A. Gray, G; isotype, C; fragments, mounted with other fragmentary collections). EXCLUDED SPECIES Leucelene alpina (Nutt.) Greene, Fl. Franciscana pt. 4: 384. 1807. = Lon- actis alpina (Nutt.) Greene, Pittonia 3: 245. 1897. (Also known as Aster scopulorum A. Gray.) THE GENUS DICHAETOPHORA A. GRAY AND ITS RELATIONSHIPS Luoyp H. SHINNERS INTRODUCTION As has already been noted under Chaetopappa and Leucelene, the nature of the pappus does not offer a very reliable clue to the relationships of many genera of the Composirak. Among the heterochromous ASTEREAE, a somewhat better, though still imperfect, arrangement of the indigenous genera may be made according to the form of the receptacle. In Boltonia, Dichaetophora, Astranthium, Aphanostephus, Egletes, and some species of Erigeron, the receptacle is hemispherical to conical, instead of flat or merely slightly convex. This sets off a somewhat miscellaneous group of genera which, though not all obviously closely related, at least do not show closer affinities elsewhere in the tribe, except for the species of Hrigeron, and perhaps Boltonia. A further small but possibly significant point of re- semblance among them lies in the manner of wilting of the rays, which remain extended or twist irregularly in withering, or are quickly deciduous, without coiling. In Chaetopappa, Leucelene, Aster, Machaeranthera, Sider- anthus, Isopappus, Chrysopsis, and probably a majority of the ASTEREAE, the rays curl under from tip toward base, unless they are extremely short or have not first become fully expanded. The heads will often appear discoid when beginning to go to seed, the curled rays having become quite inconspicuous. Dichaetophora was placed under Boltonia by Bentham and Hooker, chiefly because of the flattened, broadly winged, ribless achenes with pappus of squamellate awns, nearly identical in the two genera. In many other significant points, Dichaetophora is closer to Astranthium, showing particular resemblance to A. integrifolium (Michx.) Nutt. In Boltonia, the phyllaries are unequal, imbricated in 3—6 series, somewhat coriaceous ex- cept on the margin, with prominent and sometimes almost keel-like mid- vein; the involucre is narrowed and more or less pointed at base; the disk corollas are not very abruptly contracted to form the tube; the pubescence of the achenes is not glandular; and the plants are typically rather tall, much-branched perennials. In Dichaetophora campestris and Astranthium integrifolium, the phyllaries are equal in about 2—3 series, rather thin and flat; the involucre is broad and shallow; the disk corollas are very abruptly contracted to form a short basal tube, and abruptly enlarged at the at- tachment to the achene;' the achenes bear peculiar gland-tipped hairs; ‘ This is so indistinctly shown in Miss Larsen’s figure (pl. 2, fig. 9) as to be rather misleading. Her sketch of a ray floret (fig. 8) shows it with the type of style found only in the disk florets. go 1946] SHINNERS: DICHAETOPHORA gt and the plants are low annuals, with solitary heads terminating long naked peduncles. The ribless achene of Astranthium is not winged, but the vesti- gial squamellate pappus forms a small elliptic ring in the middle of the summit, reminiscent of Dichaetophora. There is remarkable mimicry in superficial appearance between small plants of Dichaetophora and Astranthium, and small plants of some species of Aphanostephus. But Aphanostephus is set off by a number of marked differences in detail. The phyllaries are unequal and imbricated in about 3-4 series; the tube of the disk corollas is peculiarly thickened or swollen below the middle or at base; the achenes are columnar and truncate at both ends or cupped at summit, with incomplete superficial ridges, though not true ribs; the pappus forms a ring around the whole broad summit of the achene, and the pubescence of the achene, if present, is not glandular. An often useful means of distinguishing young plants is the direction of the pubescence on the peduncles. In Aphanostephus it is horizontally spreading to downwardly appressed; in the other two it is widely spread- ing, ascending, or upwardly appressed. The species of A phanostephus are low annuals or perennials, with broad shallow involucres, and outer phyl- laries a third to three-fourths as long as the inner, unlike the narrower pointed involucre with very short outermost phyllaries found in Boltonia, and the achenes of the two genera, as already described, are quite dissimilar. In Erigeron tenuis T. & G., the receptacle is occasionally so strongly convex as to appear nearly conical in age, forming a transition to several species of west Texas and Mexico having a decidedly conical receptacle. There is an independent series of variations in pappus, from the wholly capillary simple one of EL. philadelphicus L., through the capillary double pappus of FH. pumilus Nutt., or the partly squamellate double pappus of £. Bigelovii A. Gray and EF. coronarius Greene, to the wholly squamellate pappus of those species (mostly with flat or merely convex receptacle) placed under Achaetogeron, and including forms in which the pappus is quickly deciduous or even quite wanting. In all of them the phyllaries are uniformly very narrow and all of similar shape, in 2 or 3 equal series, or an outer row of equal phyllaries only slightly shorter than the inner; the rays are numerous, narrow (sometimes also very short), and in 2-3 series (except in a few species with flat or convex receptacle); and the achenes have 2 or more true ribs, prominent from base to summit. The genus Egletes is characterized by a conical receptacle, like that of Boltonia, Dichaetophora, Astranthium, and Aphanostephus, and achenes with true ribs, like those of Erigeron. In most other respects, it shows relatively little resemblance to any of these genera. The phyllaries are broad, unequal, usually without distinet midvein, in several series, forming an involucre greatly resembling that of Pluchea in the INULEAR-PLUCHEINAE. There is further resemblance in the multiseriate reduced marginal pistillate florets of most of the species, suggesting what one might expect to have been an ancestral form of the head of Pluchea, in which only a few central florets are perfect and well developed, the more numerous marginal pistil- Q2 WRIGHTIA [Vox. 1, No. 2 late ones having very reduced filiform corollas. The frequent presence of viscous or glandular hairs, and the largely tropical distribution of both genera, also suggest possible relationship. The following key will outline the more important differences between the indigenous North American genera of heterochromous ASTEREAE having a hemispherical or conical receptacle. 1. Achenes with true ribs extending from base to summit; marginal pistillate corollas (often very reduced) in 2 to many series.................--- 2. 2. Phyllaries broad (lanceolate to oval), usually without definite mid- vein unless toward the base, unequal, in 3~5 series; vegetative parts often viscid or glandular and aromatic; chiefly tropical plants of lowlands, South America north to Mexico and Texas (lower Rio treatide) and in the: Weel indies: 265k ee ee: Egletes. 2. Phyllaries narrow (rather narrowly lanceolate to lance-linear), the midvein usually distinct from base to tip, in 2 or 3 equal series, or an outer row of equal phyllaries slightly shorter than the inner; vegetative parts not viscid or glandular and aromatic; chiefly temperate zone plants, the Mexican species montane, with a few RINNE is ee as Shes en ees Erigeron (in part). 1. Achenes without true ribs, at most with incomplete superficial ridges; marginal pistillate corollas i in 1-2 series, conspicuously radiate........ +: 3. Achenes columnar, quadrate or terete, truncate at both ends or cupped at summit; pappus forming a ring around the whole broad summit of the achene: pubescence of peduncles horizontally spread- ing or reflexed, never upwardly appressed........ Aphanostephus. 3. Achenes more or less flattened, broader than thick; pappus in a small elliptic ring in the center of the narrowed summit of the achene; pubescence of peduncles widely spreading to upwardly appressed, MOTEL -FOCR On sk ad ah ce i oe 4. i Eevee ee OL 10 ae Borie ee ee he ge ee Ce 5. Achenes not winged; pappus very short or obsolete. ..... ey Oe oe tee eee ore A stranthium. 5. Achenes winged; pappus of two prominent squamellate awns and several very small ones........ Dichaetophora. 4. Phyllaries unequal, imbriecated in about 3-6 series, the outermost at maturity not more than a third as long as the innermost. . - . sek 0h ee eee eae eC AN ha Oh aa ees Boltonia. SYSTEMATIC TREATMENT DicuartorHora A. Gray, Plantae Fendlerianae (Mem. Amer. Acad. 1.5. 4): 73-74. 1849. Boltonia sect. Dichaetophora (A. Gray) Bentham & Hooker, Gen. Plant. 2: 269. 1873. Annual from a slender taproot. Stems usually several or many, suberect or decumbent at base, 4-24 em. tall, simple or sparingly forked, term1- nating in long peduncles naked for 2-9 em. below the heads when in full flower or fruit; sparsely to rather densely pubescent with spreading, as- 1946| SHINNERS: DICHAETOPHORA 93 cending, or upwardly appressed hairs. Basal leaves (often withering before the plant has finished flowering) oblanceolate, obtuse, entire; the blades 1.0-2.3 em. long, 4-12 mm. wide, the narrow petiolar base 0.6-2.3 em. long. Stem leaves alternate, similar to the basal ones but smaller, the upper gradually reduced, the uppermost bract-like ones narrowly oblong or lance- linear, 4-16 mm. long, t.o-2.5 mm. wide. Heads solitary and terminal on the stems or branches. Involucres 3.0— 4.5 mm. high, broadly campanulate to saucer-shaped; phyllaries in two series, equal, lanceolate, acute, about o.6—-1.2 mm. wide; thinly chartaceous or membranaceous, green except for the scarious margin, sparsely and somewhat hispidly appressed pubescent on the back, serrulate-ciliate on the margins and weakly spine-tipped. Receptacle conical to subglobose, naked, glabrous, slightly pitted and roughened by the points of attach- ment of the achenes. Heads heterogamous and heterochromous. Ray florets about 16-24, uniseriate, pistillate, fertile, ligulate; ligules white, rarely lilac-tinged, especially on the back, 5-9 mm. long; tube about o.5 mm. long; style branches flattened, linear, obtuse, o.4 mm. long. Disk florets hermaphrodite, fertile; corollas yellow, tubular, about 2.5 mm. long, abruptly constricted just above the flared and slightly thickened base to form a very short tube about o.2 mm. long, the cylindric-campanulate limb (making up most of the corolla) equally 5-lobed at summit, the lobes about 0.6 mm. long. Style branches flattened, 0.6 mm. long, tipped by a narrowly acute, dorsally granular-scabrous appendage making up a third of the total length. Anthers rounded at base, tipped by a narrowly acute hyaline appendage about o.25 mm. long. Achenes appearing nearly flat, oval or orbicular in outline: the thickened center lanceolate in outline, densely pubescent with ascending, whitish, glandular-capitate hairs, surrounded by a broad coriaceous wing-margin equal in width to the central portion at either side, narrowed at top and bottom, glabrous on the faces, densely fringed with glandular-capitate hairs; total width of achene 1.6—-2.0 mm. Pappus consisting of two ant- rorsely scabrous, erect or diverging, equal or slightly unequal awns o.7— 1.1 mm. long, near the middle of the truncate or slightly emarginate sum- mit of the achene, usually with several additional minute ones 0.25 mm. or less long, forming an elliptical ring with the long awns on either end, at the wing-margins. Ciliation on the margins of the achene often longer toward the summit and simulating a supplementary pappus. Small spring-flowering annuals (February-May, or stragglers occasion- ally later) of the Rio Grande Valley and tributaries in southern Texas and northern Mexico, growing in mesquite prairies, meadows, or other low, sandy or silty open ground. Type and only species: D. campestris A. Gray. To their Boltonia section Dichaetophora, Bentham and Hooker as- signed three species, one not named and none formally transferred to Boltonia, two being mentioned simply as Dichaetophora Gray, and Poly- 04 WRIGHTIA [Vou. 1, No. 2 actidium Seemannii Sch. Bip. Gray’s genus was restored to its original form by him in the Synoptical Flora, where Polyactidium is listed as a synonym of Erigeron. From the description given by Bentham and Hooker of the unnamed species based on Coulter’s no. 278 from California, and of Polyactidium Seemannii (“achaenia margine calloso-nerviformi,”’ “‘achaenia nerviformi-marginata,” and “statura elatior et ligulae numerosissimae Erigerontis sect. Phaenactidis’’), both plants are evidently species of Erigeron. DicHAETOPHORA CAMPEsTRIS A. Gray, Pl. Fendl. (Mem. Amer. Acad. n.s. 4): 73-74. 1849. Boltonia campestris (A. Gray) Benth. & Hook. ex Hemsley, Biol. Cent.-Amer. Bot. 2: 120. 1881. Of the three Mexican collections of Dr. Josiah Gregg cited by Gray in the original description of Dichaetophora campestris, that from Bishop’s Hill, near Monterey, Nuevo Leon, Feb. 6, 1847, has been selected as type (in Gray Herbarium, under number 48; an evident duplicate in the herb- arium of the Missouri Botanical Garden, under number 135). Additional specimens examined: CHIHUAHUA: valley of Rio Conchos, Gregg 499, April 21, 1847 (Mo; duplicate in Gray Herb. under number to). (Gregg’s diary of this date gives the locality as en route from Jimenez to Santa Rosalia, along the Rio Florida branch of the Conchos, 10 miles southeast of Santa Rosalia.) COAHUILA: Rio Grande Valley near Diaz, C. G. Pringle 8303 (C, G, Mo). Muzquiz, Ernest Marsh, spring, 1935 (T; mixed with Ap- hanostephus ramosissimus). NUEVO LEON: Monterey, Gregg 49 (G), Dr. E. Palmer 501 in 1880 (G). Battlefield between Citadel and Teneria (Tenorio?), Monterey, Gregg 134 (Mo). San Juan, Pringle 13720 (G). TEXAS: Atascosa Co., near Campbellton, Eula Whitehouse 10382 (SMU). Brooks Co., 4 miles s. of Falfurrias, K. M. Wiegand & M. C. Wiegand 2432 (G). Frio Co., Dilley, B. C. Tharp, Feb. 27, 1930 (T); Devine-Dilley, Tharp, April 6, 1930 (T); 11 miles w. of Dilley, L. H. Shinners 7411 (SMU). Jim Hogg Co., outside city limits of Hebbronville on Highway 50, Whitehouse 10383 (SMU). Maverick Co., Eagle Pass, V. Havard 16 (G), H. C. Hanson 368 (G, Mo, T). Webb Co., Laredo, J. Reverchon 4005 (Mo); near Bruni, F. A. Warren 1187 (USNA). REFERENCES Bentham, G., and J. D. Hooker. Genera Plantarum. Vol. 2, pt. 1. 1873. (Bol- tonia, p. 269.) Gray, Asa. Synoptical flora of North America. Vol. 1, pt. 2. 1884. (Dt- chaetophora, pp. 166-167.) Greene, Edward Lee. [Reduction of Achaetogeron to Erigeron.] Pittonia 2: 168-169. 1801. Gregg, Josiah. Diary and letters of Josiah Gregg. Excursions in Mexico and California 1847-1850. Edited by Maurice Garland Fulton. 396 pp. 1944- Larsen, Esther Louise. Astranthium and related genera. Ann. Mo. Bot. Gard. 20: 23-44. 1933. REVISION OF THE GENUS APHANOSTEPHUS DC. Lioyp H. SHINNERS INTRODUCTION It is eloquent testimony of the confusing variations in the genus that Aphanostephus was described as three different genera by the same author almost simultaneously, and that one of its first known species was described under different names in two genera in two different tribes; yet every one of its eleven species, in one or another of its growth forms, has been or could easily be mistaken for one or several of the others. About its generic status, however, there need be little question. It possesses the flattened, triangular-appendaged style branches of the disk florets characteristic of the AsTEREAE; it does not have the cone-tipped style branches of the An- THEMIDEAE, in which tribe De Candolle placed his synonymous genus Leu- copsidium. As mentioned in connection with Dichaetophora, it belongs to the small group of heterochromous ASTEREAE having a conical or hemis- pherical receptacle. It differs strikingly from all the genera to which it shows other resemblances in having peculiar, ribless, columnar achenes. In superficial appearance it suggests Hrigeron or Astranthium, but differs from them in having unequal phyllaries, as well as in the achenes. A minor characteristic, often helpful in assigning immature plants to the proper genus, is the widely spreading to downwardly appressed pubescence of the peduncles and stems. In the plants likely to be mistaken for it, excepting a few species of Erigeron, the pubescence varies from spreading to upwardly appressed. As in a number of other genera of the Southwest and Mexico, the species exhibit much ecological and especially seasonal polymorphy, while detailed technical differences show close convergence or overlap. Comparisons must be made only between properly comparable forms. Casual examination of an assortment of herbarium specimens assigned to several species may give the impression that an incoherent jumble has been divided at random into several equally incoherent jumbles, with no distinctions that hold good. Aphanostephus skirrhobasis (DC.) Trel., in normal spring form, at a stage when the earliest heads to flower have started to go to seed, is readily separable from equivalent plants of A. arizonicus A. Gray by the larger involucres, uneven pappus, and hardened, inflated corolla bases. But less mature plants, with slightly smaller involucres, less distinct pappus, and corolla bases not yet hardened or inflated, may easily be misnamed; so also might partly developed late-season plants, which produce secondary branchlets with smaller heads than those of the spring form. Very dwarf, unbranched, single-headed, early spring plants of the two species show 95 96 WRIGHTIA [VoL. 1, No. 2 searcely any pronounced differences in detail, and are closely similar in general appearance. A. arizonicus and A. ramosissimus DC. often, and other northern species less frequently, show signs of injury like that of grasses in which an insect larva has eaten through the culm inside an upper leaf sheath, causing the inflorescence to turn brown and wither before it has fully expanded. In plants of Aphanostephus, many of the involucres take on an ovoid shape, remain closed, and turn brown and dry, together with part of their peduncles. This abnormality seems to develop only in late spring and summer, and is especially frequent and striking on much branched plants with numerous heads. Specimens of such abnormal plants belonging to two or three different species, laid side by side, will seem very much alike; and if no fully mature normal heads are present, it will be hard to find any convincing grounds for calling them more than one species. As a further complication, it is most probable that crossing takes place between species. All of them usually occur in abundance, and several may grow together; they prefer eroding ground, blowouts, road- sides, and other disturbed places, which in such genera as Aster and Soli- dago are known to be especially favorable to the appearance of hybrids. Specimens have been collected, or found in herbarium material, which from appearance, locality of collection, and (when known) associated species, suggested crosses between Aphanostephus arizonicus and A. humilis (Benth.) A. Gray, A. arizonicus and A. skirrhobasis, A. Kiddert Blake and A. ramosissimus, and A. ramosissimus and A. skirrhobasis. Most unfortunately for the taxonomist, intermediates between A. ramosisst- mus and A. skirrhobasis are very hard to distinguish from A. arizonicus, except by geography. Both supposed parents are abundant and widespread, and in west-central Texas, where their ranges meet or overlap that of A. arizonicus, the number of plants of uncertain identity is especially high. An abundance of material and field observation have made it possible to untangle many of the perplexing variations in the United States, and treat them taxonomically with some assurance. Much less assurance is felt with regard to the more numerous species of Mexico. They evidently vary greatly with relation to the rainy season, which may come at different times in different parts of the country, so that calendar date by itself is not the simple and helpful clue that it is farther north. A majority of them apparently tend to be coarser and stouter than the northern species; at least, a good many specimens show only basal branches from what must have been a quite large and bushy plant, but there is no information as to habit or root system. A discouragingly large number of collections show out- of-season shoots from the bases of old plants, never very satisfactory for exact determination, and offering little but unalloyed difficulty, especially when accompanied by no information about habitat or about the plant itself. The untangling of the variations among the Mexican species has been fraught with much uncertainty, and has depended a great deal upon analogy with what is better known of the species of the United States. It has been found impossible to construct a simple dichotomous key on the 1946] SHINNERS: APHANOSTEPHUS 97 basis of available material; a synopsis in quasi-key form is offered as substitute. SYSTEMATIC TREATMENT APHANOSTEPHUS DC., Prodromus 5: 310. 1836. Keerlia DC., Prodr. 5: 309-310. 1836. [This genus merged with A phanostephus under the latter name by Asa Gray, Plantae Wrightianae Part I (Smith- sonian Contr. vol. 3 art. 5): 93, 1852, and Proc. Amer. Acad. 16:81-82, 188o.| Leucopsidium DC., Prodr. 6: 43. 1837. Aphanostephus subg. Euaphanostephus Blake, Contr. Gray Herb. 53: 24. 1918. (Dr. Blake’s second subgenus is retained as Section 1, below.) Annuals or perennials from taproots, or perennials forming clumps from vertical, ultimately obliquely branched, woody caudices. Leaves alternate, simple, entire to deeply pinnatifid, sessile or petioled, in a few species more or less clasping at base. Heads solitary and terminal on the branchlets; peduncles with widely spreading to downwardly appressed, but never with ascending or upwardly appressed hairs. Involucre saucer- shaped to broadly hemispherical or (especially in fruit) broadly urn- shaped; of narrowly lanceolate to oblong-oblanceolate, broadly acute to acuminate, chartaceous, scarious-margined phyllaries imbricated in about 3-5 series, the outermost about 13~%4 as long as the innermost. Receptacle depressed-hemispherical to conical, naked, roughened by the raised points of attachment of the achenes. Heads heterogamous, radiate, heterochro- mous, many-flowered. Rays uniseriate or sub-biseriate, pistillate, fertile, ligulate; ligules lance-linear or lance-oblong to elliptic, slightly emarginate or denticulate at the narrowed apex, white to lavender or rose-purple (most commonly white within, partly or wholly colored without), per- sistent, in drying becoming inrolled sideways or longitudinally furrowed, remaining extended instead of coiling from tip toward base; tube short; style branches linear or lance-linear, obtuse or subacute. Disk florets hermaphrodite, fertile; corollas yellow, tubular, narrowly funnel-form or with more or less distinet cylindric basal tube and urn-shaped limb, equally 5-lobed at summit; base of tube in some species with thickened or hardened walls, becoming slightly to greatly enlarged in fruit; style branches flat- tened, tipped with a short, broadly triangular, acute appendage; anthers obtuse at base, tipped with an acute, hyaline but centrally or basally thickened appendage. Achenes of disk and ray similar, columnar, subterete to quadrate, slightly expanded toward the summit, often curved (peculiar- ities of growth due to pressure on the conical or rounded receptacle), glabrous or sparsely pubescent, with 4—12 distinet to very obscure grooves or superficial ridges not differing from the rest of the achene surface in texture and usually not extending completely from base to summit (im- mature achenes may appear strongly ribbed longitudinally as a result of shrinkage in drying). Pappus an uneven scaly crown around the periphery of the broad summit of the achene, ciliate or the lobes sharp-pointed or prolonged into narrow awn tips, in all o.25-1.8 mm. long; or pappus a 98 WRIGHTIA [Vou. 1, No. 2 rather minute crown of equal hairs or ciliae o.1-0.25 mm. long. Pappus similar in disk and ray, or that of the ray shorter. Type species: A. ramosissimus DC. OUTLINE OF THE MEXICAN SPECIES OF APHANOSTEPHUS! 1. Rays 40-65 or more; plant a clump-forming perennial, the woody caudex becoming much branched, the divisions oblique or almost creeping; hairs of the peduncles rather coarse, hispid, from swollen bases, widely spread- ing, o.2-0.6 mm. long; lower leaves toothed to deeply pinnatifid, passing rather abruptly into the narrowly oblanceolate or lance-oblong to linear, entire middle and upper ones; plants of extreme northeastern Mexico Cicero Lean. COnnain) «i450 oe fe kk oo Bee 11. A. Riddellit. t. Rays 16-42; plant annual or perennial from a taproot; hairs of the peduncles relatively fine and soft, from slightly swollen bases, widely spreading to downwardly appressed, o.2—1.0 mm. long; lower leaves toothed to deeply pinnatifid, passing gradually into the smaller, less divided middle and REO OB ae a a ees oe ee cs oe a wee 2. 2. Middle phyllaries o.6—-1.1 mm. wide; phyllaries all with uniformly very narrow dark centers; achenes about 1.0-1.4 mm. long; rays 5.0-7.5 mm. long (see also key to U. 8. species below for further differences between A. ramosissimus and A. arizonicus)..............0006- ne 3. Rays 30-40; involucres 4.5-7.0 mm. high; plants of San Luis ROOGRy soho fe 4. A. potosinus. 3. Rays 16-32; involucres 3.6-5.0 mm. high; plants of Tamaulipas Rey ONG Se GG ek 5. A. ramosissimus. 2. Middle phyllaries 1.1-2.0 mm. wide; dark centers of the phyllaries varying greatly from very narrow in the innermost to broad in the middle and outer; achenes about 1.3-2.0 mm. long; rays 6-9 mm. long (the following five species could not be fitted into a oe WIE ee ee ne eee ee cia ee 4a. Plants of northern Mexico (Chihuahua, Coahuila; doubtiully Durango and Zacatecas), stems normally erect or ascending, at first simple, ultimately much branched; peduncles naked for 1.5-8.0 em.; hairs of peduncles widely spreading or slightly deflexed, 0.3-1.0 mm. long; uppermost leaves moderately to not at all clasping, o.5—1.5 em. long; disk corollas 1.8—2.5 mm. long; achenes about 1.5 mm. long............ 6. A. arizonicus. 4b. Plants of north-central Mexico (Durango to Guanajuato, doubtfully to Hidalgo); stems or main branches usually de- cumbent to ascending, usually not much branched except as a result of injury; peduncles naked for 0.3-3.5 cem.; hairs of peduncles o.2-0.5 mm. long, widely spreading to downwardly appressed; uppermost leaves more or less clasping, o.2—0.8 cm. long; disk corollas 2.6-3.0 mm. long; achenes 1.3—1.5 mm. long Gn PA ea ee eae Pe eee 7. A. humilis. 4c. Plants of southwestern and possibly central Mexico (Jalisco, doubtfully east to Queretaro) ; stems normally erect from curved bases, sparingly branched, the branches rather closely as- ‘Excepting A. skirrhobasis var. thalassius Shinners, coast of northeastern Tamaulipas. — ot. 1946] SHINNERS: APHANOSTEPHUS 99 cending; peduncles naked for 2.0-7.5 em.; hairs of peduncles 0.3~-0.8 mm. long, widely spreading to slightly deflexed; upper- most leaves with slightly broadened, rounded, clasping bases, 0.7~2.0 em. long; disk corollas about 2.9 mm. long; achenes £020 TN. LOOR eo 5 ae oe 8. A. jaliscensis. 4d. Plants of southwestern and south-central Mexico (Michoacan to Mexico State); stems ascending; peduncles naked for 1.5- 5.5 em.; hairs of peduncles usually rather sparse, o.2-0.6 mm. long, deflexed to downwardly appressed; uppermost leaves more or less clasping, 0.6-1.5 em. long; disk corollas about 2.8 mm. long; achenes about 1.5 mm. long; middle and inner phyllaries sharply acute to rather long-acuminate........ 9. A. ramosus. 4e. Plants of south-central Mexico (Orizaba region of Vera Cruz, Puebla, Tlaxcala, Mexico State, Hidalgo); stems numerous, prostrate with ascending tips, simple or sparingly branched; peduncles naked for 2-5 cm.; hairs of peduncles o.2-0.4 mm. long, widely spreading, deflexed, or downwardly subappressed, usually dense; uppermost leaves little or not at all clasping, 0.6-1.5 em. long; disk corollas about 2.5 mm.long;achenesabout 1.5 mm. long; middle and inner phyllaries rather broadly acute to. A. pachyrrhizus. KEY TO THE UNITED STATES SPECIES OF APHANOSTEPHUS This key is based chiefly on spring-flowering plants sufficiently ad- vanced for the earliest heads to have begun to form seed. Very dwarf, precociously flowering, unbranched plants of early spring show scarcely any distinguishing characters. Plants which have passed into a second summer- or fall-flowering phase lack the lower leaves, usually have more numerous, smaller, fewer-flowered heads, with fewer, shorter rays The perennial A. Riddellii T. & G. commonly flowers twice, like Leucelene ertcoides (Torr.) Greene, and is the one most likely to be collected in summer or fall. 1. Plant conspicuously hispid with coarse, jointed, translucent hairs, those of the stem and branches 0.7-2.2 mm. long (easily broken), spreading at right angles; heads somewhat crowded, the short peduncles naked for 3-12 mm., or in age for as much as 25 mm.; rays 12-18; plants of south- western Oklahoma and adjacent Texas................. 1. A. pilosus. 1. Plant soft-pubescent to hispid-pubescent, the hairs widely spreading to deflexed or downwardly subappressed, o.2-1.0 mm. long; heads not crowded, the peduncles naked for 1.5—10.0 cm., or in age as much as 15 ONL: TAYE 1666 OF TROTG. ec Brink eee ee i ee ees oes 7 2. Pappus uneven and scaly, the scales variously lacerate-ciliate, acute, or prolonged into awn-like tips, in all o.25-1.8 mm. long; corolla bases becoming whitened, hardened, and inflated in age to twice or more their original diameter By ie a ee ee ce 3. Involucres (of well- developed plants) 6.0-8.2 mm. high; phyllaries in about 4-5 series; rays about 8-15 mm. long, 1.5-2.3 mm. 100 WRIGHTIA [VoL. 1, No. 2 wide; mature achenes about 1.5—2.2 mm. long; pappus an ir- regular sealy crown, or of about 5 unequal, acute scales 0.25- TON oe ea iy sein Ne oes cole ees 4. 4. Plants of the interior, Texas to New Mexico, Kansas, Missouri, and Arkansas; various in habit and pubescence, but pubescence of involucres rather coarse and hispid... . Dee ie ee ee oh a ea 2a. A. skirrhobasis. 4. Plants of the Gulf Coast, Tamaulipas to Louisiana and Florida, occasionally extending inland on railroad ballast; typically low and spreading, densely covered with soft, gray, felty pubescence, that of the involucres finer and Srher Tuk it Ahe Species. os. fo es ig eae ess SRO rare Ge ae See 2b. A. skirrhobasis var. thalassius. 3. Involucres (4.5—)5.0-6.3 mm. high; phyllaries in about 3 series; rays broad for their length, 6-8 mm. long, 1.8-2.4 mm. wide; achenes about 1.5 mm. long; pappus typically of 5 or 10 acute, acuminate, or awn-tipped (sometimes also lacerate-ciliate) MURIOR Gy eM 1G ee 3. A. Kidderv. 2. Pappus a ring of very short, equal hairs or ciliae o.1-2.5 mm. long; corollas only slightly hardened or thickened, at the base or slightly MOON ee ee ee es a ee < 5. Rays 16-42; achenes 1.3-1.5 mm. long; hairs of peduncles rather fine and soft, from slightly swollen bases, sparse to dense, widely spreading to deflexed, 0.3-1.0 mm. long; lower leaves saliently toothed to deeply pinnatifid, passing gradually into the smaller and less divided middle and upper ones, the upper- most 0.5—2.0 em. long, 0.3-2.0 mm. wide; plants annual or PeVeDTIAL 010 BEAONIGES no ee ee oe ee 6. 6. Involucres 3.3-5.0 mm. high; middle phyllaries 0.6-1.1 mm. wide; achenes about 1.35 mm. long, with brown lines or low ridges; pappus about o.15 mm. long; plants of the Rio Grande Plain of southern Texas, less abundantly on the Edwards Plateau and north to the Panhandle (ap- parently hybridizes with A. skirrhobasis, A. Kiddert, and possibly A. arizonicus; hybrids with the former may closely resemble the last mentioned species).........-- PES ee re aR ee ee ere eee 5. A. ramosissimus. 6. Involucres 4.6-7.0 mm. high; middle phyllaries 1.1-1.8 mm. wide; achenes about 1.5 mm. long, rather prominently ridged or grooved; pappus about 0.25 mm. long; plants of Arizona, New Mexico, and western Texas (possibly hybridizes with This work is a part of that undertaken as a Fellow of the Guggenheim Memorial Foundation, 1946-47. 166 1947] CoRRELL: ORCHIDS OF TEXAS 167 HABENARIA CLAVELLATA (Michx.) Spreng., Syst. Veg. 3: 689. 1826; Correll in Lundell, Flora of Texas 3, 3: 159, pl. 3. 1944. Cass Co., 7 miles east of Hughes Springs, November 1, 1946, Eula White- house 17694. Harrison Co., boggy woods, Caddo Lake State Park, July 14, 1946, D. S. Correll 13225. Tyler Co., boggy woods, Angelina National Forest, east of Rockland, July 17, 1946, Correll 13305. HABENARIA CRISTATA (Michx.) R. Br. in Ait., Hort. Kew., ed. 2,5: 194. 1813; Correll in Lundell, Flora of Texas 3,3: 160, pl. 4. 1944. Jasper Co., seepage slope in forest, Magnolia Springs, July 18, 1946, D.S. Correll 13323. Nacogdoches Co., in deep woods, south of Nacogdoches, Sep- tember 24, 1938, Eula Whitehouse 11179. Shelby Co., Mangum Lake, July 22, 1940, H. B. Parks, Jr. RX 1803. Tyler Co., boggy meadow along stream bank, Angelina National Forest, east of Rockland, July 17, 1946, Correll 13296. HaBenaria FLAVA (L.) R. Br. in Spreng., Syst. Veg. 3: 691. 1826; Correll in Lundell, Flora of Texas 3, 3: 161, pl. 5. 1944. Harrison Co., rich moist hardwoods along a small stream, 3 miles west of Harleton, July 13, 1946, D. S. Correll 13199. HABENARIA LACERA (Michx.) Lodd., Bot. Cab. 3: pl. 229. 1818. Fig. 6. Orchis lacera Michx., Fl. Bor.-Am. 2: 156. 1803. Platanthera psychodes Lindl., Gen. & Sp. Orch. Pl. 294. 1835, exclude synonym Orchis psycodes. Platanthera lacera (Michx.) D. Don in Sweet, Hort. Brit., ed. 3: 650. 18309. Fimbriella lacera (Michx.) Farwell, Eleventh Ann. Rpt. Commissioners Parks and Boulevards Detroit 55. 1900. Blephariglottis lacera (Michx.) Farwell, Second Rpt. Mich. Acad. Sci. 42 (October 10, 1901); Rydberg in Britton, Man. Fl. North. States and Can., ed. 1: 296 (October 24, 1901). Plant glabrous, rather stout, 2.5-7.5 dm. tall (often propagating by means of root-shoots). Roots slender, fleshy, from thickened tuberoids. Stem somewhat ribbed, leafy below, bracted above. Leaves rather rigid, erect, oblong-linear to oblong-obovate or linear-lanceolate, with the basal part sheathing the stem, 7-21 em. long, 1.5—5 em. wide. Raceme loosely or densely flowered, 3-26 em. long, 3—4.5 em. in diameter; floral bracts usually equaling the pedicellate ovaries, rarely exceeding the flowers, narrowly lanceolate to linear-lanceolate, acuminate, 1-4 em. long. Flowers pale yellowish green or whitish green, with rather stout curving pedicellate ovaries which are 1.5~-2 cm. long; dorsal sepal ovate to elliptic, concave, 4~5 mm. long, 3-4 mm. wide; lateral sepals obliquely ovate, obtuse, 4-6 mm. long, about 3 mm. wide; petals linear-oblong to narrowly oblong- spatulate, entire or rarely toothed at the truncate or rounded apex, rarely retuse, slightly oblique, 5-7 mm. long, mostly less than 2 mm. wide; lip deeply tripartite, ro—16 mm. long, 13-17 mm. wide across the lateral lobes; lateral lobes deeply cut’(usually to the base) into three divisions, with the divisions subdivided again; mid-lobe slender, clavate to narrowly cuneate [Vou. 1, No. 3 WRIGHTIA 168 Mt, Any “apy \) ey CON Tefiams [HALE |rioro| MoT [corr | “We : ta = el oes Amar | “On . ARCH TAS Pay ewes | A \4o, ICOCN |HOCH |LuaB 05. |DIKM. |AING | "Woy ATL 5 sytem ne ~ v 17 — H [wise [DENTOMCOLLIN-TRUNT | oon. 5 Icass TOARITERAY|LYNN |GARTA| MENT [SONE|HASK. |HROC.|TOUNG a RA \HOD Topsw Lan ae Pann. [TARR [OALLAS|RO U GAINES SAWS TBORD. |SCUR [FISHERLIONES [SHACK )S PH. PALO xauF |= aasON ~ SM ee WN JELLIS = ANDREWS Tin HOW vaatcant feastiD JERaTH\t000 |/° : aa RUSK Jeane fe mitt oe “ aROIN f_ ~~ SUTFOM pm | NIMBLE < “sy =\qbvre ae PRE Sag > TERRELL GiLLESPIE |S avis? ve es an >, mm dS - JALVERDE |EOWAROS |AERR ae “S eat austin) =H + P 7 xs as = A 32 : 8 o> Chg ¢ Mey ar “ye ear 4 Ok mW fa MIMNEY |UVALOE |\MED is = “) is f (an < ey s q 4 as jG, i =) \\ z \ TARR \ sig ‘S Ki = \ g 2 \ ~ a = \, Ss ) , \ Fig. 15. Plant Zones 1. Timber Belt 4. Blackland Prairies ag nd § Pe 2. Coastal Prairies 5. Edwards Plateau 3. Rio Grande Plains 6. Trans-Pecos 7. Plains Country upper portions, but usually covered with a heavy growth of mesquite, scrub oaks, chaparral, cacti, and other small trees and shrubs. The Blackland Prairies (Zone 4)! comprise the blackland and adjacent woodland areas of central Texas, including the Grand Prairie and two broad strips of post oak woodlands running from north to southwest ward on each side of the blacklands, known as East and West Cross Timbers. ‘The term “Blackland Prairies” is used in this paper as an abbreviation for “ Black- land [North Central] Prairies and Adjacent Woodlands.”’ 1940] CorRRELL: TExAs PreRIDOPHYTA 251 Ferns are uncommon here as in most prairie country. However, in the northwestern part of this zone is found some elevated rocky country, es- pecially in Palo Pinto County and vicinity, and there are additional scattered prominent rock outcrops throughout the zone which support quite a few rock-inhabiting species. | The Edwards Plateau (Zone 5) is for the most part an elevated, semi- mountainous, rolling or broken and irregular limestone region with a num- ber of deep spectacular canyons cutting through the Balcones Escarpment on the south. However, in the eastern portion the so-called “granite dis- trict,’ comprising Burnet, Llano and Mason counties, is characterized by extensive mountainous granite outcrops. Although geologically dissimilar from the rest of the region, this district is included for convenience as a part of the Edwards Plateau. The Cedar Brakes and part of the Mesquite Belt occur in this zone, and are characterized by cedars, scrub oaks and mesquite. Because of the many large canyons and rugged terrain, this area is botanically of much interest and has consequently been visited by many botanical collectors. The ferns here are primarily lithophilous, being repre- sented mainly by various species of Cheilanthes, Notholaena, and Pellaea. The Trans-Pecos (Zone 6) includes the only true mountain regions in Texas. The region is of low rainfall. In consequence, the various mountain ranges support on their slopes a xerophytic vegetation. However, on the higher summits, valleys, and slopes where a more mesophytic environment exists are found forests of conifers and hardwoods. The numerous mountain ranges have been only partially explored, if at all. The Guadalupe Range in Culberson County comprises the southernmost extension of the Rocky Mountain mass. The ferns of this area are almost wholly lithophilous. Although we perhaps know more about the ferns of the Chisos Mountains in Brewster County and the Davis Mountains in Jeff Davis County, there is little doubt that these, as well as the entirely unexplored ranges, will produce additional species of ferns, particularly of Mexican affinity, when further field exploration is undertaken. The Plains Country (Zone 7) or Staked Plains, as they are commonly called, covers a vast area of northwestern Texas. Tremendous st retches of flat or only slightly rolling land are to be found here. The region is com- posed of two rather distinct plains, the highest being farthest west. The eastern plain drops several hundred feet rather abruptly and irregularly from the western plain to leave large exposed rock faces and ledges, called “rim rock,” a favorite habitat for a few of the ferns occurring in the Plains Country. Palo Duro Canyon in Randall and Armstrong counties, and the breaks of the Canadian River to the north also provide excellent habitats for the few ferns found in the Plains Country. On the whole, however, this region, like the Coastal Plains country, is essentially a fernless area. Fig. 16 summarizes in a graphic form our knowledge thus far of the distribution of species by counties. Each dot represents a species or variety found within a particular county. It reveals not only the regions of dense and sparse distribution, but also shows the regions which have been botani- 252 WRIGHTIA | [Vou. 1, No. 5 cally explored in so far as ferns are concerned. The results shown on this map are probably indicative of the distribution of other plant groups— based on collecting. As is shown, the Timber Belt, Edwards Plateau, and Trans-Pecos regions contain the greatest number of species, which though i Fig. 16. Distribution of ferns and fern allies. Each dot represents a species or variety within a particular county. The dark lines outline the various plant zones. often infrequent are more or less uniformly distributed in each section. The greatest number of species are found in Brewster County which has 49. The Plains Country and Coastal Plains have very few species of ap- parently sparse distribution. The Trans-Pecos and Edwards Plateau have 30 species in common, and the Edwards Plateau and Timber Belt have 13 species in common. Only 7 species are found in all three regions just mentioned, these being Asplenium resiliens, Cheilanthes alabamensis, C. Eatoni, C. tomentosa, Equisetum prealtum, Pellaea atropurpurea, and P. Wrightiana. Although no one species has been found in all the plant zones in Texas, there are several which are found in 5 or 6 of the 7 zones. Those found in 6 zones are A diantum Capillus-Veneris, Asplenium resiliens, Cheilanthes tomentosa, Equisetum prealtum, Marsilea uncinata, M. vestita, and Pellaea atropur- 1949] CORRELL: TEXAS PTERIDOPHYTA 253 purea. Those found in 5 zones are Cheilanthes alabamensis, C. horridula, Dryopteris normalis, Equisetum laevigatum, Notholaena sinuata var. in- tegerrima, Pellaea Wrightiana and Polypodium polypodioides var. Michauxi- anum. Several species are confined to only one zone. These are discussed under the section dealing with each species. In order to present the information regarding distribution in a con- venient and immediately usable form, all of the species, varieties, and forms are listed in the table below and the plant zones in which each occurs are denoted by cross-marks (X). It is thus possible to find quickly the general distribution of each entity. Following the table is an annotated list giving in more detail the distribution of the ferns and fern allies in each plant zone and a brief description of their habitats. Species, varieties, forms Plant Zones (see Fig. 15) 1h ee es oe, ee ae Adiantum Capillus-Veneris. 2... se eae > Ee 4. Oe ae eit BARIUM GRGHOIIIS i Ge ok poco ns as ees D8 Adiantum tricholepis f. glabrum............-.-. D.¢ NIE TRONINE NA a he iar oes oye ee x emt Vier ge Carico fon cae es oo Xx MOPANOIGN, DIBLYHSUION. 8 ec ee pee 4 ~ Paonia vend a ee Petty EIIONOUIAT. 60 a ok eee D4 MARTTI, GRUNIOMIOINRE: § ok 255s Bese < oe D.G2 ko, p< PEROT CRINGE ee og a ce oe oe aw x Pere MIMO og ee ee ee 2.4 Botrychium dissectum var. tenuifolium.......... a o.4 BOVOUIEM ViTADIBIN. .. .<- =. 2. =<. 6-26 s - x > % i I es i ee ».4 : SCimnlantues dlahamensic. >... oe i ee eee x8 5 a he 4 | xX RrnOUtine WAIN: 2 on ee Se 2.4 >. .s x a Cheilanthes Eatoni f. castanea...........----+-- xX : ps xX Pa TSRG eth e eos ee i ee . x Ree ec tiee PNA Ie ee re ee b. iaarcthen HOrriauls ys ck ee er ee + hep & te me, Ge. @ Ctieianthes Kauttomal. 5.5 eke se ee eee ye 4 CURL NON IBNORR: 66s oe one nr ee 2.4 . Cheilanthes lendigera.........-.------+++ 2 eee > Cheilanthes leucopoda..........-.--+-22:s0200> x - Cheilanthes Lindheimeri........-.--------++--> Be. ee € = : Camanithes tomentosa. oid. oS ns ie ee eae zx ALS : X | 2 Ret asiiling VilGbAs c=. 6 oboe ce ee ee 2 = | Ceimnthes Whotonl.o: . 5. eis oe eee ee = Cheilanthes Wrightii.........------+---+++-°* = | aVecoptens (raging... .6. cen - X | Cystopteris fragilis var. protrusa.....-..-------> | », . | Cystopteris fragilis f. simulans.........--------- : x! PCY OrOrIA GHIALALN. «| oe ie ee x . | De Guterik Mamta a eee ee % 2 | Dryopteris Filix-mas..........----+-:--- aes ee ee 254 WRIGHTIA Species, varieties, forms [Vou. 1, No. 5 EIVFODRTIE NEFARORUPETO. 6 vas Se rea ee es SUPE IE HOVIDAUB es oS ba ek eee Dryopteris thelypteris var. Haleana............. Se POD URTIE VETRIOUNO cs in ca kt Se ees Shae = Pais MAURIE yO a te oe eo Equisetum kansanum f. caespitosum............. Equisetum kansanum f. variegatoides............ PH CROW INOUE eee eet ess oe sie ch ROMO ORIN i sa ee Equisetum prealtum f. Drummondii............. Equisetum prealtum f. texana.................. FaOWbee ENO UNA. gd ee ThGbibe TCILEOOGH. 6-4 ois Sees oh ie oe ra er BINNS oe se ey Gh oe es ECO ROOIUIT BOTIOERUM 626, isi cia nee es EPeCnOUIU MODCCHIOIUED.. 6 Fo 5 i is Lycopodium alopecuroides var. pinnatum Lycopodium carolinianum PN GreOrY TCL. oo a ee ee eek RENTON TARGIONOdN ee ee ee Brahe PR a re oy, DRM OH BONEUI eis een ee ics ee Dette MUU eo eG en ort es pe; PR OMrete CAMA ee, Notholaena Aschenborniana INDE NASON ois a i a eee ee ee PYOIANCUR CPO, oi Si ke ek Notholaena limitanea INOGIUBODA NOMIELR Se ea a Notholaena Schaffneri var. Nealleyi Notholaena sinuata............... Notholaena sinuata var. cochisensis............. Notholaena sinuata var. integerrima............. NOtnOIeene Dteneyi | 6 ee, Onotien HpnmnNis 6. ee eee ee, Ophioglossum crotalophoroides Ophioglossum Engelmanni.....-...........:... Ophioglossum nudicaule var. tenerum Ophioglossum vulgatum Usmunds: cinpatmnoindn 6 oo oo A ke Osmunda regalis var. spectabilis Pennen MIGpUIDIRR 6 ee Pellaea cardiomorpha Pellaea glabella eR oR eee ML idl A Ree ey aN OM ei ee ie ie Oe gg gg ag Plant Zones (see Fig. 15) eae money Pd es es am.< aia Ar aia ~“ nnn nr 24 x |X x x ¥ 24 mM MMe Meee OM A AP ro OM a bd bd bd mr 5 x xX wx ais A A Mig aislecee be | pe pape pa Anes MA AA Me Ar A 7 aia AA anata 4 nn A 1940] CorRELL: TEXAS PTERIDOPHYTA 255 Species, varieties, forms 7 Plant Zones (see Fig. 15) ; ae eo aT z ; eit ane 6 ee Pellaea longimucronata........... ska ittasetencae tee X Pellaea microphylla........ Mae Wee teerats baer. 4 x Pease ovata (0. eee eras ae yee XX : Pern Taringa... oe ee p poneon Wriehtiana. ic. $2 0 ies en bees xX >... : Puinerophiebia auriculata...:.... 0... vases omnes . Phanerophiebia umbonata.. i... 2)... 0a < 9s - ae o> 4 me. PUPUIateR MIDerinniN iy eo Se ce ee D, . eo podium erythrolepis... co. .6065 sk eee = ae , Polypodium polypodioides var. Michauxianum. .. . ah EE EG Polypodium thyssanolepis..... 6.2.25 6-25 34's . 2 Polystichum scrostichoides.......:..--..-++++-- x oe Polystichum acrostichoides f. incisum........--- + xX Pe : Pteridium aquilinum var. pseudocaudatum....... ie. ee x = Pteridium aquilinum var. pubescens........---+- 2 ree Eifel 8 es it ee Vo ge toes xX x Lae Mee OHY Hite ge ey ee eee ee Ne bee. @ ae Beets Arisoniia. oe Se ees X | = Mees Corvin ate Selaginella lepidophylla.............--+++++ 50:5 | Pi DUNNER WN tee a sc es i ee eas | = | Selaginella mutica var. limitanea........-..----- eS Selaginella mutica var. texana.........-------°> | . MEIARINGlA NOOMEXIGANA.. 2.2. 3. ese et | S| Rrenelia Hu GP ee i ee : e& oe Selaginella Riddellii..,.........-----+2 2000-0 ee Peele TaMpDOl. <5. kk rk et | | sf ca be meanerbens Sheldon. 200 es cee te eee | I | Melarinella Underwoodii.........-- +--+ ----- <2 | a Pemenelia Wrightii. 0.55.6. eet ee | | Hoes ortarrs heracieuolin. --.<. ce s it | | = Woodsia mexicana’... ..... 6.6. hee’, Tae rae | "i | ee | WOUGME Gia i ae ea ta ee te *) ee Woodsia Plumerne....-..-:4.0-0r treet yy a te Woodwardia virginica... 002.05 +. eer be ee 2 eo POLYPODIACEAE. Fern Family Aprantum [Tourn.] Linnaeus 7 Adiantum modestum ApIANTUM CapILLus-VENERIS L., Sp. Pl. 1096. 1753- Adi Underw. Wherever a calcareous substratum, and continuously available moisture occur, bob normalis are almost invariably found. Since this Se ta ie most frequently in the canyons of nal aioe r is ine bluffs exceedingly common there. It often covers the wa iB ae ame ae kept wet by seepage springs, and is common On Frocks ¢ streams. especially limestone, and sufficient this species and Dryopteris type of habitat occurs 256 WRIGHTIA [Vou. 1, No. 5 Coastal Prairies: Harris County. This lone station occurs along a deep shaded spring branch near Houston where there is an outcrop of calcareous sandstone. Rio Grande Plains: In four of the extreme northern border counties. Blackland Prairies: Infrequent and of scattered distribution. Edwards Plateau: Common, mainly in the southeastern part. Trans-Pecos: On rocks along streams in Brewster, Culberson and Presidio Counties. Plains Country: Infrequent in spring-fed canyons in Armstrong and Ran- dall Counties; also in several central and southern counties. ADIANTUM TRICHOLEPIS Fée, 8™° Mém. Fam. Foug. 72. 1854-57. On moist or dry limestone cliffs, in creeks among rocks and among boulders on open slopes, usually in partial shade. Edwards Plateau: Medina County. Reported by Reverchon (1903) as hav- ing been collected by Bigelow in Val Verde County. Clute (Fern Bull. 12:44. 1904) also reported this species, as forma glabrum Clute, from the mouth of the Pecos River, where it was collected by J. H. Ferriss. I have not seen this collection. ASPLENIUM Linnaeus ASPLENIUM PatMeErI Maxon, Contr. U. 8. Nat. Herb. 13: 39. 1909. On moist shaded mountain slopes and in crevices of granite cliffs. Trans-Pecos: Jeff Davis County. Rare in the Davis Mountains. ASPLENIUM PLATYNEURON (L.) Oakes ex D. C. Eaton, Ferns N. Am. 1: 24. 1878. Asplenium ebeneum Ait. Although Palmer (1919) states that this species is widely distributed in both eastern and western Texas, I find that it occurs only in the Timber Belt, Coastal Prairies and Blackland Prairies regions. Most frequent in sandy loam on rich wooded banks and slopes along streams and on the edge of flooded woodlands; also on hummocks in swamps, rotting logs, talus slopes, boulders, ledges, and in thickets; rarely semiepiphytic. Timber Belt: Generally distributed and frequent. Coastal Prairies: Brazoria and Orange counties. Blackland Prairies: Generally distributed but of local occurrence. ASPLENIUM RESILIENS Kunze, Linnaea 18: 331. 1844. Asplenium parvulum Mart. & Gal. In crevices of cliffs, boulders or ledges of limestone, granite or in calcareous soils, usually in shaded but dry situation. Timber Belt: Angelina County. Coastal Prairies: Orange County. This is of rather unusual occurrence. The species was doubtless found on shell mounds. Rio Grande Plains: In several extreme northern counties. Blackland Prairies: In several western counties. Edwards Plateau: In the southern and eastern sections. Trans-Pecos: Rather generally distributed but not common. 1940] CoRRELL: TEXAS PTERIDOPHYTA 257 ASPLENIUM TRICHOMANES L., Sp. Pl. ro80o. 1753. In moist sheltered crevices of cliffs, ledges and cavernous places in rocks. Trans-Pecos: Jeff Davis County. Rare in the Davis Mountains. AtHYRiIUM Roth ATHYRIUM ASPLENIOIDES (Michx.) Eat., Man. 122. 1817. Athyrium filix- femina of authors, not (L.) Roth; Asplenium asplenioides (Michx.) D. C. Eaton ex Chapman. In sandy bogs or moist sandy woods, swamps, wet thickets and on stream banks. Timber Belt: Generally distributed and rather common. Coastal Prairies: In several northern border counties. Blackland Prairies: In several eastern border counties. Bommertia Fourn. ex Baillon BomMeria nisprpa (Mett.) Underw., Bull. Torr. Bot. Club 29: 633. 1902. Gymnopteris hispida (Mett.) Underw. On rocky slopes, in crevices of bluffs and on ledges, usually in moist leaf mold under low shrubs. Trans-Pecos: In the southern and western counties. CHEILANTHES Swartz CHEILANTHES AEMULA Maxon, Contr. U. S. Nat. Herb. 10: 495. 1908. Cheilanthes Moritziana of authors, not Kuntze. On shaded banks, on ledges in canyons, in cavities under overhanging rocks, and in rock crevices. Edwards Plateau: Comal and Val Verde counties. CHEILANTHES ALABAMENSIS (Buckl.) Kunze, Linnaea 20: 4. 1847. On limestone hillsides, in rock crevices and under protecting ledges, often associated with Pellaea atropurpurea. Timber Belt: Walker County. : Blackland Prairies: In several western and southern counties. Edwards Plateau: Common and generally distributed. Trans-Pecos: In the central and southern parts. Plains Country: Concho and Upton Counties in the extreme south. CHEILANTHES Eatoni Baker in Hook. & Baker, Syn. Fil. 140. 1867. Cheilanthes tomentosa var. Eatoni (Baker) Davenp. _ On ledges and talus slopes, bluffs and about boulders in thinly wooded areas, Timber Belt: Morris County. The occurrence of this species so far from the Edwards Plateau is unique. My wife and I found it growing on ferruginous rocks and ledges in Daingerfield State Park. Blackland Prairies: Wilson County in the extreme southwest. _ Edwards Plateau: Gillespie, Llano, Mason, and Val Verde counties. Trans-Pecos: Common and generally distributed. 258 WRIGHTIA [Vou. 1, No. 5 CHEILANTHES Eaton Baker forma castanea (Maxon) Correll, comb. nov. Cheilanthes castanea Maxon, Proc. Biol. Soc. Wash. 32: 111. 1919. When extremes of typical C. Hatoni and forma castanea are found they are so distinctive that one would immediately consider them to be specifi- cally different. Unfortunately, very little material of extreme f. castanea exists in herbaria. An overwhelming amount of material does exist, how- ever, which grades into one or the other of these two forms. It is simply an arbitrary matter as to which category they should be relegated. I have placed all plants which have all or part of their fronds tending to be gla- brescent on the upper surface of their segments into f. castanea. I have designated those plants with segments hoary and densely tomentose above as the typical form. Habitat same as that of the typical form. Timber Belt: Morris County. It occurs with the typical C. Hatoni on rocks in the Daingerfield State Park. This is an exceedingly isolated locality. Trans-Pecos: Central part, especially in Brewster County where some plants collected in the Chisos Mountains are strikingly different from typical C. Eatoni. Plains Country: Mitchell County. CHEILANTHES Fret Moore, Ind. Fil. 38. 1857. Cheilanthes lanuginosa Nutt. ex Hooker; C. gracilis (Fée) Mett. In dry crevices of boulders, cliffs and ledges of limestone or calcareous sandstone, usually exposed to intense sunlight. Blackland Prairies: Palo Pinto County. Edwards Plateau: In the central and southern parts. Trans-Pecos: Rather frequent and generally distributed. Plains Country: Generally distributed. This is perhaps the most wide- spread and abundant fern in this part of Texas. CHEILANTHES FENDLERI Hook., Sp. Fil. 2: 103, pl. 107 B. 1858. In rock crevices and at the base of boulders, in shaded canyons and on dry wooded banks. Trans-Pecos: Hudspeth and Jeff Davis counties. Rare and of local oc- currence. CHEILANTHES HORRIDULA Maxon, Am. Fern Journ. 8: 94. 1918. Pellaea aspera Baker in Hooker & Baker; P. aspera forma compacta Clute; Cheilanthes horridula forma compacta (Clute) Broun; C. aspera Hook., not Kaulf. At the time of his death, Dr. Maxon had on loan all of my collections of this species from the Edwards Plateau. He had noted that this material differed from typical C. horridula of the Trans-Pecos in having hairs on the upper surface of the pinnules which were longer, more slender and less pustulate, and he was of the opinion that it might be specifically or vari- etally different. I have since studied this and other material from the Ed- wards Plateau with Dr. Morton and we have concluded that although it is somewhat atypical it is not sufficiently distinct to be segregated. 1949] CorRELL: TEXAS PTERIDOPHYTA 259 In sheltered or exposed crevices of limestone or sandstone, where the plants are usually so firmly imbedded that they are difficult to collect intact. Rio Grande Plains: Maverick and Webb counties, along the Rio Grande; also in the extreme northern border counties. Blackland Prairies: Palo Pinto County. Edwards Plateau: Rather frequent and generally distributed. Trans-Pecos: In the southeastern half. Plains Country: Concho and Upton counties in the extreme southern part. CHEILANTHES Kautrussi Kunze, Linnaea 13: 145. 1830. In crevices of rock ledges and among boulders on hillsides. Edwards Plateau: Llano County. This is the first record of this species east of the Trans-Pecos country. Trans-Pecos: Brewster and Jeff Davis counties. CHEILANTHES LANOSA (Michx.) D. C. Eaton in Torr., Rept. U. 8. & Mex. Bound. Surv. 2: 234. 1850. On dry recky slopes and ledges of sandstone. Blackland Prairies: McLennan County. Only one locality known for Texas. CHEILANTHES LENDIGERA (Cay.) Sw., Syn. Fil. 128. 1806. In crevices on cliffs and boulders. Trans-Pecos: Brewster County. Rare in the Chisos Mountains. CHEILANTHES LEUcOpPoDA Link, Fil. Sp. Hort. Berol. 66. 1841. On exposed limestone slopes and ledges on hills, mountainsides and in canyons. Edwards Plateau: Edwards, Real and Uvalde counties. CHEILANTHES LINDHEIMERI (J. Sm.) Hook., Sp. Fil. 2: ror, pl. 107 A, 1858. On dry shaded boulders and rock ledges, among talus rubble, in canyons, and on bluffs and cliffs. Rio Grande Plains: MeMullen County. Blackland Prairies: Palo Pinto County. Edwards Plateau: In the southeastern part. Trans-Pecos: In the southern and western parts. CHEILANTHES TOMENTOSA Link, Hort. Berol. 2: 42. 1833. In clefts of ledges or boulders, on wooded hillsides, usually on sand- stone, granite or siliceous rocks. Timber Belt: Tyler and Walker counties. Rio Grande Plains: Atascosa County, in the extreme northeastern part. Blackland Prairies: Rather generally distributed but of local occurrence. Edwards Plateau: Southeastern part. Trans-Pecos: Southern half. Plains Country: Mitchell County, in the extreme south. CHEILANTHES VILLOSA Davenp., Cat. Davenp. Herb. Suppl. 45. 1883. In dry crevices of limestone ledges, cliffs and boulders, or among granite boulders. 260 WRIGHTIA [Vox. 1, No. 5 Edwards Plateau: Val Verde County. Trans-Pecos: Generally distributed but uncommon. CHEILANTHES WooTont Maxon, Proc. Biol. Soc. Wash. 31: 146. 1918. In dry rocky places, often at the base of boulders. Trans-Pecos: El Paso, Jeff Davis and Presidio counties. Rare and of local occurrence. CuHEILANTHES Wricutit Hook., Sp. Fil. 2: 87, pl. 100 A. 1858. On exposed talus slopes, in crevices of boulders and in rocky ravines. Trans-Peecos: In the southern and western sections. Cysroprerts Bernhardi CYSTOPTERIS FRAGILIS (L.) Bernh., Neu. Journ. Bot. Schrad. 17: 26, pl. 2, f. 9. 1806. Filix fragilis (L.) Gilib. In sheltered crevices in cliffs, on moist banks and wooded talus slopes. Trans-Pecos: Culberson and Jeff Davis counties. It is also represented in Culberson County by forma simulans Weatherby. CYSTOPTERIS FRAGILIS var. PROTRUSA Weatherby, Rhod. 37: 373. 1035. On banks in oak forest. Blackland Prairies: Gonzales County. Dryopteris Adanson DRYOPTERIS CRISTATA (L.) Gray, Man., ed. 1, 631. 1848. In marshes, bogs, swamps, thickets and meadows, and on springy wooded slopes. Timber Belt: Bowie County. A sterile frond of this species is mounted on a Gray Herbarium sheet with a sterile frond of Osmunda cinnamomea. They were collected on the edge of a bog near Texarkana by E. J. Palmer. DRYOPTERIS DENTATA (Forsk.) C. Chr., Danske Vid. Selsk. Skr. VIII, 6: 24. 1920. Dryopteris parasitica of authors, not (L.) Kunze. On rocky wooded slopes, on hummocks in swamps, and along wooded streams. Coastal Prairies: Harris County. Dryopreris Finix-mas (L.) Schott, Gen. Fil. Pl. 67. 1834. In cool rocky mountain woods, on shaded talus slopes, and ravine walls and ledges. Trans-Pecos: Jeff Davis County. Rare in the Davis Mountains. DRYOPTERIS HEXAGONOPTERA (Michx.) C. Chr., Ind. Fil. 270. 1905. Phegopteris hexagonoptera (Michx.) Fée. On sandy wooded slopes and in ravines along streams, and on the margin of bogs. Timber Belt: San Augustine County. Only one station known for this species in Texas. 1949| CorRELL: TEXAS PTERIDOPHYTA 261 Dryopreris NorMALIS C, Chr., Ark. for Bot. 9!': 31. 1910. Dryopteris patens of authors, not (Sw.) Kuntze; D. normalis var. Lindheimeri (A. Br.) C. Chr.; D. augescens (Link) C. Chr. var. Lindheimeri (A. Br.) Broun; Nephrodium patens of authors, not (Sw.) Desv. On the edge of sandy creeks, in deep canyons, on wet bluffs and ledges, in swamps and woods, and at the base of dripping limestone bluffs. This species is practically always associated with Adiantum Capillus-Veneris. Timber Belt: In the eastern and southern sections, especially in the south. Coastal Prairies: In the central and eastern parts. Rio Grande Plains: In the extreme northern counties. Blackland Prairies: Mostly in the southern part. Edwards Plateau: In the southern and eastern sections. DrYOPTERIS THELYPTERIS (L.) Gray var. Haleana (Fern.) Broun ex Weatherby, Am. Fern Journ. 26: 95. 1936. Nephrodium thelypteris of authors, not Linnaeus. In open sandy bogs, swamps and meadows, or in open low woodlands, Timber Belt: Harrison and Marion counties. Reported from Houston and Liberty counties by Palmer (1919). Coastal Prairies: Jefferson County. Blackland Prairies: Waller County. < Dryopreris verstcotor (R. P. St. John) Broun, Index N. Am. Ferns 82. 1938; Brown & Correll, Ferns and Fern Allies La. 59. 1942. In our work on Louisiana ferns, Brown and I regarded those plants whose characters appeared to have been derived from both Dryopteris dentata and D. normalis as putative hybrids of those species. Plants which fall in this category have been collected in eastern Texas. They occur in similar habitats to those of their putative parent species. Timber Belt: Jasper and San Jacinto counties. Coastal Prairies: Orange County. LoRINSERIA Presl LoRINSERIA AREOLATA (L.) Presl, Epim. Bot. 72. 1851. Woodwar dia areolata (L.) Moore; W. angustifolia J. Sm. In sandy bogs or low sandy woods, swamps, marshes, on seepage slopes or along streams. Timber Belt: Generally distributed and rather common. Coastal Prairies: In the eastern part. : Blackland Prairies: In several eastern and southern counties. Nepurouepris Schott NEPHROLEPIS EXALTATA (L.) Schott, Gen. Fil., pl. 3. 1834- This species has escaped from cultivation and has apparently become established in several localities in Texas. It is hardy out-of-doors in Orange, where lawns are sometimes covered by its long, lanceolate fronds. ae In open or dense woods, on limestone ledges and on hummocks in swamps; commonly epiphytic. 262 WRIGHTIA (Vou. 1, No. 5 Timber Belt: Nacogdoches County. Edwards Plateau: Real County. Noruouaena A. Braun NoTHoLAENA ASCHENBORNIANA Klotzsch, Linnaea 20: 417. 1847. On rocky slopes and in deep limestone canyons. Edwards Plateau: Val Verde County. Trans-Pecos: Brewster and Culberson counties. NoTHOLAENA AUREA (Poir.) Desv., Mém. Soc. Linn. Paris 6: 219. 1827. Notholaena bonariensis (Willd.) C. Chr.; N. ferruginea Desv. On talus slopes and in crevices of ledges and cliffs, on canyon walls and about the base of boulders. Trans-Pecos: In the western and southern parts. NorHOLAENA CANDIDA (Mart. & Gal.) Hook., Sp. Fil. 5: 110. 1864. On limestone ledges and in crevices of canyon walls and bluffs, on open rocky slopes, at the base of cap-rock and about boulders. Edwards Plateau: In the central and southern parts. Trans-Pecos: Brewster County. NoTHOLAENA DEALBATA (Pursh) Kuntze, Am. Journ. Sci. II, 6: 82. 1848. Pellaea dealbata (Pursh) Prantl; Notholaena nivea var. dealbata (Pursh) Davenp. This species is a borderline case in that it could be just as well placed in the genus Pellaea as in Notholaena. In dry crevices on cliffs and boulders of limestone and other calcareous rocks, usually in partial shade. Blackland Prairies: Ellis County and several other western border counties. Edwards Plateau: In the eastern part. Plains Country: Reported by Palmer (1919) from Cole County. NotrHoLaeNna Grayt Davenp., Bull. Torr. Bot. Club 7: 50. 1880. On talus slopes, in rock crevices and rocky barrens of granite or lime- stone. Edwards Plateau: Burnet and Uvalde counties. Trans-Pecos: Brewster and Presidio counties. NoTHOLAENA Greco (Mett.) Maxon, Contr. U. S. Nat. Herb. 17: 606. 1916. Notholaena Pringlei Davenp. In exposed crevices of rocks. Trans-Pecos: Brewster County. The lone station is in Boquillas Canyon. NOTHOLAENA LIMITANEA Maxon, Am. Fern Journ. 9: 70. 1919. On limestone ledges and in talus rubble at the base of limestone cliffs. Trans-Pecos: Brewster County. Rare in the Chisos Mountains. NOTHOLAENA NEGLECTA Maxon, Contr. U. S. Nat. Herb. 17: 602. 1916. In dry crevices of limestone ledges and cliffs, and on rocky slopes. 1949] CoRRELL: TEXAS PTERIDOPHYTA 263 Edwards Plateau: Val Verde County. Trans-Pecos: Brewster County. NorHoLaENA ScHAFFNERI (Fourn.) Underw. var. NEALLEYI (Seaton) Weatherby in Johnston, Journ. Arn. Arb. 24: 315. 1943. Notholaena Schaffneri of authors, not (Fourn.) Underwood. On exposed or partially shaded rock ledges. Edwards Plateau: Val Verde County, vicinity of the Devils River. Trans-Pecos: Reported by Bush (1903) from Presidio County. NoOTHOLAENA SINUATA (Lag.) Kaulf., Enum. Fil. 135. 1824. On dry gravelly slopes, about boulders, on rock ledges, and in crevices of low cliffs. Rio Grande Plains: Atascosa and McMullen counties, and in several north- ern border counties. Edwards Plateau: Generally distributed and fairly common. Trans-Pecos: Generally distributed and rather frequent. Plains Country: In several extreme southern counties. NOTHOLAENA SINUATA var. COCHISENSIS (Goodding) Weatherby in Johns- ton, Journ. Arn. Arb. 24: 314. 1943. In rubble of open dry mountain slopes, exposed talus slopes, in dry erevices of bluffs and cap-rock. Edwards Plateau: Sterling and Uvalde counties. Trans-Pecos: Generally distributed and frequent. Plains Country: Northward to Motley County; sparsely distributed. NOTHOLAENA SINUATA Var. INTEGERRIMA Hook., Sp. Fil. 5: 108. 1864. In rock crevices of ledges, boulders and bluffs, and on talus slopes. Rio Grande Plains: Maverick and Webb counties, along the Rio Grande. Blackland Prairies: Palo Pinto County and in the extreme southwest corner. Edwards Plateau: Rather generally distributed and frequent. Trans-Pecos: Generally distributed and frequent. aS Plains Country: Northward to Motley County; sparsely distributed. NoTHOLAENA STANDLEYI Maxon, Am. Fern Journ. 5: 1. 1915. Notholaena Hookeri Eaton. ; On dry rock ledges, talus slopes and among and on boulders, on cliffs and canyon walls. Edwards Plateau: Llano and Val Verde counties. Trans-Pecos: Generally distributed and rather frequent. OnocLeEA Linnaeus ONOCLEA SENSIBILIS L., Sp. Pl. 1082. 1753. : i : In swampy open woods, sandy bogs, moist depressions, along streams and on the edge of lakes. Timber Belt: Generally distributed and rather common. Coastal Prairies: Harris and Jefferson counties. 264 WRIGHTIA [VoL. 1, No. 5 Blackland Prairies: Wilson County. Its occurrence here is rather unusual because of its isolation from the Timber Belt. Peviaka Link PELLAEA ATROPURPUREA (L.) Link, Fil. Sp. Hort. Berol. 59. 1841. This is perhaps the most widely distributed fern in Texas. It is found to a greater or lesser degree in all floral zones except the Coastal Prairies, where rock outcrops are exceedingly rare. On rock ledges, among rocks, on cliffs, banks and talus slopes of lime- rock, and in open rocky woods, usually in partial shade. Timber Belt: San Augustine and Shelby counties. Rio Grande Plains: In several northern border counties. Blackland Prairies: In several northern and central counties. Edwards Plateau: Generally distributed and common. Trans-Pecos: Rather frequent and of general distribution. Plains Country: Of sparse though general distribution. PELLAEA CARDIOMORPHA Weatherby in Johnston, Journ. Arn. Arb. 24: 309. 1943. Pellaea cordata (Cav.) J. Smith, not Fée. In shallow cavernous places in cliffs, on wet bluffs and in crevices and on ledges of camyon walls. Trans-Pecos: Brewster, Jeff Davis and Presidio counties. Rare and of local occurrence. PELLAEA GLABELLA Mett. ex Kuhn, Linnaea 36: 87. 1860. This is apparently the first report of this species from Texas. In crevices of cliffs and on ledges of limestone or caleareous sandstone. Plains Country: Potter County. My wife and I found a large colony of this species on a limestone outerop along John Rey Creek. PrELLAEA INTERMEDIA Mett. ex Kuhn forma pusescens (Mett.) Broun, Index N. Am. Ferns 132. 1938. Pellaea intermedia of authors, not Mett. ex Kuhn; P. intermedia var. pubescens Mett. ex Kuhn. On dry rocky slopes and ledges of limestone, cliffs and about boulders. Edwards Plateau: Val Verde County. Trans-Pecos: Rather frequent and generally distributed. Plains Country: Upton County, in the extreme south. PELLAEA LONGIMUCRONATA Hook., Sp. Fil. 2: 143, pl. 115 A. 1858. Among dry rocks, in shelter of boulders and in crevices of cliffs. Trans-Pecos: El Paso and Hudspeth counties. Rare and of local occurrence. PELLAEA MICROPHYLLA Mett. ex Kuhn, Linnaea 36: 86. 1869. Pellaea pulchella of authors, not Fée. On limestone ledges in gorges and canyons, and on high limestone hills, face of cap-rock, in shaded. or exposed situations. Reputedly poison- ous to sheep. Blackland Prairies: Brazos County. This station represents an unusual ex- tension of range from the west. 1940] CoRRELL: TEXAS PTERIDOPHYTA 265 Edwards Plateau: In the northern and western parts. Trans-Pecos: Frequent and generally distributed. PELLAEA OVATA (Desv.) Weatherby, Contr. Gray Herb. 114: 34. 1936. -Pellaea flexuosa (Kaulf.) Link. On dry ledges and talus slopes of limestone or caleareous rocks, on or at the base of cliffs, in rich soil in open woodlands. Blackland Prairies: Palo Pinto County and in several extreme southwestern border counties. Edwards Plateau: In the southern and eastern counties. Trans-Pecos: Brewster and Terrell counties. PELLAEA TERNIFOLIA (Cav.) Link, Fil. Sp. Hort. Berol. 59. 1841. On shaded talus slopes and ledges, in the protection of boulders and shrubs. Trans-Pecos: Brewster County. Although reported from other localities in Texas, this species is apparently confined to the Chisos Mountains, where it is rare. Reports from outside the Chisos Mountains are based on specimens of P. Wrightiana, which this species closely resembles. PeLLAEA WRIGHTIANA Hook., Sp. Fil. 2: 142. 1858. In crevices of ledges, shelter of boulders and on rocky ridges, usually im exposed situations. Timber Belt: Tyler County. This isolated locality is an unusual extension of range from the west. My wife and I found a number of plants on exposed sandstone boulders in the Angelina National Forest. Blackland Prairies: Palo Pinto County, in the northwestern part. Edwards Plateau: In the southern and eastern counties. Trans-Pecos: Brewster, El Paso, Jeff Davis and Presidio counties. PHANEROPHLEBIA Presl PHANEROPHLEBIA AURICULATA Underw., Bull. Torr. Bot. Club 26: 212, pl. 360. 1899. Aspidium juglandifolium of authors, not Kunze ex Klotzsch. On cool damp cliffs and in cavernous recesses in granite rocks, and under overhanging ledges in canyons. Trans-Pecos: Brewster, El Paso and Jeff Davis counties. PHANEROPHLEBIA UMBONATA Underw., Bull. Torr. Bot. Club 26: 211. 1899. In cool shaded canyons in maple forests, in moist ravines and on rock ledges. Trans-Pecos: Brewster and Jeff Davis counties. PortyropiuM [Tourn.] Linnaeus Po.ypoptum ERYTHROLEPIS Weatherby, Contr. Gray Herb. 65: 11. 1922. On cliffs of porphory rocks. Trans-Pecos: Jeff Davis County. Rare in the Davis Mountains. Potypopium potypoprorpeEs (L.) Watt var. MicHauxtanum Weatherby, 266 WRIGHTIA [Vou. 1, No. 5 Contrib. Gray Herb. 124: 31. 1939. Polypodium polypodioides of au- thors, not (L.) Watt; P. incanum of authors, not Swartz. This, the only truly epiphytic fern found in Texas, occurs on various species of trees, especially oaks, occasionally on rock ledges, boulders, and mossy banks, in moderately shady and damp locations. Timber Belt: Generally distributed and common. Coastal Prairies: In the eastern half. Rio Grande Plains: Atascosa County. Blackland Prairies: Mainly in the south-central part where it is sparsely distributed. Edwards Plateau: Uvalde County. POLYPODIUM THYSSANOLEPIS A. Br. ex Klotzsch, Linnaea 20: 392. 1847. On open rocky slopes and in crevices of boulders. Trans-Peeos: Brewster and Jeff Davis counties. Rare. PorysticHuM Roth POLYSTICHUM ACROSTICHOIDES (Michx.) Schott, Gen. Fil., pl. 9. 1834. On rich wooded hillsides or steep shaded banks of ravines along streams, along sandy creeks and in swamps. Timber Belt: Generally distributed and rather common, especially in the northern part; forma incisum (Gray) Gilbert occurs in several counties. Coastal Prairies: In the eastern half. Blackland Prairies: Waller County and in several eastern border counties. PTERIDIUM Scopoli PreRIDIUM AQUILINUM (L.) Kuhn var. pseupocaupaTuM Clute, Fern Bull. 8:39. 1900. Ptertdium latiusculum var. pseudocaudatum (Clute) Maxon; P. aquilinum of authors, not (L.) Kuhn; P. caudatum of authors, not (L.) Maxon; Pteris aquilina var. pseudocaudata Clute. In open sandy woods, sandy loam of rocky seepage slopes, and flat pinelands. Timber Belt: Common and generally distributed. Coastal Prairies: In the northeastern border counties. Blackland Prairies: Mainly in the south-central counties. PTERIDIUM AQUILINUM Var. PUBESCENS Underw., Our Nat. Ferns, ed. 6, gt. 1900. Pteridium aquilinum var. lanuginosum (Bong.) Fernald. On rocky open wooded mountain slopes, and rich banks along streams. Trans-Pecos: Jeff Davis County. Rare on Mt. Livermore, Davis Mountains. Preris Linnaeus PTERIS MULTIFIDA Poir. ex Lam., Encye. Bot. 5: 714. 1804. : This species has escaped from cultivation and become established 10 southeastern Texas. In sandy soil in woods and on masonry. 1949] CorRELL: TEXAS PTERIDOPHYTA 267 Timber Belt: Hardin County. Coastal Prairies: Jefferson County. TrecTaARIA Cavanilles TECTARIA HERACLEIFOLIA (Willd.) Underw., Bull. Torr. Bot. Club 33: 200. 1906. Tectaria trifoliata (L.) Cav.; Aspidium trifoliatum D. C. Eaton, not Swartz. In limestone sink-holes and caverns in limestone bluffs. This species occurs in such sinks as the Devil’s Sink-hole in Edwards County, where partial shade and protection from extremes of heat and cold exist. Edwards Plateau: Edwards and Uvalde counties. Reported by Bush (1903) from Comal County. Woopsta R. Brown Woops!IA MEXICANA Fée, Mém. Fam. Foug. 66. 1854. On shaded or sometimes sunny rock ledges and in moist crevices of cliffs. : Trans-Pecos: In the southern and western parts. Woobpsta optusa (Spreng.) Torr., Cat. Pl. in Geol. Rept. N. Y. 195. 1840. Among rocks or along rather dry shaded ledges, on rock outcrops and cliffs, commonly in sandstone or granite regions; also on well-drained wooded banks, talus slopes or sandy knolls in woods. Timber Belt: Mostly in the northern and western parts. Blackland Prairies: Generally distributed and rather frequent. Edwards Plateau: In the eastern part. Trans-Pecos: Reported by Palmer (1930) from Mt. Livermore, Davis Mountains, Jeff Davis County. If this report is correct it represents an iso- lated locality for the species. Woopsta PLuMERAE Lemmon, Bot. Gaz. 7: 6. 1882. On moist shaded cliffs and ledges, talus slopes and at the base of boulders. Trans-Pecos: In the southern and western counties. Plains Country: Randall County. Rare and isolated from the Trans-Pecos in Palo Duro Canyon. Woopwarpia J. Smith Woopwarpia VirGINICA (L.) J. Sm., Mém. Acad. Turin 5: 412. 1793. Anchistea virginica (L.) Presl. In sphagnous bogs, swamps, moist thickets, and along streams. Timber Belt: Rather generally distributed. Coastal Prairies: In the extreme northeastern border counties. Blackland Prairies: Gonzales, Lee and Milam counties. SCHIZAEACEAE. Curly-grass Family ANEMIA Swartz ANEMIA MEXICANA Klotzsch, Linnaea 18: 526. 1844. On dry partially shaded steep hillsides, banks of dry ravines, on face 268 WRIGHTIA VOL: FUNG = or at base of cliffs and boulders, under overhanging limestone ledges in dry soil. Edwards Plateau: In the southern and southeastern parts where it is rather common. LyGoprumM Swartz LyGopruM JAPoNicuM (Thunb.) Sw., Journ. Bot. Schrad. 18007: 106. 1801. In moist sandy woods, thickets and along ditches, where it has escaped from cultivation and become naturalized. Timber Belt: Jasper County. Coastal Prairies: Jefferson and Orange counties. OSMUNDACEAE. Cinnamon-fern Family Osmunpba [Tourn.| Linnaeus OSMUNDA CINNAMOMEA L., Sp. Pl. 1066. 1753. In swamps, bogs, wet depressions in woods, on stream-banks, springy slopes and wet rock ledges. Timber Belt: Generally distributed and common. Coastal Prairies: In the extreme northeastern counties. Blackland Prairies: In several of the eastern and southern counties. OSMUNDA REGALIS L. var. SPECTABILIS (Willd.) Gray, Man., ed. 2. 600. 1856. Osmunda regalis of authors, not Linnaeus. In bogs, moist woods, and on springy slopes, hummocks in swamps, stream banks and rocky seepage slopes. Timber Belt: Generally distributed and common. Coastal Prairies: In the easternmost counties. Blackland Prairies: In several central and eastern counties. OPHIOGLOSSACEAE. Adder’s-tongue Family BotrycHiuM Swartz BorrycHIuM DISsEcTUM Spreng. var. TENUIFOLIUM (Underw.) Farwell, Papers Mich. Acad. Sci. 3: 89. 1924. Botrychium obliquum of authors, not Muhl. ex Willdenow; B. obliquum Muhl. var. tenuifolium (Underw.) Gilbert. In low wet woods, swamps, moist sandy pine woods, frequently on or about rotting wood. Timber Belt: In several eastern and southern counties. Coastal Prairies: Jefferson County. BorrYcHIUM VIRGINIANUM (L.) Sw., Journ. Bot. Schrad. 18007: 111. 1801- In alluvial or well drained woodlands, swamps and thickets. Timber Belt: Uncommon but generally distributed. Blackland Prairies: Dallas and Red River counties. Edwards Plateau: Kerr County. This unusually remote locality from the Timber Belt is probably part of a floral relict area. 1949] CorrRELL: Texas PreRIDOPHYTA 260 OpHIoGLossuM [Tourn.| Linnaeus OPHIOGLOSSOIDES CROTALOPHOROIDES Walt., Flora Caroliniana 256. 1788. Ophioglossum pusillum Nutt. In damp pastures, moist sandy soil of open pine forest, and on grassy slopes. Timber Belt: Reverchon (1903) reported it from Newton County. Blackland Prairies: Bastrop and Harris counties. Very rare or, perhaps more correctly, overlooked because of its small size. OPHIOGLOSsuM ENGELMANNI Prantl, Ber. Deut. Bot. Ges. 1: 351. 1883. Usually found in large colonies in thin black soil on limestone barrens or ledges, rocky woodland slopes, in cedar brakes, or in clayey soil along streams. Timber Belt: Central and western parts. Blackland Prairies: Rather frequent and generally distributed. OPHIOGLOSSUM NUDICAULE L. f. var. TENERUM (Mett.) Clausen, Mem. Torr. Bot. Club 197: 146. 1938. Ophioglossum tenerum Mett. ex Prantl. On grassy slopes and in wet meadows, damp depressions in pinelands and on the edge of bogs. Timber Belt: Hardin County. Rare. OPHIOGLOSsUM VULGATUM L., Sp. PIL. 1062. 1753. In moist open woods, meadows, alluvial woodlands and swamps. Timber Belt: Harrison County. Reported by Bush (1903) from Upshur County. Coastal Prairies: Jefferson County. SALVINIACEAE. Salvinia Family AzouLa Lamarck AZOLLA CAROLINIANA Willd., Sp. Pl. 5: 541. 1810. On still water of swamps, ponds, lakes and in slow-moving water of streams. This species, as well as those of Marsilea, are probably dissemi- nated in part by water-fowl. Coastal Prairies: Orange, Matagorda, and several other northern border counties. Rio Grande Plains: Cameron County. : Blackland Prairies: Dallas County and several southern counties. Trans-Pecos: Jeff Davis and Presidio counties. MARSILEACEAE. Pepperwort Family MarsILea Linnaeus MarsILEa Macropopa Engelm. ex A. Br., Am. Journ. Sci. II, 3: 56. 1847. This species is apparently confined to southern Texas, where it occurs in mud or sandy soil and water of swamps, marshes, woodland bogs, ditches, streams, and on the edge of ponds and lakes. 270 WRIGHTIA [Vou. 1, No. 5 Coastal Prairies: In several counties of the extreme western part. Rio Grande Plains: Rather common and widespread. Blackland Prairies: In the extreme southwestern part. Edwards Plateau: In several counties in the extreme south. MARSILEA TENUIFOLIA Engelm. ex Kuntze, Am. Journ. Sci. I, 6: 80. 1848. Marsilea vestita var. tenuifolia (Engelm.) Underw. & Cook. On the edge of lakes, in shallow beds of creeks, and in periodically inundated depressions, particularly in old buffalo wallows. Blackland Prairies: Travis County. Edwards Plateau: In the extreme eastern counties. Plains Country: Mitchell and Taylor counties, near the southern boundary. MARSILEA UNCINATA A. Br., Flora 22: 304. 1839. This species, unlike M. tenuzfolia and M. vestita, apparently needs a constant supply of water since it is found only in or on the edge of per- manent ponds, along spring branches and in shallow water of brooks, ditches and bayous. Coastal Prairies: Aransas County, in the extreme west. Rio Grande Plains: Dimmit County and in counties on the northern and eastern borders. Blackland Prairies: Of general but sparse distribution. Edwards Plateau: Val Verde County and in several of the southern and eastern border counties. Trans-Pecos: Jeff Davis County. Plains Country: Potter County. MarsiLea vestTITaA Hook. & Grev., Ic. Fil. 2: pl. 159. 1831. M arsilea mucronata A. Br. This, the most widespread Marsilea in Texas, usually occurs in black waxy mud along streams and rivers, in and about ponds, in silt of lakes, and in ditches or depressions such as old buffalo wallows in prairies which are periodically imundated. Although a temporary pool may become powdery dry the bony sporocarps of this species remain undamaged until water again makes the depression a quagmire. Some plants closely resemble those of M. macrocarpa, with which it is occasionally confused. However, the solitary sporocarp readily distinguishes it from that species. Coastal Prairies: Aransas, Galveston and Wharton counties. Rio Grande Plains: Rather frequent and generally distributed. Blackland Prairies: Sparsely distributed from Dallas County southward. Edwards Plateau: In the eastern and western parts. : Trans-Pecos: El Paso and Pecos counties. Plains Country: Lynn, Randall, Taylor and Tom Green counties. Pruvuuaria Linnaeus PILULARIA AMERICANA A. Br., Monatsb. Kén. Akad. Wiss. Berlin, 1863: 435- 1864. ; In shallow temporary pools on rocky flats and depressions in clayey prairies. Edwards Plateau: Burnet County. Rare. 1949| CorRELL: TEXAS PTERIDOPHYTA 271 EQUISETACEAE. Horsetail Family Eaqutsetum [Tourn.| Linnaeus EQUISETUM KANSANUM Schaffner, Ohio Nat. 13: 21. 1912. In clayey or sandy soil of prairies, sloughs, along irrigation ditches, lake-shores, on bluffs, and among grasses on marshy banks. Blackland Prairies: Somervell County. It is represented here by forma caespitosum (A. A. Eaton) Broun. Edwards Plateau: Bexar and Blanco counties. Trans-Pecos: Represented in several southern and central counties by the typical form as well as forma caespitosum and forma variegatotdes (A. A. Eaton) Broun. Plains Country: Represented in several northern and central counties by the typical form and forma caespitosum. EQUISETUM LAEVIGATUM A. Br., Am. Journ. Sci. & Arts 46: 87. 1844. Equisetum hyemale L. var. intermedium A. A. Eaton. Fig. 17. An unusual abnormality in Equisetum prealtum, in which nodal and spiral sheaths occur in the same plant. Left: plant collected in Tarrant County, Texas; right: close-up of lower half of plant to show detail. 272 WRIGHTIA [Vou. 1, No. 5 On sandy banks of lakes and streams, seepage slopes, in meadows, prairies, and rocky creek beds of canyons. Coastal Prairies: Harris County. Blackland Prairies: In the north-central and southern parts. Edwards Plateau: In the central part. Trans-Pecos: Jeff Davis County. Plains Country: Potter and Randall counties. EQUISETUM PREALTUM Raf., Fl. Ludov. 13. 1817. Fig. 17. Equisetum robus- tum A. Br.; E. hyemale of authors, not Linneaus; EL. hyemale var. robustum (A. Br.) A. A. Eaton; E. hyemale var. affine (Engelm.) A. A. Eaton. Along moist sandy banks of streams, in alluvial flats and on seepage slopes. Timber Belt: Harrison County. Coastal Prairies: Harris County. Blackland Prairies: Represented in the northern and southern parts by the typical form and forma Drummondii (Milde) Broun and forma texana (Milde) Broun. Edwards Plateau: In the southern and northern regions. Trans-Pecos: El Paso and Jeff Davis counties. : Plains Country: Randall and Taylor counties. It is also represented in Taylor County by forma Drummondii and forma texana. LYCOPODIACEAE. Clubmoss Family Lycopropium [Dill.| Linnaeus LycopopruM ADPRESSUM (Chapm.) Lloyd & Underw., Bull. Torr. Bot. Club 27: 153. 1900. Lycopodium alopecuriodes var. adpressum Chapm. In sandy bogs, depressions in prairies and savannahs, and in moist open pinelands. Timber Belt: In the southern and west-central part. Coastal Prairies: Jefferson and Orange counties. Blackland Prairies: In several east-central counties. LYCOPODIUM ALOPECUROIDES L., Sp. Pl. 1102. 1753. This is apparently the first report of this species from Texas. In bogs, sphagnous meadows and prairies, moist open pinelands, and on seepage slopes. Coastal Prairies: Orange County. LyYCOPODIUM ALOPECUROIDES Var. PINNATUM (Chapm.) Lloyd & Underw., Bull. Torr. Bot. Club 27: 155. 1900. This is apparently the first report of this variety from Texas. Habitat similar to that of the typical form. Blackland Prairies: Travis County. LycopopIUM CAROLINIANUM L., Sp. Pl. 1104. 1753. In sphagnous meadows, depressions in savannahs and prairies and open moist pinelands. 1949] CorRELL: TEXAS PTERIDOPHYTA 273 Timber Belt: Jasper County. The specimens which my wife and I obtained in a savannah north of Kirbyville probably represents the first collection of this species from Texas. SELAGINELLACEAE. Spikemoss Family SELAGINELLA Beauvois SELAGINELLA Apops (L.) Spring ex Mart., Fl. Bras. 17: 119. 1840, as “apus’’. Selaginella ludoviciana A. Br. In sandy bogs, swamps, wet meadows, and about springs and along streams in seepage areas. Timber Belt: In several eastern and southern counties. Coastal Prairies: In the extreme northeastern border counties. Blackland Prairies: Austin and Ellis counties. Edwards Plateau: Eastern and southern parts. SELAGINELLA ARIZONICA Maxon, Smiths. Mise. Coll. 72°: 5, pl. 3. 1920. On rocky slopes and boulders. Edwards Plateau: Mason County; representing an isolated locality from the Trans-Pecos. Trans-Pecos: Brewster, Culberson and Presidio counties. SELAGINELLA Coryr Weatherby, Am. Fern Journ. 36: 51. 1946. On large boulders and ledges. Trans-Pecos: Brewster and Jeff Davis counties. SELAGINELLA LEPIDOPHYLLA (Hook. & Grev.) Spring, Mém. Acad. Brux. 24': 72. 1850. On dry limestone ledges and bluffs, around and in the protection of boulders and on rocky slopes. Edwards Plateau: Crockett, Uvalde and Val Verde counties. Trans-Pecos: Rather frequent and of general distribution. SELAGINELLA MuTIcA D. C. Eaton ex Underw., Bull. Torr. Bot. Club 25: 128. 1898. In crevices of shaded cliffs and ledges. Trans-Pecos: Confined to several southern and western counties. SELAGINELLA MUTICA Var. LIMITANEA Weatherby, Journ. Arn. Arb. 25: 414. 1944. On shaded rocky hillsides and crevices of cliffs and ledges. Trans-Pecos: El Paso and Jeff Davis counties. SELAGINELLA MUTICA var. TEXANA Weatherby, Journ. Arn. Arb. 25: 414. 1944. On shaded rocky hillsides and in crevices of cliffs. Trans-Pecos: Brewster County. SELAGINELLA NEOMEXICANA Maxon, Smiths. Mise. Coll. 72°: 2, pl. 1. 1920. On dry rock ledges, cliffs and canyon walls, slopes and boulders. 274 WRIGHTIA , [Vou:. t, Novs Trans-Pecos: El Paso and Presidio counties. Rare. SELAGINELLA PILIFERA A. Br., Ind. Sem. Hort. Berol. App. 1857: 20. 1857. Selaginella Pringlei Baker; S. pilifera var. Pringlet (Baker) Morton. On rocky slopes and floors of canyons in thinly wooded areas, and in crevices of canyon walls. Trans-Pecos: Apparently confined to several counties in the south and central parts. . SELAGINELLA RippeLLi Van Eseltine, Contr. U. 8. Nat. Herb. 20: 162, 1918. In sandy or gravelly soil in open woods, on sunny ledges and open rocky slopes, in shallow pockets among boulders, and on high dry sandy ridges. Timber Belt: In the southern part where it occurs along an irregular ridge of sandstone outcrops. Rio Grande Plains: Atascosa County, in the northwest part. Blackland Prairies: In the southern half.~ Edwards Plateau: In the eastern part. SELAGINELLA RUPINCOLA Underw., Bull. Torr. Bot. Club 25: 129. 1898. On shaded cliffs and ledges of granite, open rocky hillsides, and in depressions on large boulders. Trans-Pecos: Brewster, Jeff Davis and Presidio counties. SELAGINELLA SHELDONI Maxon, Proc. Biol. Soc. Wash. 31: 171. 1918. On rock ledges and talus slopes, edge of rim-rock, in soil or on rocks along streams, commonly forming large mats. ' Edwards Plateau: In the eastern and western parts. Trans-Pecos: Western and southern parts. Plains Country: Mitchell County. SELAGINELLA UnpEerRwoopit Hieron. ex Engelm. & Prantl, Pflanzenfam. 1: 714. 1901. On shaded cliffs and ledges, and floors of canyons. Trans-Pecos: Brewster and Jeff Davis counties. SELAGINELLA Wricuttt Hieron., Hedwigia 39: 208. 1900. On dry rocks, open hills, exposed flat rock outcrops of limestone, and on walls of canyons, frequently forming large mats. Edwards Plateau: Rather generally distributed and frequent. Trans-Pecos: Central and southern parts. Plains Country: Upton County, in the extreme south. ISOETACEAE. Quillwort Family Isornres Linnaeus ISOETES LITHOPHILA Pfeiffer, Ann. Mo. Bot. Gard. 9: 135, pl. 16. 1922: In shallow depressions and temporary pools and on rock outcrops 20 mountains of granite. d 1949] CoRRELL: TEXAS PTERIDOPHYTA 275 Edwards Plateau: Burnet and Llano counties. IsOETES MELANOPODA Gay & Durieu, Bull. Soc. Bot. France 11: 102. 1864. In shallow ponds, old buffalo wallows, wet thickets and woods, in seasonal streams and temporary sedge-grass puddles in meadows and prairies, and in temporary pools on granite outcrops. Timber Belt: Angelina County. Coastal Prairies: Harris County. Blackland Prairies: Bastrop, Dallas and Tarrant counties. Edwards Plateau: Burnet and Llano counties. EXCLUDED SPECIES Athyrium Filix-femina (L.) Roth ex Mertens, Arch. Bot. 2': 106. 1799. This species was perhaps first reported from Texas by Bush (1903). Misdetermined specimens of Athyrium asplenioides were undoubtedly the basis for this report. I have seen only one of the specimens cited by Bush; his number 985. It was typical A. asplenioides. Palmer’s (1919) citations of this species are also referable to A. asplenioides. Athyrium Filix-femina is a northern species which has not been found south of Colorado. Cheilanthes microphylla Sw., Syn. Fil. 127. 1806. I have referred to Chetlanthes alabamensis all material from Texas previously identified as this species. It may be that I am in error here. However, it is my opinion that we do not have any collections from Texas sufficiently distinct to separate them from C. alabamensis. Cheilanthes myriophylla Desv., Berl. Mag. 5: 328. 1811. Reverchon (1903) and others reported this fern from Texas. The re- ports of this primarily South American species in Texas are undoubtedly based on misdeterminations of some native Texas species. Diplazium acrostichoides (Sw.) Butters, Rhod. 19: 178. 1917. Athyrium acrostichoides (Sw.) Diels ex Engelm. & Prantl. In June, 1903, Bush reported this species as having been collected by Reverchon in Wood County in the Timber Belt. However, Reverchon, who published a list of the ferns and fern allies in Texas in April, 1903, did not include this species. I have seen no specimens from farther west than West Feliciana Parish, in eastern Louisiana. It seems best to me to exclude this species from the flora of Texas until its occurrence in the state is verified. Dryopteris ludoviciana (Kunze) Small, Ferns Southeastern States 281. 1938. Dryopteris floridana (Hook.) Kuntze; Nephrodium floridanum Hook. This species was reported by Reverchon (1903) as having been collected by J. M. Fetherolf in Newton County in the Timber Belt. It was also re- ported from Texas by Cory and Parks (1937). I have not seen any speci- mens of this species west of East Baton Rouge Parish, Louisiana. 276 WRIGHTIA [Vou. 1, No. 5 Dryopteris noveboracensis (L.) Gray, Man., ed. 1. 630. 1848. Cory and Parks (1937) reported this species from the Timber Belt. I have seen no specimen from the state. Its occurrence in northeastern Texas, however, is not improbable. I have seen a specimen collected by F. L. Harvey from swamps of southern Arkansas. Dryopteris spinulosa (O. F. Mill.) Watt var. intermedia (Muhl.) Underw. Our Nat. Ferns, ed. 4. 116. 1893. Dryopteris intermedia (Muhl.) Gray. A specimen of Athyrium asplenioides in the S. M. Tracy Herbarium collected in Nacogdoches County by H. B. Parks and labeled Dryopteris intermedia is probably the basis for Cory and Parks’ (1937) report of this northern and eastern plant in Texas. Gymnopteris Ehrenbergiana (Klotz.) C. Chr., Ind. Fil. 341. 1905. Gymno- gramma Ehrenbergiana Klotz. Reverchon (1903) reported this central Mexico species from Texas. This report was undoubtedly based on misdetermined specimens of Bom- meria hispida. Lycopodium inundatum L. var. Bigelovii Tuckerm., Am. Journ. Sei. 45: 47. 1843. A specimen of sterile moss (Polytrichon sp.) from Mangum Lake, Shelby County, collected by H. B. Parks and identified as this species (in S. M. Tracy Herbarium) is probably the basis for Cory and Parks’ (1937) report of this northeastern plant in Texas. Lycopodium lucidulum Michx., Fl. Bor.-Am. 2: 284. 1803. Cory and Parks (1937) reported this species from the Timber Belt and Coastal Prairies. Undoubtedly, the report of this eastern and northern species in Texas is based on misdeterminations of some native Texas species. Lygodium palmatum (Bernh.) Sw., Syn. Fil. 154. 1806. A specimen of Lygodium japonicum in the 8. M. Tracy Herbarium from Orange County, identified as L. palmatum, is probably the basis for Cory and Parks’ (1937) report of this northern and eastern species in Texas. Notholaena Fendlerit Kunze, Farnkr. 2: 87, pl. 136. 1851. Bush (1903) and others reported this species from Texas. I have not seen any specimen from the state. Its occurrence, however, in the Trans- Pecos is not improbable since its range extends into New Mexico. Pellaea mucronata D. C. Eaton in Torr., Rept. U. 8. & Mex. Bound. Surv. 2. $44, 3650. Broun (1938) included Texas in the range of this species. I have, how- ever, seen no specimen from Texas. This species is usually found on shady rock ledges and slopes. _ Polypodium furfuraceum Schlecht. & Cham., Linnaea 5: 607. 1830. Cory and Parks (1937) reported this species from the Trans-Pecos. Undoubtedly, the report of this central Mexico species in Texas is based on a misdetermination of some native Texas species. 1949] CORRELL: TEXAS PTERIDOPHYTA 277 Polypodium virginianum L., Sp. Pl. 1085. 1753. Cory and Parks (1937) reported this species as occurring in the Black- land Prairies. I have seen no specimen from Texas. I have seen specimens, however, as far southwest as Logan County, Arkansas. Polypodium vulgare L. forma pygmaewm (Schur) Broun, Index N. Am. Ferns 144. 1938. Broun (1938) includes western Texas in the range of this variety. I have seen no specimens from Texas. However, since the typical form is found in New Mexico, its occurrence in the Trans-Pecos is not improbable. Selaginella arenicola Underw., Bull. Torr. Bot. Club 25: 541. 1808. This species was first reported from Texas by Reverchon (1903) as growing in sand and among rocks in the granite region of Llano County. In 1r919, Palmer cites specimens from Hardin and Leon Counties. The specimen from Leon County, which I have seen, is S. Riddelliiz. Cory and Parks (1937) were the last to include this species in the flora of Texas. According to Clausen (1946), with whom I agree, this species is confined to Georgia and Florida. Reports from Texas are, for the most part, based on misdetermined plants of S. Riddellii which closely resembles S. arenicola in habit. Selaginella densa Rydb., Mem. N. Y. Bot. Gard. 1: 7. 1900. Cory and Parks (1937) reported this species from the Trans-Pecos. I have seen no specimen from Texas. Its occurrence in western Texas, how- ever, is not improbable since the species is found in New Mexico, Selaginella rupestris (L.) Spring, Flora 21: 149, 182. 1838. Reverchon (1903) was perhaps the first to report this species from Texas, where he said it occurred on dry exposed rocks in the southwest. Later, Palmer (1919) cited specimens from the state. These specimens, however, were either misdetermined S. Sheldoni (Palmer 11389) or S. Wrightit (Palmer 10146). In 1937, Cory and Parks included this species in their catalogue of Texas plants. In 1946, Clausen gives the range of this species as only south to central Oklahoma and northern Arkansas. I do not believe this species occurs in Texas. Selaginella tortipila A. Br., Ann. Sci. Nat. V, 3: 271. 1865. Selaginella Sherwoodii Underw. Cory and Parks (1937) reported this species as occurring in the Timber Belt and Coastal Prairies. Undoubtedly, the report of this eastern Pied- mont and Blue Ridge Mountains species in Texas ‘is based on misdetermi- nations of some native Texas species. POSSIBLE FUTURE ADDITIONS TO THE FERN FLORA OF TEXAS The following species are cited merely to call the attention of collectors to the possibility of their occurrence in Texas. Thus forewarned, botanical collectors and Texas students of ferns can be on the watch for these species. 278 WRIGHTIA [Vou. 1, No. 5 Adiantum pedatum L., Sp., Pl. 1095. 1753. Specimens of this species have been sillected near Page, Le Flore County, Oklahoma, about 60 miles north of Red River County, Texas. It is usually found on rich wooded slopes. Asplenium pinnatifidum Nutt., Gen. 2: 251. 1818. This species occurs on rocks near Robbers Cave, Latimer County, Oklahoma, about 60 miles north of Lamar and Red River counties, Texas. It is usually found in dry crevices of non-caleareous rocks. Cystopteris bulbifera (L.) Bernh., Neu. Journ. Bot. Schrad. 1°: to. 18or. Specimens have been collected above Gray’s Ranch (Otero or Eddy County), Guadalupe Mountains, New Mexico. The Guadalupe range ex- tends into Culberson County, Texas. This species is usually found on shaded ledges or talus slopes of limestone. Dryopteris marginalis (L.) Gray, Man., ed. 1. 632. 1848. This species has been found near Cache, Comanche County, Oklahoma, about 30 miles north of Wichita County, Texas. It has also been collected near Page, Le Flore County, Oklahoma, about 60 miles north of Red River County, Texas. It is usually found on shaded ledges, talus slopes or in rocky woods. ACKNOWLEDGMENTS I wish to express my appreciation to the curators of the various her- baria who have kindly cooperated in loaning me specimens. To Mr. C. A. Weatherby and Dr. C. V. Morton I am indebted for advice and assistance in determining some of the most difficult collections. LITERATURE CITED Broun, Maurice. 1938. Index to North American Ferns. 217 pp. Bush, Benjamin F. 1903. A List of the Ferns of Texas. Bull. Torr. Bot. Club 30: 343-358. Clausen, Robert T. 1946. Selaginella, Subgenus Euselaginella in the South- eastern U. 8. Am. Fern Journ. 36: 65-82. Cory, Victor L. and Harris B. Parks. 1937 (1938). Catalogue of the Flora of the State of Texas. Bull. Texas Agr. Expt. Sta. 550, 130 pp., Map. Palmer, Ernest J. 1919. Texas Pteridophyta. Am. Fern Journ. 9: 17~2?) 50-56, 81-85. . 1927. Ferns of the Davis Mountains, Western Texas. Am. Fern Journ. 17: 77-85. [see additions in Am. Fern Journ. 20: 138-142. 193° (1931), also from Chisos Mts.| Reverchon, Julian. 1903. The Fern Flora of Texas. Fern Bull. 11: 33~38- Division of Plant Exploration and Introduction Bureau of Plant Industry, Soils, and Agricultural Engineering Agricultural Research Administration U.S. Department of Agriculture NOTES REQUEST FOR SEED OF LOBELIA CARDINALIS Wray M. BowpEeNn Viable seeds of any of the four varieties of Lobelia Cardinalis subsp. graminea (Lam.) MeVaugh will be much appreciated; see Ann. Mo. Bot. Gard. 27:347. The seed should be very ripe when collected. If unripe, the plants may be dug and transferred to a convenient location, or the flowering stalks may be cut and placed in jars of water in a sunny place until ripe. An herbarium specimen from one or more of the plants from which the seed is taken will assist in identification of the material. In case the collector would like part of the material returned to him after it has been identified, extra specimens should be collected and a note attached requesting return of duplicate material. Seeds may be sent by letter; herbarium specimens should have a customs declaration form stating that the contents are “dried plant specimens, for scientific pur- poses only.’”’ The appearance of the pressed flowers will be much im- proved if a piece of waxed paper is placed against the inflorescence for the first 24-48 hours. Mail Address: Dr. Wray M. Bowden, Division of Botany and Plant Pathology, Central Experimental Farm, Ottawa, Canada.