UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN MATURAL HIST. SURVEY Botany O. 18 FLORA COSTARICENSIS William Burger, Editor Family #200 Acanthaceae L. H. Durkee Family #201 Plantaginaceae William Burger December 31. 1986 Publication 1372 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 114 36 144 146 i . 101 !i 13 ; 142 II 47 .! FLORA COSTARICENSIS Family #200 Acanthaceae Family #201 Plantaginaceae Other Fieldiana: Botany Titles Available in the Flora Costaricensis Series, William Burger, Editor Family #40, Casuarinaceae; Family #41, Piperaceae. By William Burger. 1971; 227 pages, 14 illus. Publication No. 1140, $18.25 Family #42, Chloranthaceae; Family #43, Lacistemaceae; Family #44, Salicaceae; Family #45, Garry- aceae; Family #46, Myricaceae; Family #47, Juglandaceae; Family #48, Bataceae; Family #49, Be- tulaceae; Family #50, Fagaceae; Family #51, Ulmaceae; Family #52, Moraceae; Family #52a, Can- nabaceae; Family #53, Urticaceae. By William Burger, Luis D. Gomez, Donald E. Stone, and John G. Furlow. 1977; 291 pages, 30 illus. Publication No. 1270, $23.25 Family #15, Gramineae. By Richard W. Pohl. 1980; 608 pages, 224 illus. Publication No. 1313, $45.00 Family #54, Podostemaceae; Family #55, Proteaceae; Family #56, Olacaceae; Family #57, Opiliaceae; Family #58, Loranthaceae; Family #59, Aristolochiaceae; Family #60a, Hydnoraceae; Family #60b, Rafflesiaceae; Family #6 1 , Balanophoraceae; Family #62, Polygonaceae; Family #63, Chenopodiaceae; Family #64, Amaranthaceae; Family #65, Nyctaginaceae; Family #66, Phytolaccaceae; Family #67, Aizoaceae; Family #68, Portulacaceae; Family #69, Basellaceae; Family #70, Caryophyllaceae. By William Burger, Job Kuijt, Kerry Barringer, Luis D. Gomez, and Richard Baker. 1983; 255 pages, 42 illus. Publication 1350, $25.50 Order by publication number, or ask for a free copy of our price list. Address all requests to: FIELD MUSEUM OF NATURAL HISTORY Division of Publications Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496, U.S.A. FIELDIANA Botany NEW SERIES, NO. 18 FLORA COSTARICENSIS William Burger, Editor Family #200 Acanthaceae L. H. Durkee Professor of Biology Department of Biology Grinnell College Grinnell. Iowa 501 12 Family #201 Plantaginaceae William Burger Curator, Vascular Plants Department of Botany Field Museum of Natural History Chicago, Illinois 60605-2496 Accepted for publication March 14, 1985 December 31, 1986 Publication 1372 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1986 Field Museum of Natural History Library of Congress Catalog Card Number: 78-172358 ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations ACKNOWLEDGMENTS vi ACANTHACEAE by L. H. Durkee 1 Conspectus of the Subfamilies and Genera in Costa Rica 2 Illustrations of Acanthaceae 3-20 Key to the Genera of Acanthaceae in Cos- ta Rica 21 Acanthus 23 Aphelandra 23 Asystasia 30 Barleria 31 Blechum 32 Bravaisia 33 Buceragenia 33 Carlowrightia 34 Chaetochlamys 35 Chaetothylax 35 Chamaeranthemum 36 Dicliptera 37 Dyschoriste 40 Elytraria 41 Graptophyllum 41 Habracanthus 42 Hansteinia 43 Henrya 45 Herpetacanthus 46 Hygrophila 47 Hypoestes 47 Justicia 48 Louteridiwn 62 Megaskepasma 62 Mendoncia 63 Nelsonia 65 Odontonema 66 Poikilacanthus 67 Pseuderanthemum 67 Razisea 70 Ruellia 72 Sanchezia 81 Spathacanthus 81 Streblacanthus 82 Teliostachya 83 Tetramerium 83 Thunbergia 84 Trichanthera 86 PLANT AGINACEAE by William Burger .. 87 Plantago 87 Illustration of Plan taginaceae 89 INDEX . 91 1. Acanthus mollis; Aphelandra: A. auran- tiaca var. aurantiaca; A. aurantiaca var. stenophylla; A. campanensis; A. scabra\ A. dolichantha 3 2. Aphelandra: A. golfodulcensis; A. leo- nardii; A. lingua- bovis; A. seibertii; A. sin- clairiana; A. storkii 4 3. Aphelandra: A. tonduzii; A. tridentata; Asystasia gangetica; Barleria micans; Blechum: B. costaricense; B. brownei ... 5 4. Bravaisia integerrima; Buceragenia glan- dulosa; Chaetothylax leucanthus; Cha- maeranthemum durandii; Dicliptera: D. imbricata; D. iopus 6 5. Dicliptera: D. pallida; D. podocephala; D. skutchii; D. trifurca; D. unguiculata; Dys- choriste valeriana; Elytraria imbricata . . 1 6. Graptophyllum pictum; Habr acanthus sil- vanticus; Hansteinia: H. blepharorhachis; B. sessilifolia; H. stricta; H. ventricosa . . 8 7. Henrya scorpioides; Herpetacanthus pan- amensis; Hygrophilia costata; Hypoestes phyllostachya; Justicia: J. angustibractea- ta; J. aurea 9 8. Justicia: J. brenesii; J. candelariae; J. car- thaginensis; J. chamaephyton; J. comata; J. costaricana 10 9. Justicia: J. crenata; J. brandegeana; J. macrantha; J. metallica; J. oerstedii; J. orosiensis 11 10. Justicia: J. parvibracteata; J. pectoralis; J. pittieri; J. refractifolia; J. sarapiquensis; J. skutchii 12 1 1 . Justicia: J. spicigera; J. tinctoria; J. ton- duzii; J. trichotoma; J. urophylla; J. val- erii 13 12. Louteridium costaricensis; Mendoncia: M. brenesii; M. costaricana; M. lindavii; M. retusa; M. tonduzii 14 1 3. Megaskepasma erythrochlamys; Nelsonia canescens; Odontonema tubiforme; Po- ikilacanthus macranthus; Pseuderanthe- mum atropurpureum; Pseuderanthemum cuspidatum 15 14. Pseuderanthemum: P. pittieri; P. praecox; P. standleyi; Razisea: R. citrina; R. spi- cata; R. voilburi 16 15. Ruellia: R. biolleyi; R. cooperi; R. gemi- niflora; R. golfodulcensis; R. inundata; R. jussieuoides 17 16. Ruellia: R. metallica; R. nudiflora; R. pa- lustris; R. paniculata; R. pittieri; R. prae- clara 18 17. Ruellia: R. stemonacanthoides; R. ton- duzii; R. tubiflora var. hirsuta; R. tubiflora var. tetrastichantha\ Spathacanthus hoff- manni; Streblacanthus monospermus ... 19 18. Teliostachya alopecuroidea; Tetramer- ium nervosum; Thunbergia alata; Tri- chanthera gigantea 20 19. Plantago australis 89 Acknowledgments This is the fifth issue of the Flora Costaricensis. The first dealt with Piperaceae (Fieldiana, Bot. 35, 1 97 1). The second included families numbered 42 through 53, Chloranthaceae through Urticaceae (Fieldiana, Bot. 40, 1977). The third issue con- tained the Gramineae by Richard Pohl (Fieldiana, Bot., new series, No. 4, 1980). The fourth issue included families numbered 54 through 70, Podo- stemaceae through Caryophyllaceae (Fieldiana, Bot., new series, No. 13, 1983). We are especially grateful for the financial assistance of the National Science Foundation, which has aided this program for many years, both at Field Museum and in Cos- ta Rican fieldwork. The project at Field Museum has been supported most recently by NSF grant DEB-8103184, through the Biological Research Resources Program. The staff and the facilities of the Museo Nacional de Costa Rica have been a central resource and most helpful to our work for more than two decades. This issue includes the large and attractive Acanthaceae family. We are especially fortunate to have the account of this family prepared by Dr. Durkee, a specialist in the family who has spent considerable time in Costa Rica and earlier published an account of the Acan- thaceae for the Flora of Panama. Technical assistance and fieldwork for Durkee have been financed by grants from Grinnell Col- lege and the Associated Colleges of the Midwest. Work on the Acanthaceae was supported by loans from the following herbaria: Museo Nacional de Costa Rica, Duke University, Field Museum of Natural History, Missouri Botanical Garden, New York Botanical Garden, and the Museum of Nat- ural History, Smithsonian Institution. Robin Chazdon and Sarah Dewey provided able tech- nical assistance; Susan Durkee (SD), Laurie Engle, Kelly Hanick (KH), Lyla Kimura (LK), Kathleen Neff (KBN), Christopher Rasmussen (CAR), and Anthony Sheeder (TS) were the illustrators for the Acanthaceae. All of these individuals were un- dergraduates at Grinnell College during the time of their work. VI FLORA COSTARICENSIS Family #200 Acanthaceae Family #201 Plantaginaceae ACANTHACEAE By L. H. Durkee REFERENCES— C. E. B. Bremekamp, The delim- itation of the Acanthaceae. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. C. 56: 533-546. 1953. C. E. B. Bremekamp, Delimita- tion and subdivision of the Acanthaceae. Bull. Bot. Surv. India 7: 21-30. 1965. A. Cronquist, Acan- thaceae, pp. 963-966. In An Integrated System of Classification of Flowering Plants. 1981. L. H. Durkee, Acanthaceae. In Flora of Panama. Ann. Missouri Bot. Gard. 65: 156-283. 1978. E. C. Leonard, Acanthaceae. In P. C. Standley, Flora of Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1188-1263. 1938. E. C. Leonard, The Acan- thaceae of Colombia. Contr. U.S. Natl. Herb. 31. 1951-1958. G. Lindau, Acanthaceae. In A. Engler and G. Prantl (eds.), Nat. Pflanzenfam. 4(3b): 274- 354. 1895. Herbaceous to suffrutescent plants, some twining, less often shrubs, rarely trees. Leaves simple, opposite (ex- cept Elytraria of the subfamily Nelsoniodiae), exstipu- late, often with cystoliths on the upper surfaces of lam- inae (most evident in dried specimens). Flowers irregular to nearly regular, perfect, borne in spikes, racemes, pan- icles, cymes, or singly and often subtended by a con- spicuous bract with 2 bracteoles; calyx synsepalous at least basally, persistent, the segments (3, 4) 5(-16); the corolla gamopetalous, the limb 5-lobed or 2-lipped, rare- ly 1 -lipped; stamens adnate to the corolla tube, 4 and didynamous or 2 only, staminodes often present in flow- ers with 2 stamens; rarely all 5 stamens fully developed (none in ours); the anther cells 2 or 1, longitudinally dehiscent, the pollen sacs juxtaposed or sometimes su- perposed; ovary 2-celled, superior, 2-10 ovules in each cell, the style filiform, simple, the stigmas 1 or 2. Fruit a loculicidal capsule (except Mendoncia with its dru- paceous fruits), often explosively dehiscent; seeds usually flattened, mostly borne on hook-shaped funicles, some- times on papilliform funicles, the testa smooth or rough- ened, often mucilaginous when wetted. The family Acanthaceae includes about 250 genera and 2,500 species, widespread in both New and Old World Tropics, with only a few species occurring in warm temperate regions. The family is divided into four subfamilies according to Lin- dau. More recent considerations have raised each of the subfamilies Mendoncioideae, Nelsonioi- diae, and Thunbergioideae to family ranks. I am in agreement with Cronquist who recognizes the Mendonciaceae, but who feels that the Nelson- ioideae and Thunbergioideae represent stages along the way between the Scrophulariaceae and the Acanthoideae, the fourth subfamily of Lindau. While they have clear affinity with the Acanthoi- deae, they are anomalous with either family, but insufficiently distinct to warrant separate status. Furthermore, Cronquist does not feel that the con- necting forms are sufficiently numerous to warrant uniting the Acanthaceae and the Scrophulariaceae. The subfamily Mendoncioideae is included in this treatment as a matter of convenience since it fol- lows traditional usage. The Acanthaceae can be recognized by the prominent cystoliths appearing with magnifica- tion as small, cylindrical concretions or short, raised lines on the upper surface of the dried leaves, younger stems, the branches of the inflorescence, and the calyx. Genera lacking these are few and have other exceptional features such as the fruit. The genus Mendoncia is unique, with a drupa- ceous fruit, for example. The only genera lacking BURGER: FLORA COSTARICENSIS cystoliths and having the more typical capsular fruit with hooklike funicles are the genera Aphe- landra and Megaskepasma. Seeds are frequently ejected forcefully by the hook-shaped funicles when the mature capsules are moistened by rainfall. In opened or dehisced capsules, the funicles can be seen as hook-shaped projections. Many species of the family also have imbricate bracts subtending the flowers. These bracts may be green, yellow, bright red, or orange. While most genera of the family are restricted to either New or Old World Tropics, a few are found in both. These include Justicia and Ruellia, the largest genera in the family, with about 410 and 250 species, respectively. Typically, most of these species are found in the tropics and sub- tropics, with a few extending into temperate areas. The major economic importance of the family is horticultural. Cultivation is often restricted to greenhouses, however, since in spite of attractive foliage and flowers, they are difficult to grow. Jus- ticia brandegeana, the shrimp plant, with its showy spikes of reddish bracts is perhaps the best known in the United States. Seeds of Thunbergia alata, the clock vine or black-eyed susan, are frequently sold in gardening stores in the United States, and in the tropics several other species are grown for their attractive foliage and flowers. Many other genera are cultivated; these include: Acanthus, Aphelandra, Crossandra, Eranthemum, Fittonia, Graptophyllum, Hypoestes, Pachystachys, Ruellia, Sanchezia, and Strobilanthes. The following summary characterizes each of the four subfamilies as recognized here and indi- cates the Costa Rican genera belonging to each. Conspectus of the Subfamilies and Genera in Costa Rica SUBFAMILY I. Ac ANTHOIDEAE— Cylindrical silic- ified cystoliths commonly present in epidermal cells of stems and leaves. These are best observed under magnification of dried specimens. They are sometimes mistaken for strigose hairs. Among the Costa Rican genera, Aphelandra and Megaske- pasma lack cystoliths. Seeds borne on hooklike funicles in explosively dehiscent capsules. Most of the genera and species of the family are found in this subfamily: Acanthus, Aphelandra, Asystasia, Barleria, Blechum, Bravaisia, Buceragenia, Car- lowrightia, Chaetochlamys, Chaetothylax, Cha- maeranthemum, Dicliptera, Dyschoriste, Grapto- phyllum, Habracanthus, Hansteinia, Henrya, Herpetacanthus, Hygrophila, Hypoestes, Justicia, Louteridium, Megaskepasma, Odontonema, Poi- kilacanthus, Pseuderanthemum, Razisea, Ruellia, Sanchezia, Spathacanthus, Streblacanthus, Te- liostachya, Tetramerium, Tricanthera. SUBFAMILY II. MENDONCIOIDEAE— Mostly high- climbing vines without cystoliths. Upper surfaces of laminae sometimes with stellate bases of the hairs. Flowers enclosed by two fairly conspicuous bracts that are at first connate; calyx reduced to an entire or low-lobed annulus. Fruit a drupe: Mendoncia. SUBFAMILY III. NELSONIOIDEAE— Herbs without cystoliths in stems or leaves. Leaves opposite or alternate. Inflorescences mostly in bracteate spikes. Flowers with a more or less equally five-parted calyx. Fruit a capsule with papilliform funicles: Elytraria, Nelsonia. SUBFAMILY IV. THUNBERGIOIDEAE— Vines or shrubs. Cystoliths lacking in stems and leaves. Flowers with an annular or toothed calyx; corollas usually conspicuous; stamens four with two-celled anthers. Fruit a beaked capsule with pulvinate fu- nicles. The genera of this subfamily are all of Pa- leotropical origin, and Neotropical species are cul- tivated or escaped: Thunbergia. FIELDIANA: BOTANY ACANTHUS mollis 2cm APHELANDRA aurantiaca var. aurantiaca 5cm APHELANDRA aurantiaca var. stenophylla 1cm APHELANDRA campanensis APHELANDRA scabra APHELANDRA dolichantha FIG. 1. Acanthaceae: species of Acanthus and Aphelandra (in part) in Costa Rica. The 5-cm scale refers to foliage and inflorescences, but note that Acanthus mollis is at a different scale. BURGER: FLORA COSTARICENSIS golfodulcensis 5cm seibertii sinclairiana FIG. 2. Acanthaceae: species ofAphelandra (second part) in Costa Rica. FIELDIANA: BOTANY APHELANDRA tonduzii APHELANDRA tridentata ASYSTASIA gangetica BARLERIA micans BLECHUM costaricense BLECHUM brownei 5cm FIG. 3. Acanthaceae: species of Aphelandra (concluding part), Asystasia, Barleria, and Blechum in Costa Rica. BURGER: FLORA COSTARICENSIS BRAVAISIA integerrima BUCERAGENIA glandulosa CHAETOTHYLAX eucanthus CHAMAERANTHMUM durandii DICLIPTERA 5cm imbricata FIG. 4. Acanthaceae: species of Bravaisia, Buceragenia, Chaetothylax, Chamaeranthemum, and Dicliptera (first part) in Costa Rica. DICLIPTERA iopus FIELDIANA: BOTANY DICLIPTERA pallida 5cm DICLIPTERA podocephala DICLIPTERA DICLIPTERA trifurca DICLIPTERA unguiculata DYSCHORISTE valeriana ELYTRARIA imbricata m / FIG. 5. Acanthaceae: species of Dicliptera (concluding part), Dyschoriste, and Elytraria in Costa Rica. BURGER: FLORA COSTARICENSIS GRAPTOPHYLLUM pictum HANSTEINIA blepharorhachis HANSTEINIA sessilifolia - HANSTEINIA stricta HANSTEINIA ventricosa FIG. 6. Acanthaceae: species of Graptophyllum, Habracanthus, and Hansteinia in Costa Rica. FIELDIANA: BOTANY HENRYA scorpioides HERPETACANTHUS panamensis HYGROPHILA costata HYPOESTES phyllostachya JUSTICIA angustibracteata FIG. 7. Acanthaceae: species ofHenrya, Herpetacanthus, Hygrophilia, Hypoestes, and Justicia (first part) in Costa Rica. BURGER: FLORA COSTARICENSIS JUSTICIA brenesii 5cm candelariae carthaginensis comata costaricana FIG. 8. Acanthaceae: species ofJusticia (second part) in Costa Rica. 10 FIELDIANA: BOTANY JUSTICIA \\ crenata scm • metallica oerstedii orosiensis FIG. 9. Acanthaceae: species ofJusticia (third part) in Costa Rica. BURGER: FLORA COSTARICENSIS 11 JUSTICIA „„ parvibracteata 5cm pectoralis pittieri sarapiquensis FIG. 10. Acanthaceae: species ofJusticia (fourth part) in Costa Rica. 12 FIELDIANA: BOTANY JUSTICIA spicigera 5cm tinctoria tonduzii urophylla valerii FIG. 1 1. Acanthaceae: species ofJusticia (concluding part) in Costa Rica. BURGER: FLORA COSTARICENSIS 13 LOUTERIDIUM costaricense MENDONCIA 5cm MENDONCIA costaricana MENDONCIA lindavii MENDONCIA tonduzii FIG. 12. Acanthaceae: species of Louteridium and Mendoncia in Costa Rica. 14 FIELDIANA: BOTANY MEGASKEFttSMl 5cm erythrochlamys NELSONIA canescens 1cm POIKILACANTHUS macranthus 1cm PSEUDERANTHEMUM PSEUDERANTHEMUM cuspidatum FIG. 13. Acanthaceae: species of Megaskepasma, Nelsonia, Odontonema, Poikilanthus, and P sender anthemum (first part) in Costa Rica. BURGER: FLORA COSTARICENSIS 15 PSEUDERANTHEMUM pittieri 2 cm PSEUDERANTHEMUM praecox 5cm PSEUDERANTHEMUM standleyi citrina 1 cm RAZISEA spicata RAZISEA wilburii FIG. 14. Acanthaceae: species of Pseuderanthemum (concluding part) and Razisea in Costa Rica. 16 FIELDIANA: BOTANY RUELLIA biolleyi y 1cm 5cm geminiflora 1cm golfodulcensis inundata jussieuoides FIG. 15. Acanthaceae: species of Ruellia (first part) in Costa Rica. BURGER: FLORA COSTARICENSIS 17 RUELLIA metallica 5cm nudiflora palustris paniculate 1cm pittieri praeclara FIG. 16. Acanthaceae: species of Ruellia (second part) in Costa Rica. 18 FIELDIANA: BOTANY 5cm RUELLIA stemonacanthoides RUELLIA tonduzii RUELLIA tubiflora var. hirsuta RUELLIA tubiflora var. tetrastichantha 1cm STREBLACANTHUS monospermus FIG. 17. Acanthaceae: species of Ruellia (concluding part), Spathacanthus, and Streblacanthus in Costa Rica. BURGER: FLORA COSTARICENSIS 19 1cm TELIOSTACHYA alopecuroidea 5cm TETRAMERIUM nervosum TRICHANTHERA gigantea THUNBERGIA alata FIG. 18. Acanthaceae: species of Teliostachya, Tetramerium, Trichanthera, and Thunbergia in Costa Rica. 20 FIELDIANA: BOTANY Key to Genera of Acanthaceae in Costa Rica 1 a. Fertile stamens 4 2a Ib. Fertile stamens 2 1 7a 2a. Plants usually scandent; seeds not borne on hooklike funicles 3a 2b. Plants not scandent, usually erect herbs or shrubs; seeds borne on hooklike funicles or, if on papilliform funicles, capsules not beaked 4a 3a. Fruit a drupe Mendoncia 3b. Fruit a beaked capsule with 2-4 seeds Thunbergia 4a. Corolla 1 -lipped; stamen filaments thick and prominent; leaves sinuate pinnatifid Acanthus 4b. Corolla 2-lipped, subequal or regular; stamen filaments narrow; leaves ovate, obovate, elliptic, or oblong, but not sinuate-pinnatifid 5a 5a. Calyx spathaceous, 2-lobed; fruits 6.5-8 cm long Spathacanthus 5b. Calyx 4- or 5-parted, not spathaceous; fruits less than 4 cm long 6a 6a. Anthers 1 -celled; cystoliths lacking in the leaves Aphelandra 6b. Anthers 2-celled in at least 1 pair of stamens; cystoliths present in the leaves 7a 7a. Calyx segments 4, unequal; bracts with spiny margins Barleria 7b. Calyx segments 5, either equal or unequal; bracts lacking spiny margins 8a 8a. One pair of stamens with 2-celled anthers, the other pair of stamens with 1 -celled anthers or sterile 9a 8b. Both pairs of stamens with 2-celled anthers lOa 9a. Corolla bilabiate; bracts loosely imbricate and conspicuous, 9-14 mm long Herpetacanthus 9b. Corolla nearly regular; bracts lax and inconspicuous, 1.5-2 mm long Chamaeranthemum lOa. Shrubs or trees; calyx segments often apically rounded or obtuse 1 la 1 Ob. Herbs, often more or less suffrutescent (rarely shrubs); calyx segments subulate or lanceolate, usually apically acute or acuminate 1 2a 1 la. Anthers basally rounded, not spurred; capsules 4-seeded, sericeous Tricanthera lib. Anthers basally spurred; capsules 8-seeded, glabrous Bravaisia 1 2a. Corolla conspicuously bilabiate, 1 cm long or less; capsules sessile (bilocular from base) . . 13a 12b. Corolla regular or nearly so; capsule stipitate (basal portion solid for at least 2 mm) . . 14a 1 3a. Inflorescence a dense spike; anther cells unequal; seeds 4 per capsule . . Teliostachya 13b. Inflorescence fasciculate in the leaf axils; anther cells equal; seeds 12-16 per capsule Hygrophila 1 4a. Inflorescences in 1 -sided racemes; corollas yellow with purplish throats Asystasia 1 4b. Inflorescences in cymes, heads, panicles, spikes, or flowers axillary; corollas variously colored, not yellow with purplish throats 1 5a 1 5a. Flowers in closely imbricate bracted, 4-sided spikes; bracts conspicuous, ovate . . Blechum 1 5b. Flowers otherwise; bracts mostly inconspicuous and linear 1 6a 1 6a. Calyx segments fused for '/j-'/i their length; anther cells basally spurred Dyschoriste \ 6b. Calyx segments parted nearly to base; anther cells basally rounded Ruellia 17a. Anthers 1 -celled, or if 2, one always much smaller than the other and usually abortive 18a 1 7b. Anthers 2-celled, the cells about equal in size 25a 1 8a. Corolla less than 5 mm long; staminodes present Buceragenia 1 8b. Corolla more than 5 mm long; staminodes absent 1 9a 1 9a. Inflorescences condensed, headlike, bracts white-ciliate, linear subulate 20a 1 9b. Inflorescences open, bracts otherwise 2 la 20a. Calyx segments 4; corolla ca. 1 5 mm long Chaetothylax 20b. Calyx segments 5; corolla more than 20 mm long Chaetochlamys 2 la. Corolla blue; inflorescence a loose, terminal thyrse (cymose panicles) Habracanthus BURGER: FLORA COSTARICENSIS 21 2 1 b. Corolla white, red, magenta, orange, or yellow; inflorescence of single flowers, spikes, racemes, compact thyrses, or racemose panicles 22a 22a. Leaves pink-dotted; corolla with 3-lobed upper lip and 1 -lobed lower lip Hypoestes 22b. Leaves lacking pink dots; corolla with 1 - or 2-lobed upper lip and 3-lobed lower lip . . 23a 23a. Inflorescences paniculate; corolla tube usually abruptly enlarged above the ovary Hansteinia 23b. Inflorescences a spike or narrow spikelike thyrse; corolla tube expanding gradually from the base 24a 24a. Inflorescences spicate; calyx 4-parted Streblacanthus 24b. Inflorescences a narrow spikelike thyrse; calyx 5-parted Razisea 25a. Inflorescences condensed, headlike; bracts white-ciliate, linear subulate 26a 25b. Inflorescences open, bracts otherwise 27a 26a. Calyx segments 4; corolla ca. 1 5 mm long Chaetothylax 26b. Calyx segments 5; corolla more than 20 mm long Chaetochlamys 27a. Cystoliths lacking in the leaves; seeds not borne on hooklike funicles 28a 27b. Cystoliths present in the leaves; seeds borne on hooklike funicles 30a 28a. Bracts large and showy, more than 20 mm long; corolla more than 30 mm long Megaskepasma 28b. Bracts small and inconspicuous, less than 10 mm long; corolla less than 15 mm long . 29a 29a. Bracts stiff and awn-tipped; leaves mostly more than 10 cm long Elytraria 29b. Bracts flaccid and awnless; leaves mostly less than 8 cm long Nelsonia 30a. Staminodes present 3 la 30b. Staminodes absent 35a 3 la. Anther cells basally spurred; flowers in fascicles of 3-8, loosely enclosed by bracts Sanchezia 31b. Anther cells not basally spurred; flowers not in fascicles enclosed by bracts 32a 32a. Bracts conspicuous with spiny margins, more than 2 cm long; calyx 4-parted, segments unequal Barleria 32b. Bracts inconspicuous, without spiny margins, less than 1 cm long; calyx 5-parted, segments equal 33a 33a. Corolla limb subequally 5-parted, tube narrowly cylindrical, limb spreading; stamens included (except Pseuderanthemum atropurpureum) Psenderanthemum 33b. Corolla limb 2-lipped, tube expanding; stamens exserted 34a 34a. Leaf blades mottled; corollas 4.5-8 cm long; plants cultivated Graptophyllum 34b. Leaf blades green; corollas 1.5-3 cm long; plants mostly wild Odontonema 35a. Flowers in dense spikes; bracts imbricate, often 4-ranked 36a 35b. Flowers borne in lax spikes, panicles, cymes, or thyrses; bracts not imbricate and not 4-ranked 38a 36a. Bracts more than 2 cm long; calyx segments widely dissimilar Barleria 36b. Bracts less than 2 cm long; calyx segments similar 37a 37a. Bracts 4-ranked; floral bracts not uniting to form an involucre Tetramerium 37b. Bracts not 4-ranked; floral bracts uniting almost to apex forming an involucre .... Henrya 38a. Stems 6-angled in cross section; calyx subtended by 2 partially united bracts Dicliptera 38b. Stems rounded to 4-angled; bracts distinct 39a 39a. Corolla lobes 4; mature dried capsules black or dark purple Carlowrightia 39b. Corolla lobes 5; mature dried capsules yellowish brown to dark brown 40a 40a. Calyx 3-parted Louteridium 40b. Calyx 5-parted 41a 4 1 a. Bracts large, to 44 mm long Megaskepasma 4 1 b. Bracts small, less than 20 mm long 42a 42a. Corolla more than 6 cm long; leaf pairs unequal (excluding J. trichotoma which has unequal leaf pairs); pollen polyporate Poikilacanthus 42b. Corolla less than 6 cm long (all species except /. aurea are less than 5 cm long); leaf pairs equal, with the exception of/, trichotoma; pollen mostly 2-porate, some 3- or 4-porate Justicia FIELDIANA: BOTANY Acanthus L. REFERENCE— T. H. Everett, The New York Bo- tanical Garden Illustrated Encyclopedia of Hor- ticulture 1: 21. 1980. Herbs or small shrubs. Leaves petiolate; laminae broad and pinnatifid. Inflorescences in terminal spikes; flowers sessile, borne singly in the axils of the bracts; bracteoles 2 per flower; calyx 4-merous. segments unequal; corolla 1 -lipped, the lip 3-lobed; stamens 4, filaments thick and prominent, anthers 1 -celled, wooly. Fruit a capsule. This is a genus of about 20 species in subtropical and tropical regions of the Mediterranean, Asia, and Africa. Two species are often cultivated for their appearance, A. montanus T. Anders and A. mollis L. Acanthus mollis L., Sp. pi. 639. 1753. Figure 1. Herbs to 70 cm tall, erect. Leaves mostly basal, pet- ioles to 2 1 cm long, subquadrangular, 3 mm wide, spar- ingly pilose; laminae to 60 cm long, 30 cm broad, cor- date, smuatc-pmnatifid. dentate, sparingly pilose, more so on the veins. Inflorescences in large, terminal spikes to 65 cm tall; peduncle quadrangular to 35 cm long, 5 mm thick, sparingly pilose, rachis similar; floral bracts ovate, to 3.5 cm long, 1.5 cm broad, apically acute, spine- tipped, puberulous, margins with 6-8 spreading, spiny teeth to 5 mm long; bracteoles linear, 15-20 mm long, 1.5-2 mm broad, spine-tipped, puberulous; upper calyx segment obovate, 4.5 cm long, 2 cm broad, apically 2-lobed, lobes 25 mm long apically acuminate, lower segment obovate, 3.5 cm long, 1.2 cm broad, apically 2-lobed, lobes 22 mm long, apically acuminate, lateral lobes orbicular, 50 mm in diameter, all segments puber- ulous; corolla whitish, rose or lilac, 1 -lipped, to 4 cm long, the lip 3-lobed, lobes suborbicular, 4 mm long, middle lobe to 1 9 mm broad, lateral lobes 1 4 mm broad, glabrous, tube short, 4 mm long, glabrous with a circle of hairs 2 mm long at point of attachment of stamens; stamens included within upper calyx lobe, filaments sig- moid, to ca. 20 mm long, 2 mm thick, prominent, in- durate, glabrous; anthers 10 mm long, 2 mm broad. Fruits not seen; seeds 4. A common planting in yards and flower gardens of the Meseta Central. This species is a native of Italy. Aphelandra R. Br. REFERENCES— D. C. Wasshausen, The Genus Aphelandra, Smithsonian Contr. Bot. No. 18. 1975. L. A. McDade, Systematics and reproductive bi- ology of the Central American species of the Aphe- landra pulcherrima complex (Acanthaceae). Ann. Missouri Bot. Card. 71: 104-165. 1984. Shrubs or suffrutescent herbs. Leaves opposite; lam- inae usually large, oblong to elliptic, margins toothed, lobed, crenate or entire. Inflorescences of terminal or axillary spikes, spikes often conspicuous with showy bracts and flowers; bracts most often imbricate, certain species bearing dorsal clusters of submarginal glands (ex- tra-floral nectaries) which may be conspicuous, margins of bracts entire, toothed or ciliate; bracteoles various, rudimentary in a few species; flowers with 5-merous ca- lyx, segments separate nearly to base, frequently lanceo- late and striate-veined, posterior segment usually broad- er than others; corolla straight or curved, the limb usually bilabiate, upper lip erect, 2-lobed or entire, lower lip reflexed or spreading, 3-lobed, the middle lobe often larger than the laterals; stamens 4, usually exserted, but usually not beyond the upper lip of the corolla; anthers narrow, 1 -celled, often pilose dorsally and held together at apices by intertwining hairs. Fruits a capsule, usually clavate, 4-seeded. Aphelandra is one of the larger genera of the family and is endemic to tropical America. Species of the genus are often quite showy and yet they are seldom cultivated because of difficulties in cul- tivation. In Costa Rica the genus is represented in a variety of forested habitats from sea level to elevations of 1 500 m, including the dry forests of the Pacific watershed, cloud forests of higher ele- vations, and in the rain forests of the Caribbean watershed. Aphelandra scabra of the Pacific dry forests is probably the most abundant and wide- spread species in the genus. Key to Species of Aphelandra la. Bracts toothed 2a Ib. Bracts entire 8a 2a. Bracts with submarginal glands 3a 2b. Bracts without submarginal glands 4a 3a. Bracts more than 20 mm long; petioles 1 0-30 mm long A. storkii 3b. Bracts less than 1 6 mm long; petioles 0-5 mm long A. scabra BURGER: FLORA COSTARICENSIS 23 4a. Bracts less than 1 5 mm long; corolla less than 2 cm long 5a 4b. Bracts more than 20 mm long; corolla more than 5 cm long 6a 5a. Leaves mostly less than 10 cm long; bracts more than 10 mm long A. seibertii 5b. Leaves more than 10 cm long; bracts less than 8 mm long A. tonduzii 6a. Bracts with 1-2 pairs of teeth above midpoint; peduncle more than 10 mm long A. trident at a 6b. Bracts with spine-tipped, serrate margins; peduncle short, less than 5 mm long 7a 7a. Leaves elliptic to ovate, 3-8.5 cm broad A. aurantiaca var. aurantiaca 7b. Leaves lanceolate, 1.5-2.5 cm broad A. aurantiaca var. stenophylla 8a. Bracts with submarginal glands 9a 8b. Bracts without submarginal glands 1 3a 9a. Submarginal glands consisting of numerous, minute glands appearing as a dull, alveolar spot lOa 9b. Bracts with submarginal glands represented by 1 or more well-defined, elliptical, shiny areas 12a lOa. Petioles short, to ca. 1 cm long; corolla puberulous; capsules clavate . . A. lingua-bovis lOb. Petioles 2-8 cm long; corolla papillose; capsules oblong or ovate 1 la 1 la. Bracts acuminate to acute, 7-9.5 mm long; calyx segments 9-1 1 mm long A. leonardii 1 Ib. Bracts obtuse to acute, 10-1 1 mm long; calyx segments 15-17 mm long A. campanensis 12a. Bracts 6-13 mm long, drying to light or dark brown color; younger stems strigose A. golfodulcensis 1 2b. Bracts 1 6-20 mm long, drying to light orange color (orange-red when fresh); younger stems velutinous A. sinclairiana 13a. Bracts more than 3 cm long; young stems quadrangular, puberulous to glabrous . A. dolichantha 1 3b. Bracts less than 8 mm long; stems terete, pilose A. tonduzii Aphelandra aurantiaca (Scheidw.) Lindl. in Ed- wards, Bot. Reg. 31: pi. 12. 1845. Hemisandra aurantiaca Scheidw., Bull. Acad. Roy. Sci. Bru- xelles9:22. 1842. Figure 1. Suffrutescent herb to shrub, erect, 1 m tall; young steins tetrangular, older stems terete, internodes between leaf- bearing nodes 2-6 cm long, 2-4 mm thick, younger stems glabrous to velutinous. Leaves petiolate; petioles to 1 cm long, glabrous; laminae elliptic to ovate with undulate to entire or sinuate-repand margins, frequently thick- ened, 8-28 cm long, 2.5-12.5 cm broad, apically acu- minate, basally attenuate, occasionally obliquely decur- rent, glabrous except for occasional puberulence along veins, costa and lateral veins prominent, reticulations conspicuous on upper surface and obscure beneath. In- florescences in mostly terminal, quadrifarious spikes to 1 7 cm long, 3 cm broad, excluding the corollas; peduncle short, to 3 mm long, 2 mm thick, puberulous, rachis similar, flattened at the nodes; bracts elliptic or ovate, imbricate, 2.5-3 cm long, 0.75-1 cm broad, apically acute, basally cuneate, puberulous-glandular on both surfaces, reticulations prominent on inner surface, margins ser- rate, serrations tipped with spines to 1.5 mm long; brac- teoles narrowly lanceolate, to 10 mm long, 0.5 mm broad, puberulous on both surfaces, striate-ridged. Flowers with calyx segments 1 cm long, posterior segment oblong, 2.5 mm broad, with a 3-toothed apex, lateral and anterior segments linear-lanceolate, 1 mm broad, all segments puberulous with striate ridges culminating at the base in callus tissue ca. 1 mm thick; corolla red, orange-red, or scarlet, tube to 4 cm long, 3 mm broad, bilabiate, upper lip to 2 cm long and 1 cm broad above base, oblong- ovate, apiculate at apex, forming a partial hood over the anthers, lower lip 3-lobed, elliptic middle lobe 2 cm long, 1.2 cm broad, apically acute, lateral lobes elliptic-ovate, to 1 .2 cm long, 0.75 cm broad, apically acute, puberulous on both surfaces; style to 4.6 cm long, sparingly puber- ulous with ascending hairs, ovary puberulous at apex. Fruits 1 .6 cm long, sparingly puberulous, seeds hispid- ulous. Found in dense forests from Mexico to Bolivia. Aphelandra aurantiaca is recognized by its large, spiny toothed bracts which lack submarginal glands. It is one of the most easily recognized species of the genus and does not closely resemble any of the other Costa Rican species. Aphelandra aurantiaca (Scheidw.) Lindl. var. au- rantiaca The characteristic features of this variety are elliptic to ovate laminae 8-21 cm long, 3-8.5 cm broad with undulate to entire margins that are usually thickened. 24 FIELDIANA: BOTANY This variety is found in dense forests from Mex- ico to Bolivia. In Costa Rica it ranges from the wet evergreen forests of the Caribbean watershed to the moist evergreen forests around the Meseta Central and into Guanacaste to Liberia, at ele- vations near sea level to above 1 500 m. Flowering collections have been made from August through January. Aphelandra aurantiaca (Scheidw.) Lindley var. stenophylla Standley, Publ. Field Columbian Mus., Hot. Ser. 4: 324. 1929; Field Mus. Nat. Hist., Bot. Ser. 10: 59. 1931. Aphelandra re- panda Nees in DC., Prodr. 11: 728. 1847. The characteristic features of this variety are lanceo- late leaves, laminae 20-28 cm long, 1.5-2.5 cm broad, gradually narrowed apically and basally, margins sin- uate-repand. Commonly found in wet forests from Central America to Bolivia. Collected mostly in Costa Rica in the Meseta Central region from an altitude of 1 300 m down to elevations of 700 m near Tilaran, Guanacaste. Aphelandra campanensis Durkee, Ann. Missouri Bot. Gard. 65: 162. 1978. Figure 1. Shrub to 3.5 m tall; internodes between leaf-bearing nodes 2.5-8 cm long, 3.5-7.5 mm thick, tetrangular, sparingly to moderately strigose, lenticels corky ca. 1 mm long. Leaves with strigose petioles to 5 cm long; laminae elliptic, 14-30 cm long, 7-12.5 cm broad, apically acu- minate, basally attenuate, margins entire or undulate, glabrous to sparingly strigose above, strigose beneath, more so on the costa and veins. Inflorescences in ter- minal spikes borne singly or in clusters, to 24 cm long, 1-1.5 cm broad excluding corollas; peduncles short, 5- 13 mm long, sparingly strigose; rachis glabrate; bracts rhombic-ovate, 10-11 mm long, 7-8 mm broad, apically obtuse to acute, apiculate, glabrate and minutely rough- ened with strigose hairs in some, the nerves prominent except toward the apex, with 2 submarginal glandular areas at about the midpoint, each consisting of a compact cluster of numerous, minute, dull brown glands making an elliptic, alveolar spot 1.5-2.0 mm long and 0.5-0.75 mm broad, bract margins subhyaline, entire; bracteoles falcate-lanceolate, 9-10 mm long, 2.5-3.25 mm broad, keeled, striate-nerved, densely hirsute along the keel, marginal area glabrous, subhyaline, sparingly papillose. Flowers with the calyx segments conspicuously longer than the bracts, segments 15-17 mm long, apically ob- tuse, apiculate, striate-nerved, glabrous outside, anterior segments lanceolate, 3.5 mm broad, posterior segment ovate, 7 mm broad; corolla orange-red, bilabiate, densely papillate outside and inside, 5-7 cm long, the tube to 4 cm long, 5 mm broad basally, constricted to 3.5 mm broad just above the apex of the ovary, expanding to 1 1 mm broad at the mouth, the upper lip ovate, ca. 1 9 mm long, 9 mm broad, apically with 2 acuminate lobes ca. 6 mm long, the lower lip spreading at maturity, the mid- dle lobe ovate, to 23 mm long, 1 1 mm broad, apically acuminate, the tip keeled, the lateral lobes triangular, the free portion 1.5 mm long, 2 mm broad, fused to the upper lip for most of their length, the tip rounded; sta- mens extending to the tip of the upper lip, the anthers 9 mm long, 1.5 mm broad, acute at both ends; style and ovary glabrous. Fruits ovate, ca. 20 mm long, 6 mm broad, the tip rounded, the surface shiny and punctic- ulate; seeds suborbicular, 4 mm in diameter, the surface minutely roughened. This species occurs in lowland to mid-elevation ( 1 000 m) rain forests mainly in western provinces bordering on the Caribbean in Panama, but ex- tends into Costa Rica. Only one Costa Rican col- lection, McDade 242 (DUKE), has been observed. Flowering collections have been made from Au- gust through November. Aphelandra campanensis is recognized by its shrubby habit; compact terminal inflorescences with imbricate bracts that lack marginal teeth, are apically obtuse to acute, and have two submarginal glandular areas, each consisting of numerous mi- nute, dull, alveolar-appearing glands; its calyx lobes conspicuously longer than the bracts; and its 5- to 7-cm long corollas that are orange-red and are externally papillate. It may be confused with A. leonardii, which differs in its shorter bracts (7 to 9.5 mm long vs. 10 to 1 1 mm), which are apically acute to acuminate, its shorter calyx segments (9 to 1 1 mm long vs. 1 5 to 17 mm), and its bright red corollas. It may also be confused with A. lin- gua-bovis, which can be distinguished by its cla- vate capsules, bright red bracts, and puberulous corollas. Aphelandra dolichantha Donnell Smith, Bot. Gaz. (Crawfordsville) 27: 438. 1899. Figure 1. Shrubs or suffrutescent herbs ca. 1 m tall; young stems quadrangular, internodes between leaf- bearing nodes 1- 5 cm long, 2.5-4 mm thick, puberulous to glabrous, hairs brownish. Leaves petiolate, petioles 0.5-2 cm long, mi- nutely strigose; laminae mostly elliptic, to 24 cm long, 10.5 cm broad, apically acuminate, basally attenuate, margins entire, glabrous above, glabrous to finely strigose below, when present, hairs more dense on costa and lateral veins, drying to a lighter green than the upper surface, costa and lateral veins prominent. Inflorescences of terminal, sessile spikes, borne singly or occasionally in small clusters, to 13 cm long, 1.5 cm broad excluding corollas; rachis puberulous; bracts closely imbricate at anthesis, spreading at time of fruiting, green when fresh, BURGER: FLORA COSTARICENSIS 25 drying reddish brown, ovate-lanceolate to oblanceolate, to 3.5 cm long, 1 cm broad, apically acuminate to sub- acute, glabrous, margins ciliolate; bracteoles linear, to 8 mm long, 0.5 mm broad, with appressed pilose hairs, 0.5 mm long plus smaller gland-tipped hairs. Flowers 6.5-7.2 cm long; calyx minute, segments linear-subulate, ca. 1.5 mm long, 0.25 mm broad, ciliolate; corolla white, the tube slender, to 5 cm long, nearly cylindrical, basally 2 mm broad, 3.5 mm broad at throat, glandular-puber- ulous, the upper lip erect, oblong, to 1 2 mm long, 6 mm broad, 2-lobed, the lobes 5 mm long, 3 mm broad at base, apically rounded, the lateral lobes oblong to 17 mm long, 6 mm broad, apically rounded, the middle lobe of the lower lip to 22 mm long, 9 mm broad, apically rounded; stamens included, filaments ca. 6 mm long, affixed just below the throat, anthers 2.5 mm long, ba- sally apiculate, adhering by terminal hairs; style includ- ed. Fruits clavate, 17 mm long, 4 mm broad, 3 mm thick, glabrous, minutely punctate, perianth persistent except early deciduous corolla; seeds flattened, ovoid to obliquely ovoid, to 5 mm long, 4 mm broad, 1.5 mm thick dark brown, smooth. Found in lowland wet evergreen forests on the Caribbean side of Costa Rica, in higher wet ev- ergreen forests of Bajo La Hondura to an altitude of 1400 m, and at 750 m near Tilaran in Guana- caste. Flowering collections have been made from April through October. The species is also found in similar habitats of Panama and Colombia. Aphelandra dolichantha is recognized by its long (more than 3 cm), narrow (1 cm) bracts, which lack submarginal glands, and its large (more than 5 cm long), narrow, white flowers. It slightly re- sembles A. storkii, but this species has bracts that are shorter, with toothed margins and submarginal glands, orange flowers, and apparently a much more restricted habitat. Aphelandra golfodulcensis McDade, Ann. Mis- souri Bot. Card. 69: 405. 1982. Figure 2. Shrubs to small trees 1-6 m tall; internodes between leaf-bearing nodes 1-4 cm long, 3-6 mm thick, terete, densely retrorse. Leaves on moderately strigose petioles to 1 cm long; laminae elliptic to oblanceolate, 8.5-30 (45) cm long, 2-8 ( 1 5) cm broad, apically acute to acu- minate, basally attenuate, margins entire or slightly un- dulate, sparsely strigose on the veins above, moderately strigose beneath. Inflorescences borne in numerous ter- minal spikes to 1 5 cm long, 1 cm broad excluding co- rollas; peduncles to 10 cm long, moderately pubescent; rachis glabrous to minutely puberulous; bracts imbricate, rhombic-ovate, apically acute, 6-10 mm long, 4-7 mm broad, green to dull brown-orange, margins ciliolate, with 2 clusters of submarginal glands just below the middle of the dorsal surface, glands ca. 0.75 mm long, 0.5 mm broad, 1-7 glands per cluster; bracteoles narrowly ovate, apically attenuate, 4-6.5 mm long, 2-4 mm broad, green, moderately puberulous. Flowers with calyx segments 6- 9 mm long, apically acute, green, finely striate, minutely puberulous, posterior segment narrowly ovate, 3-4 mm broad, anterior segments broadly lanceolate, 2-2.5 mm broad, lateral pair narrowly lanceolate, ca. 1 .5 mm broad; corolla orange to red, 6.0-7.3 cm long, puberulous, tube 3.8—4.0 cm long, 2-3 mm broad basally, expanding to 6-8 mm broad at the mouth, the upper lip erect, elliptic, 17-19 mm long, 7-11 mm broad, 2-lobed, the lobes triangular, acuminate, 6-10 mm long, the lateral lobes of the lower lip triangular, 3 mm long, 5-7 mm broad, middle lobe broadly lanceolate, spreading, 22-26 mm long, 6-9 mm broad, apically acuminate; stamens exsert- ed to just below the tip of the upper lip and enclosed by it. Fruits clavate, terete, 19-23 mm long, 5-8 mm broad, 5.5-7 mm thick, glabrous; seeds 4, slightly flattened, orbicular, 4-6 mm in diameter. According to McDade this species is found pri- marily in the wet lowlands of the Golfo Dulce region in Puntarenas Province. It extends into the Burica Peninsula of Panama, to mid-elevations above the Golfo Dulce region, and to the north in Alajuela and Guanacaste, where local conditions provide similar moist habitats. Flowering collec- tions have been made from December through March. Aphelandra golfodulcensis is recognized by its imbricate, rhombic-ovate bracts (6 to 10 mm long, 4 to 7 mm broad), which are green or dull brown- orange and minutely puberulent, its minutely pu- berulent corolla, and its glabrous green fruits. It most closely resembles A. sinclairiana, which dif- fers in its larger obovate-rhombic bracts ( 1 6 to 2 1 mm long, 14 to 20 mm broad), which are bright orange, its corollas, which are short velutinous, and its black fruits, which are pubescent. Accord- ing to McDade these two species are closely re- lated, as observed in experimental hybridizations and several collections of putative hybrids from northeastern Costa Rica. She reports further that these hybrids are morphologically intermediate between the two parental species and are sterile. Aphelandra leonardii McDade, Ann. Missouri Bot. Gard. 69: 408. 1982. Figure 2. Shrubs 1-5 m tall; younger stems quadrangular, older stems terete, internodes between leaf-bearing nodes 2-5 cm long, 1.5-2.5 mm thick, strigose, lenticels scattered, oval, ca. 0.5 mm long, corky. Leaves petiolate, petioles ca. 3 cm long, pubescence that of the costa; laminae elliptic to elliptic-obovate, 10-20 (30) cm long, 4-10 cm broad, apically acute to acuminate, basally attenuate, margins shallowly crenate to slightly undulate, glabrous above, sparingly strigose to glabrous below. Inflores- cences in terminal spikes, quadrangular, to 8 cm long, 1 26 FIELDIANA: BOTANY cm broad excluding corollas; peduncle subsessile, rachis glabrous to sparingly puberulous; bracts rhombic-ovate, imbricate, green to bright orange, 7-9.5 mm long, 5-6 mm broad, apically acute to acuminate, tip apiculate, keeled and stria te-nerved, minutely puberulous on upper bracts, lower bracts increasingly strigose mostly over the keel, with submarginal glandular areas just below mid- point consisting of numerous, minute (0.2 mm), often shiny brown to dark brown glands making an elliptic, alveolar spot 1-1.5 mm long, 0.5-1 mm broad, margins ciliolate; bracteoles falcate, keeled, to 8 mm long, 2.5 mm broad, striate-nerved, finely puberulous outside ex- cept wooly keel, glabrous inside. Flowers 6.5-7.5 cm long; calyx segments conspicuously longer than the bracts, 9-1 1 mm long, glabrous, anterior segments oblong, all others lanceolate, apically acute, anterior segments 3 mm broad, lateral segments 2.5 mm broad, posterior segment 5 mm broad, all striate-nerved; corolla bright red, bi- labiate, papillate outside and inside, tube to ca. 5 cm long, 4 mm broad basally, constricted to ca. 2 mm broad just above the ovary, expanding to 9 mm broad at the throat, upper lip erect, ca. 20 mm long, basally 9 mm broad, consisting of 2 acuminate lobes ca. 1 2 mm long, the lower lip spreading at maturity, with 3 lobes, the middle lobe lanceolate, ca. 26 mm long, 8 mm broad, the lateral lobes ca. 6 mm long, acute, connate to the upper lip for most of their length; stamens extending to ca. the length of the upper lip. Fruits oblong, flattened, glabrous, 17.5-19 mm long, ca. 5 mm broad, 3.5-4 mm thick; seeds irregularly orbicular, strongly flattened, 3.5- 6.5 mm in diameter. A shrub of the forest understory, this species is found in lowland and premontane forests in east- ern Panama and in Costa Rica from along the Pacific in Guanacaste to the Meseta Central and the Cordillera Talamanca north of San Isidro to elevations of 1 300 m. Flowering collections have been made in December only. Aphelandra leonardii can be recognized by its calyx, which is conspicously longer than the bracts, its short (7 to 9.5 mm long), entire bracts with submarginal alveolate glands, and its long (ca. 7 cm), bright red corollas, which are externally pa- pillate. It resembles A. campanensis, a largely Pan- amanian species that has been collected in adja- cent areas of Costa Rica. (See the description of that species for a summary of differences.) It also resembles A. lingua-bovis, and these differences are summarized in descriptions of both this species and A. campanensis. Aphelandra lingua-bovis Leonard, Contr. U.S. Natl. Herb. 31: 268. 1953. Figure 2. Shrubs, 1-3.5 m tall; stems terete or subquadrangular toward tips, internodes between leaf-bearing nodes 2-9 cm long, 3-5 mm thick, moderately and minutely stri- gose, the hairs upwardly appressed. Leaves petiolate, pet- ioles to ca. 1 cm long, strigose; laminae oblanceolate to narrowly elliptic, to 38 cm long, 1 2 cm broad, apically acuminate and often curved to one side, basally atten- uate, margins entire or undulate, glabrous to sparingly strigose above, hairs densest along the prominent costa and lateral veins, strigose to sparingly strigose below, lower surface drying to a lighter green than the upper. Inflorescences in terminal spikes, mostly solitary, spikes to 38 cm long, 1.2 cm broad; peduncle to 1 cm long, sericeous; rachis sparingly sericeous; bracts bright red when fresh, drying to reddish brown, rhombic-ovate, 10- 14 mm long, 8-10 mm broad, apically acute to slightly obtuse, glabrous to puberulous with upwardly appressed hairs along the prominent costa, lateral veins also prom- inent, with 2 dull, elliptic, alveolate, submarginal glan- dular areas about midway from the base, to 3 mm long, 1 mm broad, margins ciliolate; bracteoles linear-lanceo- late, to 1 5 mm long, 2 mm broad, apically acuminate, keeled, the keel densely hirsute with ascending hairs ca. 1 mm long. Flowers with the calyx ca. 1 7 mm long, the segments lanceolate, increasingly puberulous toward the apex, striate-nerved, the posterior segments 8 mm broad, the lateral segments 1 .7 mm broad, the anterior segments 2.2 mm broad; corolla bright red, or reddish yellow, 5.5- 6 cm long, puberulous, the tube subcylindric, slightly curved, 3.5 mm broad at the base, 6 mm broad at the throat, the upper lip erect, elliptic, 16 mm long, 7 mm broad, 2-lobed, the lobes apically acute, 4 mm long, 3 mm broad basally, the middle lobe of the lower lip lan- ceolate, 1 8 mm long, 6 mm broad, acute, the lateral lobes 4 mm long, partly coalescent to the upper lip, the free portions triangular, ca. 1.5 mm long, 1 mm broad, api- cally acute; stamens slightly exserted, anthers apiculate at both ends, webby-pilose at tip; ovary densely strigose, hairs ascending. Fruits clavate, 1 8-20 mm long, 4 mm broad, 3.5 mm thick, glabrous except for ascending stri- gose hairs toward the round apex, perianth persistent except the early deciduous corolla; seeds 4, dark brown, deltoid-shaped, ca. 5 mm long, 4 mm broad, margins sparingly tuberculate. Found in mature rain forest of lowland areas in southwestern Costa Rica. Most collections are from the Golfo Dulce area at low elevation, although one collection, Raven 22017 (F), has been made at Wilson's finca south of San Vito (San Vito de Java) at ca. 1300 m elevation. Flowering collec- tions made from June through March. This species is also found in Panama and Colombia. Aphelandra lingua-bovis is recognized by its long (to 38 cm), compact inflorescences with entire, bright red bracts (when fresh) with alveolate sub- marginal glands and large (ca. 6 cm long), pu- berulous, bright red corollas. It resembles A. cam- panensis and A. leonardii, which differ in their smaller inflorescences, papillose corollas, and either oblong or ovate (vs. clavate) capsules. Aphelandra sea bra (Vahl) Sm. in Rees, Cyclopae- dia 39, Aphelandra n. 3( 1 8 1 8), based on Justicia BURGER: FLORA COSTARICENSIS 27 scabra Vahl, Enum. 1: 120. 1804. Aphelandra deppeana Schlecht. & Cham., Linnaea 5: 96. 1830. A. pulcherrima sensu H.B.K., Nov. gen. sp. 2: 236. 1817. A. pectinata Willd. ex Nees, DC., Prodr. 1 1: 298. 1847. A. haenkeana Nees, DC., Prodr. 11: 298. 1847. Figure 1. Shrubs 1-3.5 m tall; young steins subquadrangular to terete, glabrous to velutinous, leafy inter nodes 3-10 cm long, 2.5 mm thick, lenticels oval to linear, mostly less than 1 mm long. Leaves petiolate, petioles sessile to 5 mm long, strigose; laminae elliptic to 25 cm long, 8 cm broad, apically acuminate, basal ly attenuate, margins en- tire to undulate, glabrous to scabridulous above, strigose to velutinous beneath, especially on costa and lateral veins. Inflorescences in dense terminal spikes to 1 9 cm long, 1 cm broad excluding corollas; peduncle subsessile; rachis pilose; bracts imbricate to 15 mm long, 6 mm broad, apically acuminate, basally cuneate, strigose out- side, puberulous inside, with 2 clusters of submarginal, mid-dorsal glandular areas with 1-10 ovular glands per area, each gland ca. 0.5 mm long, outside of bract rather strongly 5-nerved, with 4-8 marginal teeth to 1.5 mm long, margins ciliate; bracteoles lanceolate, to 6 mm long, 1.5 mm broad, keeled, velutinous along the keel. Flowers to 3 cm long, calyx segments lanceolate, to ca. 8 mm long, anterior segments ca. 1.5 mm broad, lateral seg- ments ca. 1 mm broad, posterior segment ca. 3 mm broad, all segments puberulous toward apex; corolla pink, purplish red, scarlet or red, bilabiate, to 3 cm long, grad- ually expanding to 4 mm broad at the mouth, constricted to 2 mm broad just above the ovary, puberulous outside, glabrous inside, lips ovate, upper lip 2-lobed, to 1 cm long, each lobe acuminate, 3 mm long, lower lip slightly longer, acuminate at apex; stamens with hispid filaments. exserted. Fruits elliptic, to 1 5 mm long, 6 mm broad, 3 mm thick, apex acute, glabrous to sparingly puberulous; seeds 2 per locule, flattened, irregular in shape, ca. 3 mm broad, 1 mm thick, surface minutely roughened, dark brown. This is a common and wide-ranging plant in shaded areas within mature tropical dry forests or old secondary forests, generally at lower elevations of the western coastal areas in Costa Rica, but also found in the Meseta Central region at elevations as high as 1 100 m. This species grows from south- ern Mexico to northern South America. Flowering observed year round. Aphelandra scabra is recognized by its pale green (when fresh) toothed bracts with two clusters of submarginal glands, smallish bracts (less than 1 6 mm long), short, reddish corollas (to 3 cm long), and its subsessile leaves. It bears a slight resem- blance to A. storkii, which is easily distinguishable, with its longer bracts, longer orange corolla, and much longer petioles. Aphelandra seibertii Leonard, Ann. Missouri Bot. Card. 24: 205. 1937. Figure 2. Herbs to 19 cm tall; stems quadrangular, internodes between leaf-bearing nodes 0.5-1 cm long, 2-2.5 mm thick, pilose hairs to 1 mm long, lower stem rooting at the nodes. Leaves petiolate, petioles to 1.6 cm long, densely pilose; laminae elliptic, 4.5-9.5 cm long, 1.5- 4.0 cm broad, apically acute, basally acute, margins en- tire to slightly undulate, pilose hairs to 1 mm long above and beneath, very dense along costa and lateral veins beneath. Inflorescences in 1-3 terminal spikes to 13 cm long, 1 cm broad excluding corollas; peduncles 3-5 cm long, densely yellowish pilose; rachisdensely pilose; bracts closely imbricate, spreading at maturity, rhombic, to 14 mm long, 6 mm broad, tips often purplish, margins above midpoint dentate with 2-3 erect teeth per side, each ca. 1 mm long; bracteoles lanceolate, to 9.5 mm long, 2 mm broad, keeled, translucent, pilose along the keel. Flowers 15-20 mm long, falling away at maturity when bracts are expanded; calyx segments lanceolate, subequal, ca. 5 mm long, translucent, with small glandular hairs plus pilose hairs, posterior segment 1.4 mm broad, lateral segments 0.5 mm broad, anterior segments 0.7 mm broad; corolla yellow at base, lobes lavender-tipped, puberu- lous-glandular outside, glabrous inside, bilabiate, tube funnelform, 1 1 mm long, 2 mm broad at base expanding to 4 mm broad at mouth, upper lip 6.5 mm long, 5.5 mm broad, apex with 2 rounded lobes 2 mm long, lower lip 3-lobed, middle lobe retuse, 8 mm long, 5 mm broad, lateral lobes apically rounded, 6 mm long, 2 mm broad; stamens slightly exserted, filaments pilose. Fruits oblan- ceolate, ca. 8 mm long, 3 mm broad, 1.5 mm thick, reddish brown; seeds 4, subtri angular. 3 mm long, 0.7 mm thick, puberulent. Found in rain forests in deep shade in Costa Rica and Panama. Found in the Caribbean slope at lower elevations in Costa Rica. Flowers through most of the year. Aphelandra seibertii is recognized by its low her- baceous habit, pilose stems and leaves, small leaves (mostly less than 1 0 cm long), toothed bracts, and corollas with yellow bases and purplish tips. This may be confused with Justicia chamaephyton, which can be distinguished from A. seibertii by its basally obtuse or rounded leaves, two stamens, less compact spikes, and smaller entire bracts. Aphelandra sinclairiana Nees in Benth., Bot. voy. Sulphur 146. pi. 47. 1844. Figure 2. Shrubs or small trees to 6 m tall; younger stems terete to subquadrangular at the apex, internodes between leaf- bearing nodes 1—4 cm long, 2-8 mm thick, velutinous. Leaves petiolate, petioles 1-2 cm long, pilose; laminae narrowly elliptic to near-oblanceolate, to 30 cm long, 8.5 cm broad, apically acuminate, basally attenuate, margins entire to shallowly crenate, sparingly pilose above except at costa, pilose below. Inflorescences borne in terminal and subterminal spikes in clusters of as many as 1 0 spikes, spikes to 15 cm long, 2 cm broad excluding corollas; peduncles subquadrangular, to 5.5 cm long, rachis ve- lutinous; bracts orange-red, densely imbricate, obovate- 28 FIELDIANA: BOTANY rhombic, 16-20 mm long, 14-20 mm broad, lowermost bracts apically acuminate, upper bracts often obtuse, 5- 6 rather prominent nerves with branching toward apex of bract, puberulous to velutinous dorsally, puberulous ventrally, with 2 clusters of submarginal elliptic glands at about the middle of the dorsal surface, glands ca. 0.5 mm long, 0.25-0.4 mm broad, 2-10 glands per cluster, margins entire; bracteoles lanceolate to ovate-lanceolate, 6.5-7.5 mm long, 3-3.5 mm broad, striate, subcarinate, puberulous to tomentose outside, puberulous inside. Flowers 5.5-6.5 cm long; calyx segments lanceolate, the posterior segment 7-9 mm long, 3 mm broad, lateral and anterior segments 7-9 mm long, 1.5-2 mm broad, puberulous outside, glabrous inside, striate-nerved; co- rolla purplish red, bilabiate, short-velutinous outside, tube funnelform, 4.5-5 cm long, 3.5 mm broad at base, 6.5 mm at mouth, upper lip erect, to 1 8 mm long, 9 mm broad, 2-lobed, the lobes acuminate, keeled apically, to 7.5 mm long, the lower lip spreading, middle lobe lan- ceolate, keeled, to 25 mm long, 8 mm broad, lateral lobes triangular, connate to upper lip for most of their length, acute, free portion to 1.3 mm long, 1.5 mm broad; sta- mens exserted to just below the tip of the upper lip and enclosed by it. Fruits clavate, black, to 1.8 cm long, 6 mm broad, 3 mm broad at base, puberulent, apically obtuse; seeds 4, puberulent, flattened, ca. 3.5 mm in diameter. According to McDade this species grows mainly in gaps in lowland rain forests extending to pre- montane forest and is also a successful colonizer in more extensively disturbed areas. In Costa Rica it has been collected mostly from such habitats in Limon Province, with a few older collections from Cartago and Heredia provinces. Flowering collec- tions have been made from September through April. Aphelandra sinclairiana is recognized by its large, obovate-rhombic bracts ( 1 6 to 20 mm long), which are orange-red and have two clusters of shiny, sub- marginal glands, its large (ca. 6 cm long), purplish red corollas, and its often relatively tall height (to 6 m). It is easily confused with A. golfodulcensis. (See the description of that species for a discussion of these differences.) Aphelandra storkii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1197. 1938. Figure 2. Shrub to ca. 2 m tall; young stems quadrangular, terete shortly below terminus, internodes between leaf-bearing nodes 2-4.5 cm long, 2-3.5 mm thick, pilose toward the terminus. Leaves petiolate, petioles 1-3 cm long, hirsute to suhglahrous; laminae elliptic to elliptic-obovate, to 44.5 cm long, 1 3.5 cm broad, apically acuminate, basally attenuate, margins serrulate to undulate, sparingly stri- gose above, densely strigose below, the veins prominent. Inflorescences in terminal spikes to 18 cm long, 2.5 cm broad excluding corollas; peduncles quadrangular, to 2.5 cm long, hirsute, rachis quadrangular, hirtellous; bracts oblong-ovate, to 25 mm long, 1 3 mm broad, pilosulous, with 2 clusters of 1 5-20 submarginal elliptic glands just below midpoint of dorsal surface, glands ca. 0.5 mm long, 0.2 mm broad, margins ciliolate, with 2-3 pairs of teeth to 0.9 mm long near midpoint, one of which is frequently larger than the others; bracteoles lanceolate, to 10 mm long, 2 mm broad, apically acuminate, striate- nerved, pilosulous. Flowers to 7 cm long; calyx segments subequal, 12-13 mm long, posterior segment 4 mm broad, lateral and anterior segments 2 mm broad, all apically attenuate, pilosulous, striate-nerved; corolla orange, pi- lose, the tube funnelform, 3 mm broad at base, 7 mm broad at throat, upper lip erect, to 20 mm long, 1 2 mm broad, apically 2-lobed, lobes 1 1 mm long, apically keeled and attenuate, the middle lobe of the lower lip lanceolate, to 22 mm long, 9 mm broad, apically attenuate and curved back, lateral lobes triangular, connate to upper lip for most of their length, acute, free portion to 0.8 mm long, 0.9 mm broad, stamens exserted to just below the tip of the upper lip and enclosed by it; anthers to 8 mm long, 1 mm broad, apically adherent, basally acute; filaments glabrous. Fruits clavate, 25-28 mm long, 7-8 mm broad, 4-5 mm thick, glabrous; seeds 4, ovate, ca. 6 mm long, 5 mm broad, glabrous. Found in lowland rain forest. Only one collec- tion, Opler 988 (F, NY, MO) from La Selva near Puerto Viejo of Heredia Province, has been seen other than the type collection made at Livingston along the Rio Reventazon in Limon Province. Flowering collections have been made in July and August. Aphelandra storkii is recognized by its long (more than 2 cm) gland-bearing, toothed (two to three pairs) bracts, its long orange corolla (7 cm), and its relatively large, petiolate leaves. It bears a slight resemblance to A. scabra, which has shorter bracts, corollas, and leaves; its leaves are subsessile and its corollas are reddish, but not orange. Aphelandra tonduzii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1 198. 1938. Figure 3. Herbs to 16 cm tall, stems terete, pilose, internodes between leaf-bearing nodes 3-30 mm long, 2-4 mm thick. Leaves on pilose petioles 1-3 cm long; laminae elliptic to 18 cm long, 5.5 cm broad, apically acute to obtuse, basally attenuate, margins entire, pilose on both surfaces, more so on costa and lateral veins. Inflorescences of loose axillary spikes to 20 cm long, 0.5-1 cm broad excluding corollas; peduncle subquadrangular, pilose, plus occa- sional glandular hairs, rachis subquadrangular, pilose with numerous glandular hairs; bracts lanceolate, slightly keeled, 6-8 mm long, 1.25-1.5 mm broad, puberulous with occasional glandular hairs, margins ciliolate, often with a pair of minute, marginal teeth, bracts not imbri- cate at anthesis; bracteoles subulate, ca. 4.5 mm long, 0.6 mm broad, puberulous with occasional glandular hairs. Flowers to 1 3 mm long; calyx segments to 5 mm BURGER: FLORA COSTARICENSIS 29 long, posterior segment lanceolate, 1 mm broad, lateral and anterior segments subulate, 0.5 mm broad, all striate- nerved, puberulous toward tip; corolla white in bud, be- coming purplish with age, bilabiate, tube to 8 mm long, 1 mm broad at base, 1.5 mm broad at mouth, glabrous outside, puberulous within, the 2 lobes of the upper lip obovate, 2.5 mm long, 1 mm broad, apically retuse, middle lobe of the lower lip rhomboid-ovate, 5 mm long, 5 mm broad, apically retuse, lateral lobes obovate, 4 mm long, 3 mm broad, apically retuse, minutely puberulous inside; stamens included, filaments sparingly puberu- lous, anthers bearing a tuft of hairs apically. Fruits cla- vate, ca. 8 mm long, 3 mm broad, glabrous; seeds 4, obovate, flattened, 3 mm long, 2.5 mm broad, 1 mm thick, surface hispidulous. red, puberulous, the tube 4 mm broad at base, narrowing to 2 mm just above the ovary, gradually enlarging to 7 mm broad at the mouth, the upper lip erect, elliptic, to 25 mm long, 8 mm broad, apically keeled, lower lip 3-lobed, middle lobe ovate, to 25 mm long, 17 mm broad, apically obtuse, lateral lobes oblong-obovate, to 15 mm long, 6 mm broad, apically rounded; stamens exserted to just below tips of upper lip, anthers lemon yellow, 6.5 mm long, 1 mm broad, apically acuminate with pilose hairs, basally acute; filaments sparingly pu- berulous. Fruits ovate, ca. 15 mm long, 6.5 mm broad, apically apiculate, glabrous, shiny, perianth except co- rolla persistent; seeds orbicular, ca. 4 mm broad, surface covered with tubercles. Found in cloud forest at and above 1000 m in Costa Rica and adjacent Panama. It has been col- lected over a wider range in Costa Rica, where most collections are from the Cordillera Central, and others are from the vicinity of San Isidro. Flowering collections have been made from Oc- tober through February. Aphelandra tonduzii is recognized by its small size (to 16 cm tall), its small (less than 8 mm), toothed, eglandular bracts, and its small flowers (to 1 3 mm long). Superficially, it can be confused with some forms of Elytraria imbricata, which differs in its smaller, entire, and more imbricate bracts, its smaller flowers (to 8 mm long), and its slenderly conical capsules, which lack hooklike fu- nicles. Aphelandra tonduzii also resembles A. ar- noldii of Colombia and Panama, which differs with its eglandular rachis and red flowers. Found in cloud forests and forest clearings from the San Jose area to Monteverde in Guanacaste at elevations of 850-1600 m. Flowering collections have been made from September through Decem- ber. Aphelandra tridentata is recognized by its long (2.5 to 3 cm), eglandular bracts with one or two pairs of teeth and its long (5 to 6.5 cm), bright red corollas; it is found at higher elevations (above 850 m). Older inflorescences lacking flowers could be confused with A. dolichantha, which has less compact inflorescences and also differs in its lack of toothed bracts and its longer (6.5 to 7.2 cm), white corollas; it occurs between sea level and 1000m. Asystasia Blume Aphelandra tridentata Hemsl., Biol. cent.-amer., Hot. 2: 513. 1882. Figure 3. Herb to subshrub. erect, to 2 m tall; young stems qua- drangular, internodes between leafy nodes 1-6 cm long, 2-5 mm thick, pilose to wooly. Leaves petiolate, petioles to 3.0 cm long, pilose to strigose; laminae elliptic to elliptic-oblong, 10-18 cm long, 2.5-3 cm broad, apically acuminate, basally attenuate, margins entire to undulate, strigose to glabrous above, strigose below, upper surface drying to pale green, lower surface to dark green. Inflo- rescences in solitary, dense terminal spikes to 1 1 cm long, 3 cm broad excluding corollas; peduncle quadrangular, to 1 3 mm long, wooly, rachis quadrangular, wooly; bracts oblong-lanceolate, green, 3.5-4 cm long, 0.5-1.5 cm broad, sericeous, margins entire with 1-2 pairs of teeth above the midpoint, one conspicuously larger than the rest, to 3.5 mm long; bracteoles linear, to 13 mm long, 1.3 mm broad, apically acuminate, striate-nerved, se- riceous. Flowers 5-6.5 cm long; calyx segments lanceo- late, to 1 1 mm long, puberulous, striate-nerved, poste- rior segment 3.5 mm broad, lateral segments 1.5 mm broad, anterior segments 2.5 mm broad; corolla bright Herbs or shrubs. Leaves entire, petiolate, with abun- dant cystoliths when mature. Inflorescences of secund spikes or racemes, these loose or compact, simple or branched. Flowers solitary on short pedicels, subtended by 2 small bracts and 2 bracteoles of ca. equal size; calyx 5-merous, segments linear or lanceolate; corolla white, blue, purple, rose, or yellow, tube short to funnelform, 5 lobes, subequal; stamens 4, didynamous; anthers ob- long with 2 parallel cells, calcarate or muticous at base; ovary 4-ovulate, pubescent; stigma minutely 2-parted or subcapitate; capsules elliptic, 2-4 seeded; seeds com- pressed, orbicular or irregularly angled, glabrous. This genus is not native to the Neotropics. Ac- cording to Leonard (1951) about 70 species have been described which are mostly from tropical Af- rica, Asia, and the East Indies. Asystasia resembles the genus Dyschoriste with which it shares most characteristics including the few pollen grains that have been examined. They differ in inflorescences, with Dyschoriste having 30 FIELDIANA: BOTANY flowers borne in cymose fascicles or singly at the axils. More than 1 00 species of this genus have been described, and nearly all of them occur in the Old World Tropics. Asystasia gangetica (L.) T. Anders in Thwaites, Enumeratio plantarum zeylaniae 235. 1859- 1864. Justicia gangetica L., Amoen. acad. 4: 299. 1759. Figure 3. Barleria micans Nees in Benth., Bot. voy. Sulphur 146. 1844. B. discolor Nees in Benth., loc. cit. Figure 3. Herbs, ascending, to ca. 30 cm tall; stems quadran- gular, ink-modes between leaf-bearing nodes 4.5-7.5 cm long, 1-2 mm thick, pilose. Leaves petiolate, petioles to 1 1 mm long, sparingly strigose; laminae ovate to deltoid, 3.5-5 cm long, 2-2.5 cm broad, apically acuminate, ba- sally obtuse to truncate, margins crenulate to entire, subglabrous to sparingly pilose especially on the veins above and below, numerous cystoliths visible only on the upper surface. Inflorescences in terminal, mostly se- cund racemes, to 1 3.5 cm long; flowers on short pedicels to 2 mm long; 4 bracteoles in 2 series subtending each flower, lanceolate, to 2 mm long, pilose. Flowers to 4 cm long; calyx and pedicel pilose with numerous glan- dular hairs, calyx lobes lanceolate, 5-7 mm long; corolla yellow with purplish throat, funnelform, 3.5 mm broad at base, 10 mm broad at throat, lobes semiorbiculate, ca. 10 mm broad with numerous minute glandular hairs outside, glabrous within, stamens included; anther cells each mucronulate apically and basally. 3 mm long, 1 mm broad; ovary 3.5 mm long, style 21 mm long, ve- lutinous, stigma 2-lohed. Fruit oblong, to 1 3 mm long, 2 mm broad; perianth persistent except early deciduous corolla; with 2-4 flat, oval seeds ca. 1 mm long, 0.75 mm broad, oblique at point of attachment, edged with dense mucilaginous hairs when wet. Found in cultivation or as an escape in Costa Rica and much of tropical America. Native to tropical Africa, Asia, and Malaya. In Costa Rica and Panama, it is found from near sea level to ca. 1000 m altitude. Flowering collections have been made in March, April, and May. Asystasia gangetica is recognized by its one-sid- ed racemes with large (to 4 cm long), funnelform corollas, which are yellow with purplish throats, and its minute bracteoles (2 mm long). Barleria L. Shrubs or herbs, erect, sometimes spiny. Leaves op- posite, petiolate to near sessile; laminae entire, cystoliths generally prominent and dense on both surfaces. Flowers axillary or borne in dense terminal bracted spikes; calyx deeply 4-lobed, lobes unequal; corolla 5-merous, the broad, spreading lobes mostly subequal; stamens 4, di- dynamous, adnate to base of corolla tube, generally all stamens perfect, but American species with only 2 per- fect stamens plus 2 staminodes. Fruits ovate to oblong, flattened, with 2 seeds per locule; seeds flattened, ovate to suborbicular. Herbs with erect stems to 75 cm tall; internodes be- tween leaf-bearing nodes 1.6-6.2 cm long, 1.5-4 mm thick, stems strigose, hairs appressed, to 1 mm long, cystoliths prominent and dense, 0.3 mm long. Leaves petiolate, petioles to 1.5 cm long, strigose; laminae lan- ceolate to oblanceolate, 5.5-23 cm long, 1 .5-5.2 cm broad, apically acuminate, basally attenuate, margins entire, sparsely strigose above with cystoliths, strigose below, cystoliths few, trichomes densest on veins and margins, bluish along costa upon drying. Inflorescences in dense terminal spikes to 7 cm long, 3 cm broad excluding corollas, subsessile; bracts ovate, to 3.5 cm long, 1.5 cm broad, apically acuminate, basally acute, chartaceous, costa prominent, sericeous inside, strigose outside plus occasional glandular hairs, midvein and base of each side velutinous, often bluish upon drying, margins of spiny hairs to 2 mm long; bracteoles oblanceolate, to 2 cm long, sericeous with simple and glandular hairs, margins ciliate. Flowers sessile, to 4.5 cm long; calyx segments unequal, anterior segment elliptic, to 2.5 cm long, 1 cm broad, apically bidentate, margin with spines to 2 mm long, lateral segments lanceolate, to 2 cm long, 2 mm broad, apically aristate, sericeous, posterior segment slightly longer and narrower than the anterior segment, apically acuminate, margin with spines to 2 mm long; corolla yellow, drying purple, bilabiate, the tube to 3 cm long, slender, 4 mm broad at base, lobes subequal, ob- ovate, ca. 1.5 cm long, 6 mm broad, apically rounded, upper lip of 1 lobe, lower lip of 4 lobes; perfect stamens 2, exserted, anthers 2 -celled, 5 mm long, filaments gla- brous, to 4 cm long, imperfect stamens reduced to stami- nodes ca. 0.2 mm long, staminodes with a centrally lo- cated spur; ovary pandurate, ca. 5 mm long, style ca. 4.2-4.5 cm long, glabrous, stigma cylindrical, to 1 mm long. Fruits elliptic to 2 cm long, 1.6 cm broad, 2 mm thick, apically acuminate, dark purple-brown when dry, the 4 seeds irregularly ovate, ca. 5 mm long, 0.3 mm thick, velutinous. Found in dry forest regions of all provinces in the Pacific watershed from near sea level to about 600 m of the western Meseta Central. Flowering collections made from December to April. This species ranges from west-central Mexico to Co- lombia. Barleria micans is recognized by its dense ter- minal spikes with large, conspicuous, overlapping bracts with spiny margins, these often turning dark bluish upon drying, and its conspicuous yellow (drying to purple), subequal flowers to 4.5 cm long. This species is slightly similar to Aphelandra au- BURGER: FLORA COSTARICENSIS rantiaca, but close examination will show all Aphelandra species to have four fertile stamens instead of two and bilabiate flowers. Blechum P. Browne Perennial herbs or shrubs. Leaves petiolate; laminae lanceolate to ovate, entire, undulate, crenate, or serru- late, containing numerous cystoliths on both surfaces. Inflorescences in dense terminal, quadrifarious spikes. Flowers 5-merous, calyx segments subequal, linear-su- bulate; corolla white, blue, lavender or purplish, fun- nelform with narrow tube, lobes subequal, bracts 1 per flower, imbricate, ovate-oblong to ovate-deltoid; brac- teoles 2, linear, oblanceolate or lanceolate; stamens 4, didynamous, cells of anthers parallel and equal. Fruits of elliptic capsules, flattened, dissepiment breaks away from capsule wall at maturity, seeds few to 16, reddish brown, flattened, ca. 2 mm in diameter, edges with a band of sticky, dense, mucilaginous hairs when wet, gla- brous when dry. Native to tropical America, although species such as B. brownei are reported from other tropical areas as well. Leonard (195 1) states that this genus con- sists of about six tenable species and that there is a close relationship between Blechum and Ruellia, but Blechum can be recognized by its dense, four- sided spikes and very small flowers. Key to Species of Blechum la. Corolla small, equally or barely exceeding the bracts; bracts acute at apex B. brownei Ib. Corolla relatively large, twice as long as the bracts or longer; bracts acuminate at apex B. costaricense Blechum brownei Juss., Ann. Mus. Natl. Hist. Nat. 9: 270. 1807. 1918. Ruellia blechum L., Syst. nat. 10, 2: 1120. 1759. Barleria pyramidata Lam., Encycl. 1: 380. 1783. Blechum pyrami- datum (Lam.) Urb. in Repert. Spec. Nov. Regni Veg. 15: 323. 1918. Blechum brownei f. puber- ulum Leonard, J. Wash. Acad. Sci. 32: 942. Fig- ure 3. Herbs with erect to declining stems to 50 cm tall; younger branches quadrangular, appressed-puberulent, internodes between leaf-bearing nodes 1 .2-7 cm long, 1- 2 mm thick. Leaves petiolate, petioles to 14 mm long, puberulent; laminae ovate to ovate-lanceolate, 2-6.5 cm long, 1-3 cm broad, apically acute, basally cuneate to obtuse, margins crenate to entire, both surfaces sparsely pilose to glabrous, numerous cystoliths. Inflorescences in terminal spikes, 4-sided, 1.5-7 cm long, peduncles to 1.5 cm long, appressed-puberulent; bracts ovate, 7.5-14 mm long, 6-1 1 mm broad, apically acute, basally round- ed, loosely strigose and densely puberulous, margins cil- iate; bracteoles lanceolate, 5 mm long, 1 mm broad, ciliate; calyx segments 3 mm long, 0.5 mm broad, cil- iolate; corolla white turning pale pink and lavender with age, 10-15 mm long, slightly longer than the subtending bract. Fruit obovate, ca. 6 mm long, 3 mm broad, 1 mm thick, apically acuminate, puberulous, perianth persis- tent except the early deciduous corolla; seeds 16, sub- orbicular, 1 mm in diameter. Found as a weed in disturbed areas along riv- erbanks, roadsides, lawns, and pastures; flowering year-round, it occurs in all parts of Costa Rica below 1400 m elevation. Ranging from eastern and west-central Mexico through Central and South America to the West Indies, Guam, Taiwan, and the Phillipine and Caroline islands. Blechum brownei is recognized by its four-sided, terminal spikes, its apically acute bracts, and its small flowers, which barely extend beyond the bracts. Closely resembling B. costaricense, it can be distinguished from it by its shorter flowers and acute bracts; it usually grows in a more disturbed, open habitat. Commonly called Sornia, it is used as a folk remedy in Costa Rica and Panama in the form of a tea to relieve amoebic and probably other types of dysentery. Blechum costaricense Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 168. 1855. Blechum dariense Lindau, Repert. Spec. Nov. Regni Veg. 12: 423. 1913. Figure 3. Herbs to suflrutescent at the base in older forms, to l.lm tall; young stems quadrangular, pilose, internodes between leaf-bearing nodes 2.2-6.5 cm long, 1-2 mm thick. Leaves with petioles to 2 cm long, pilose; laminae ovate to ovate-lanceolate, 2-11 cm long, 1-5 cm broad, apically acuminate, basally attenuate, margins undulate or serrulate, both surfaces pilose to sparingly pilose, nu- merous cystoliths visible on both surfaces. Inflorescences in terminal spikes, 4-sided, to 5.5 cm long, peduncles to 2 mm long, pilose; bracts ovate-deltoid, apically cuspi- date or acuminate, basally rounded, to 2 cm long, 1.5 32 F1ELDIANA: BOTANY cm broad, sparingly pilose outside and inside, margins ciliate; bracteoles linear to oblanceolate, 7-10 mm long, 0.5-1 mm broad, pilose margins, ciliate; calyx segments equal, ca. 5 mm long, 0.75 mm broad at base, ciliate; corolla 2-2.5 cm long, pale lavender to purple, extending well beyond the subtending bract. Fruit elliptic, ca. 8 mm long, 5 mm thick, apically acuminate, perianth per- sistent except the soon deciduous corolla; seeds subor- bicular, ca. 2 mm in diameter. Found in Costa Rica in rain forests of the Ca- ribbean watershed area, the Pacific slopes of Valle del General area to the Cordillera Tilaran, at el- evations of 200 to 1000 m. This species is also found rather commonly in adjacent areas of Pan- ama to sea level and in lowland areas in the Canal Zone. Flowering collections have been made from September through March. Blechum costaricense is recognized by its four- sided, terminal spikes, its broad, acuminate bracts, and its corollas, which are twice as long as the bracts. Closely resembling Blechum brownei, it can be distinguished by its larger flowers and acumi- nate bracts. Bravaisia DC. Shrubs or trees. Leaves petiolate; laminae ovate to elliptic, apically acute to acuminate, margins entire to slightly undulate. Inflorescences terminal in compact, leafy panicles. Flowers subtended by a pair of bracts; calyx 5-merous, subcoriaceous; corolla white, yellow, or purplish with a short tube and campanulate throat, 5 lobes, subequal; stamens 4, anther cells 2, contiguous, spurred at the base. Fruit a capsule with 4-8 seeds. A genus of the Neotropics, Bravaisia has five species variously reported from Mexico, Central America, Colombia, Venezuela, Peru, and the West Indies. Bravaisia integerrima (Spreng.) Standley, Contr. U.S. Natl. Herb. 23: 1335. 1926. Amasonia in- tegerrima Spreng., Syst. veg. ed. 16. 2: 765. 1825. OnchyacanthusspeciosusOersl., Vidensk. Med- del. Dansk Naturhist. Foren. Kj0benhavn 1854: 131. 1855. Figure 4. Tree to 20 m tall; branchlets quadrangular, internodes between leaf-bearing nodes 2.5-7 cm long, 2.5-5 mm thick, glabrous to tomentose on youngest portions. Leaves with petioles to 4 cm long, pubescence of petioles like that of the stem; laminae ovate to ovate-elliptic, 3.5- 28.5 cm long, 1-1 1.8 cm broad, apically acute or acu- minate, hasally acute or oblique, margins entire to slight- ly undulate, subcoriaceous, surfaces glabrous, sparingly pilose at costa and lateral veins. Inflorescences in leafy, compact, terminal panicles, to 13 cm long, 25 cm wide, peduncles quadrangular. Flowers subtended , - oval bracteoles which are ca. 2 mm long, apically obtuse, puberulent and ciliolate; calyx segments subequal, spat- ulate, 5-6 mm long, 2-3 mm broad, puberulous and ciliolate; corolla white or yellow with a purple spot in the throat, 1 .5-2.0 cm long, glabrous, tube campanulate, ca. 8 mm broad at the mouth, the limb ca. 2 cm across, lobes rounded or emarginate, ca. 10 mm long, 6-8 mm broad. Fruits clavate, flattened, 10-12 mm long, 4-5 mm broad, 3 mm thick, glabrous, perianth persistent; seeds 8, orbicular, 3.5 mm in diameter. Plants of evergreen and deciduous forest for- mations, ranging from 100 to 600 m in the Pacific coastal regions to the Meseta Central and the Llan- ura de San Carlos. Flowering collections have been made from December through May. The species ranges from Peru, Colombia, Venezuela, and north to Oaxaca and Tamaulipas, Mexico. Bravaisia integerrima is exceptional among the Acanthaceae for its large size. In addition to its large size, it is recognized by its flowers with four stamens having anthers with basal spurs. It also has eight-seeded capsules. It most closely resem- bles another woody species, Trichanthera gigan- tea, which has anthers with rounded bases and four-seeded capsules. Commonly called palo de agua in Costa Rica, it has a light-colored wood that is suitable for packing boxes and food pails. Numerous aerial roots similar to mangroves are often produced at the bases of the trunks (Leonard, 1951). Buceragenia Green man Herbs; stems erect. Leaves petiolate; laminae usually ovate or elliptic. Inflorescences spicate or spicate pani- cles, the flowers solitary or in clusters; bracts and brac- teoles minute; calyx 5-merous, segments linear, sub- equal; corolla tubular, shallowly bilabiate, posterior lip in-curved, emarginate or 2-lobed. anterior lip erect or slightly spreading, 3-lobed; stamens 2, inserted at middle of corolla tube, included; anthers usually 1 -celled, the lower cell aborted or vestigial and sterile, if fertile, always considerably smaller; staminodes 2; fruits a clavate cap- sule, stipe narrow; seeds 4, flattened, suborbicular. This is a genus of four species which extend from Mexico to Costa Rica. BURGER: FLORA COSTARICENSIS 33 Buceragenia glandulosa Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1205. 1938. Figure 4. Herbs to 1.5 m tall, stem erect, branching, subqua- drangular, glabrous to sparingly puberulous; internodes between leaf-bearing nodes 1.5-4 cm long, 2.5-3 mm thick. Leaves with petioles to 3 cm long, those subtending inflorescences subsessile, pubescence like that of the stem; laminae ovate to ovate-oblong, 5.5-18 cm long, 2-7.5 cm broad, apically acuminate, basally attenuate except the leaves immediately subtending the inflorescence which are basally rounded to cordate, margins entire, costa often reddish purple, glabrous. Inflorescences in terminal and axillary spikes or spicate panicles to 20 cm long, flowers sessile or nearly so, borne in fascicles of 2- 4 or occasionally singly; bracts subulate, 1.5 mm long, 0.5 mm broad, glabrous, bracteoles similar, but smaller; rachis glandular-puberulous; calyx 5-merous, 3 mm long, lobes subulate, 2 mm long, 0.5 mm broad at base, gla- brous; corolla small, tubular, glabrous, 3-3.5 mm long, 1 mm broad, lobes 5, more or less equal, suborbicular, 0.5 mm long, 0.25 mm broad, stamens 2, included, 1 mm long, anthers usually 1 -celled, other cell usually aborted or vestigial and sterile, if fertile, always much smaller, 1 or both cells mucronate. Fruits clavate, 1 3- 19 mm long, glabrous, stipe 6.5-10 mm long, 1.5 mm broad, expanded portion 6.5-9 mm, 2.5 mm broad, seeds flat, oval, 3 mm long, 2 mm broad, roughened. Found in cloud forests at 1200 to 1800 m, this species is reported from Mexico, Guatemala, Nic- aragua, and Costa Rica. Only three collections have been observed from Costa Rica, all from Cartage Province: Holway 400 (us, holotype), Standley 33686 (us), and Standley 35503 (us). All collec- tions were made in January or February, and all were in both fruit and flower. Buceragenia glandulosa is recognized by its her- baceous habit with terminal and axillary spikes or spicate panicles, glandular-puberulous rachises, inconspicuous flowers in clusters of two to four or occasionally single, small, five-merous, mostly equal corollas 3 to 3.5 mm long with two stamens and two staminodes, anthers two-celled, with one cell usually aborted or vestigial, sterile, and much smaller; leaves immediately subtending the inflo- rescence sessile with ovate laminae and rounded to cordate bases. It resembles Pseuderanthemum cordatum, but is distinguished from it by its mi- nute flowers and inflorescences with glandular- puberulous rachises (vs. puberulous). It also re- sembles P. cuspidatum, but may be distinguished from it by its minute flowers which occur in fas- cicles (vs. solitary flowers which are much larger). Carlowrightia Gray REFERENCES— A. Gray, Contributions to the botany of North America. Proc. Amer. Acad. Arts 13: 361-374. 1878. T. F. Daniel, Carlowrightia (Acanthaceae). Flora Neotropica 34: 1-116. 1983. Subshrubs and suffrutescent herbs. Leaves opposite, sessile to petiolate; laminae linear to ovate, entire. Flow- ers in axillary clusters or more commonly in spicate panicles. Flowers with a 5-merous calyx; corollas 4-part- ed, ranging from nearly regular to strongly zygomorphic, the lower lobe somewhat keeled and containing the an- thers and style in bud, upper lobe spatulate and banner- like, often marked with a distinctively colored eye and/ or radiating nerves; stamens 2, these follow the lower lip out the corolla tube, anther 2-celled with the cells equal and nearly parallel, opening toward the upper lip. Fruits with flattened stipes, compressed-ovoid, acuminate heads; seeds usually 4, ovoid, disklike, smooth, muricate to tuberculate, apically rounded and basally obliquely notched. This is a genus endemic to the New World, with 20 species ranging from the southwestern United States to Costa Rica. Carlowrightia arizonica A. Gray, Proc. Amer. Acad. Arts 13: 361-374. 1878. Carlowrightia costaricana Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1206. 1938. Erect, suffrutescent plants to ca. 60 cm tall; internodes between leaf-bearing nodes 2-4 cm long, 3.5-4.5 mm thick, terete or subterete, retrorse pubescence. Leaves with short, puberulous petioles 2-6 mm long; laminae ovate, 3-4.5 cm long, 1.1-2.3 cm broad, apically acu- minate, basally obtuse to rounded, margins entire to undulate, puberulous above and beneath, cystoliths vis- ible on both surfaces, more obscure below. Inflores- cences in lax terminal and subterminal spicate panicles, rachis puberulous; flowers sessile, mostly 1 per node, but occasionally 2 per node and opposite or 2 per bract; bracts and bracteoles subulate, 2-2.5 mm long, 0.5 mm broad, puberulous. Flowers with subulate, puberulous calyx segments, 1.2-1.5 mm long, ca. 0.5 mm broad; corolla bilabiate, puberulous, 9-15 mm long, ca. 1 mm broad basally, drying purplish, white with a yellowish eye bordered by purplish radiating lines on the upper lobe, narrowly oblong lobes twice the length of the tube; stamens exserted. Fruits clavate, 9-10 mm long, ca. 3 mm broad, 1 mm thick, apically attenuate, the slender stipe ca. half the length of the fruit, capsule dark purple or black when dry, glabrous; seeds 4, cordate, ca. 3 mm long, 2.8 mm broad, drying purplish, papillose. This species extends from Texas and Arizona to Costa Rica. Only one collection from Costa Rica has been observed, Standley & Valeria 44890 (us). This was in dry forest in the vicinity of Li- bano, Guanacaste, at an elevation of 260 to 360 m. Flowering during January. In other areas it is 34 FIELDIANA: BOTANY often found in rocky thickets with dry soil at sim- ilar elevations. Carlowrightia arizonica is recognized by lax spi- cate panicles, bilabiate corollas with exserted sta- mens that have near equally attached anther cells, dark purple or black capsules when dry that have a narrow stipe nearly equal to half the capsule length, and short-petioled leaves. It may be con- fused with either Justicia comata or Justicia pec- toralis, but these differ in their superposed anther cells and inflorescences with glandular puberulous rachises. Chaetochlamys Lindau Plants of deciduous forest formations in the Pa- cific coastal area at elevations of near sea level to 1 000 m in Tilaran and San Ramon and one col- lection from near the Panamanian border in Li- mon Province, Hazlett 5207 (F). Also found in similar habitats of Panama. Flowering collections have been made from December through March. Chaetochlamys panamensis can be recognized most easily by its inflorescences in sessile, terminal heads with conspicuous ciliate, linear bracts. It most closely resembles Chaetothylax leucanthus, but can be distinguished from it by its longer co- rolla (more than 20 mm long vs. 15 mm), five- parted calyx (vs. four-parted), and longer capsules (11 to 12 mm long vs. 6 mm). Herbs to suffrutescent plants. Leaves petiolate; lami- nae ovate to lanceolate. Flowers borne in short spikes that are crowded into a terminal, congested head; bracts and bracteoles narrow; calyx segments 5, equal, narrow; corolla salverform, bilabiate, upper lip 2-lobed, lower lip 3-lobed; stamens 2, exserted, anthers 2-celled, cells at- tached at different heights, the lower cells spurred. Fruits a clavate capsule; seeds 4. Chaetochlamys is native to the Neotropics, with at least seven species known from South America and Central America. Chaetochlamys panamensis Lindau, Report . Spec. Nov. Regni. Veg. 11: 124. 1912. Herbs to 60 cm tall; steins branching, erect or decum- bent, sometimes rooting at lower nodes, internodes be- tween leaf-bearing nodes 2-4 cm long, 1-1.5 mm thick, terete, glabrous to puberulous along 2 lines. Leaves with petioles subsessile to 3 mm long, petioles puberulous to glabrous; laminae lanceolate-elliptic, 2.5-10.5 cm long, 1. 1-3.2 cm broad, apically acuminate, basally attenuate, margins entire to undulate, cystoliths dense, prominent on both surfaces, glabrous to sparingly puberulous along costa and veins. Inflorescences in terminal heads, sessile; bracts linear, to 22 mm long, 0.75 mm wide, ciliate, the hairs to 1 .5 mm long; bracteoles similar, but shorter and narrower. Flowers sessile; calyx segments lanceolate, 10 mm long, 1 .5 mm wide, ciliate; corolla pink or purplish, sparingly glandular-puberulous, to ca. 4 cm long, the tube cylindrical ca. 22 mm long, 4 mm broad, the upper lip oblong, 10 mm long, 3 mm broad, apically 2-lobed, the middle lobe orbicular, 10 mm in diameter, the lateral lobes elliptic, 1 2 mm long, 8mm broad; stamens exserted to just above the mouth, anther cells unequal, inserted at different heights, the upper cell 2 mm long, basally acute, lower cells 2.5 mm long, basally subcalcarate, the connective ca. 1 mm broad. Fruits stout-clavate, 11-12 mm long, 4.5-5 mm broad, 2 mm thick, glabrous, peri- anth persistent except the early deciduous corolla; seeds 4, suborbicular, not flattened, 2-2.5 mm in diameter. Chaetothylax Nees Herbs. Leaves petiolate; laminae ovate to elliptic. Flowers more or less secund, borne in short, dense ter- minal and axillary spikes which often form panicles; bracts narrow with filiform tips, puberulous with ciliate margins; calyx 4-merous, segments resembling the bracts; corollas white to purplish, puberulous, bilabiate, lips suhcqual, upper lip narrowly ovate, lower lip deeply 3-lobed, the lobes rounded; stamens 2, slightly exserted, not exceeding the corolla lips, anthers 2-celled, upper cell oblong, lower cell reduced or vestigial or, as in the type species, hairlike, the lobes separated by a slender filiform connective. Fruit a clavate capsule; seeds 4, pa- pillose or puberulous. Chaetothylax is native to the Neotropics, oc- curring in Central and South America. There are at least eight species known. Chaetothylax leucanthus Leonard, Contr. U.S. Natl. Herb. 31: 663. 1958. Figure 4. Herbs to 60 cm tall; stems erect, terete with retrorse hairs along 2 lines, internodes between leaf-bearing nodes 1.5-3 cm long, 1-1.5 mm thick. Leaves with petioles short, to 5 mm long, strigose; laminae ovate-elliptic to elliptic, 2-10 cm long, 0.8-2.8 cm broad, apically cus- pidate, basally attenuate, margins entire to crenate, gla- brous above, cystoliths dense and prominent above. In- florescences in compact, headlike terminal and axillary spicate panicles, spikes to 4 cm long, 1 cm wide, spikes dense, more or less 1 -sided; panicles subtended by small leaves; bracts narrowly subulate, to 9 mm long, 0.5 mm broad, ciliate and moderately puberulous. Flowers with 4-merous calyx, segments narrowly subulate, to 1 1 mm long, ca. 0.4 mm wide, ciliate and minutely glandular- puberulous; corolla white, ca. 14 mm long, puberulous except at base, the tube cylindrical, 9 mm long, 0.5 mm broad, the upper lip ovate-oblong, 3.5 mm long, 1 mm BURGER: FLORA COSTARICENSIS 35 broad, apically emarginate, the lower lip 3-lobed, ca. 3.25 mm long, the lobes oblong, 2 mm long, the middle lobe 1.5 mm broad, the lateral lobes 1.25 mm broad, all apically obtuse; stamens exserted ca. half the length of the upper lip, the upper anther cell 0.75 mm long, lower cell attached 0.5 mm below the upper cell, reduced, ster- ile, filaments glabrous. Fruit clavate, 6 mm long, 1 mm broad, 2 mm thick, puberulous, perianth persistent; seeds 4, orbicular, slightly flattened, 1 mm in diameter, 0.5 mm thick, puberulous. Plants of clearings, roadsides, and trails in de- ciduous forest formations in Guanacaste and Ala- juela from near sea level to 500 m elevation. While only two collections (both by Brenes s.n.) have been observed from Costa Rica, it is assumed that this species is more common than this low number suggests, since it is rather inconspicuous and can easily be mistaken for another species which is more frequently collected. Chaetothylax leucanthus can be recognized most easily by its inflorescences in sessile, headlike pan- icles. It is easily mistaken for Chaetochlamys pan- amensis, but can be distinguished from it by its shorter corolla and capsule and its four-parted ca- lyx. Chamaeranthemum Nees Herbs or small shrubs. Leaves petiolate to sessile; lam- inae lanceolate to ovate-lanceolate, entire to undulate, cystoliths dense and prominent on both surfaces. Inflo- rescences in loose terminal spikes or spicate panicles. Flowers sessile or pedicellate, 5-merous with inconspic- uous bracts and bracteoles; corolla salverform, lobes sub- equal; stamens 4, included, anterior stamens with 2-celled anthers, posterior stamens with 1 -celled anthers or ster- ile; capsules 2-4 seeded, with a long, narrow stipe. Chamaeranthemum is native to tropical Amer- ica with eight species reported from Costa Rica and northern South America. Key to Species of Chamaeranthemum la. Leaves, stems, and rachises hirsute . 1 b. Leaves, stems, and rachises glabrous C. durandii .C. tonduzii Chamaeranthemum durandii Leonard, Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1208. 1938. Fig- ure 4. Herb to 56 cm tall; younger stems quadrangular, older stems terete, internodes between leaf-bearing nodes 2.5- 7.5 cm long, 1-1.5 cm broad, hirsute. Leaves petiolate to sessile, petioles to 6 mm long, pubescence like that of the stem; laminae ovate-lanceolate, 4.5-10 cm long, 1 .6- 4.5 cm broad, apically acuminate, basally rounded to obtuse, margins entire to undulate, sparingly pilose above, pilose below, cystoliths dense and prominent on both surfaces. Inflorescences of loose terminal spicate pani- cles, to 1 5 cm long, pubescence of peduncle and rachis that of the stem plus minute puberulence; bracts and bracteoles subulate ca. 1.5-2.0 mm long, 0.5 mm broad, puberulous. Flowers sessile, to 2 cm long; calyx segments linear-subulate, 2.5-3 mm long, 0.5 mm broad, puber- ulous-glandular; corolla pink or lavender, tube salver- form, puberulous-glandular, 10-12 mm long, ca. 1 mm broad, expanding to 1.5 mm broad at midpoint, lobes 5, subequal, oblong, ca. 8 mm long, 5 mm broad, apically rounded; stamens 4, included, anthers of anterior sta- mens 2-celled, ca. 1.5 mm long, 0.5 mm broad, apically and basally acute, filaments short, 1 mm long, glabrous, posterior stamens sterile, 1.1 mm long. Fruits clavate, 1 5 mm long, 3 mm broad, 2 mm thick, apically apiculate, puberulous; seeds 2—4, flattened, suborbicular, ca. 2 mm in diameter, slightly winged, glabrous. Found in Costa Rica in the shade of both trop- ical moist and wet forests of the Pacific watershed ranging from Valle del General and Valle del Di- quis to Quepos at elevations of 30 m to ca. 800 m. Flowering collections have been made from January, February, and March. Chamaeranthemum durandii can be recognized by its lavender corollas with five equal lobes and a narrow cylindrical tube with four short, included stamens, the posterior pair being sterile. It can be distinguished from C. tonduzii by its hirsute leaves, stems, and rachises. Chamaeranthemum tonduzii Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and in Pitt., Prim. fl. costaric. 2: 303. 1900. Herb to 42 cm tall; younger stems quadrangular, older stems terete, internodes between leaf-bearing nodes 1 .6- 4 cm long, 1 mm broad, puberulous along 2 lines. Leaves petiolate to sessile, petioles to 5 mm long, pubescence like that of the stem; laminae lanceolate to lanceolate- ovate, 3-7.5 cm long, 1.5-3 cm broad, apically acumi- nate, basally acute, margins entire to undulate, glabrous above and below, cystoliths prominent and abundant on 36 FIELDIANA: BOTANY both surfaces. Inflorescences of loose, terminal spikes or spicate panicles to 10 cm long, peduncle and rachis gla- brous; bracts and bracteoles subulate, to 2 mm long, 1 mm broad, puberulous. Flowers sessile or with pedicels to 2 mm long; calyx segments linear-subulate, 1.5-2.5 mm long, 0.5 mm broad, glandular-puberulous; corolla color not observed, salverform, tube ca. 6.5 mm long, 0.5 mm broad at base expanded to 1 .25 mm broad above midpoint, puberulous, lobes 5, subequal, oblong, api- cally obtuse, ca. 4.5 mm long, 2.5 mm broad, puberu- lous; stamens 4, included, anthers of anterior stamens 2-celled, apically and basally acute, ca. 1.5 mm long, 0.75 mm broad, filament 0.5 mm long, glabrous, pos- terior stamens sterile, 0.75 mm long. Fruits clavate, ca. 14 mm long, 3 mm broad, 1 mm thick, apically apiculate, puberulous to subglabrous; seeds 4, suborbicular, flat- tened, ca. 2 mm in diameter, including a wing ca. 0.5 mm broad surrounding the seed, glabrous. resembles C. durandii and can be distinguished from it by its glabrous leaves, stems, and rachises. Dicliptera Juss. Herbs or shrubs, stems more or less hexagonal in cross section. Leaves petiolate; laminae mostly ovate with en- tire or undulate margins. Inflorescences of cymes, spikes, panicles, or thyrses with flowers or flower clusters sub- tended by 2-4 pairs of conspicuous involucrate bracts. Flowers sessile or pedicellate, often showy; calyx 5-mer- ous, hyaline; corollas narrow, slightly funnelform, bila- biate; stamens 2, anthers 2-celled, cells often unequal; staminodes absent. Fruits of ovate, elliptic, or clavate capsules; seeds 2 or 4. Found in Costa Rica in tropical moist forests at lower elevations of the Caribbean watershed, with collections ranging from the Panamanian border to Livingston on the Rio Reventazon. Flowering collections have been made from March and July. Chamaeranthemum tonduzii can be recognized by its salverform corollas, four short, included sta- mens, with the posterior pair being sterile. It closely A total of 1 50 species of New and Old World Tropics have been described for this genus, which is most easily recognized by its hexagonal stems and reduced cymes with bracts. It has been con- fused with Blechum and Tetramerium, but can be distinguished from these by the above character- istics. Key to Species of Dicliptera la. Inflorescences of axillary cymes; corollas more than 2 cm long 2a Ib. Inflorescences of spikelike racemes or thyrses or axillary heads with peduncles; corollas less than 2 cm long 5a 2a. Inflorescences with peduncles less than 2 cm long; capsules up to 8 mm long D. pallida 2b. Inflorescences with peduncles more than 2 cm long; capsules more than 10 mm long .... 3a 3a. Floral bracts lanceolate and glandular-puberulous; flowers subsessile, pedicels ca. 1 mm long D. skutchii 3b. Floral bracts oblong, sparingly puberulous, or glabrous; flowers pedicellate, pedicels more than 2 mm long 4a 4a. Floral bracts apically rounded and apiculate; capsules puberulous; corolla puberulous D. trifurca 4b. Floral bracts apically acute to obtusish; capsules glandular-puberulous; corollas glandular-pu- berulous D. iopus 5a. Corollas white with lavender stripes; floral bracts to 7 mm long D. imbricata 5b. Corollas pinkish or lavender; floral bracts 9-14 mm long 6a 6a. Floral bracts spatulate, awned; inflorescence of spikelike racemes and/or axillary umbels D. unguiculata 6b. Floral bracts obovate or elliptic, not awned; inflorescence of peduncled axillary heads D. podocephala Dicliptera imbricata Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1210. 1938. Figure 4. Herb to 1 m tall; stems branched, sometimes rooting at lower nodes, internodes between leaf-bearing nodes 3-6 cm long, 1.5-2.5 mm thick, puberulous mostly at the angles in younger portions, basal portions terete and glabrous. Leaves with petioles to 4 cm long, puberulous; laminae ovate-elliptic, 3-10 cm long, 1-4 cm broad, apically acuminate, basally attenuate, margins entire, BURGER: FLORA COSTARICENSIS 37 glabrous, cystoliths dense and prominent on both sur- faces. Inflorescences of terminal and axillary spikelike thyrses, to 4 cm long, 1 .6 cm broad excluding corollas, 2 fanlike cymes of 3-5 flowers each at every node, each cyme subtended by a pair of oblong-lanceolate bracts 3- 8 mm long, 0.5-1 mm wide, puberulous; bracts sub- tending the flowers unequal, obovate, to 7 mm long, 3.5 mm broad, whitish hasally, greenish apically, strongly 3-nerved, puberulous with ciliate margins; outer pair of bracteoles linear, slightly falcate, 5 mm long, 0.75 mm broad, apically acuminate, ciliate, inner bracteoles lan- ceolate, 4 mm long, 0.5 mm broad, ciliolate. Flowers to 16 mm long; calyx segments linear-subulate, 3 mm long, 0.25 mm broad, minutely puberulous; corolla white, lav- ender-striped, puberulous, the tube cylindrical, 10 mm long, 1 mm broad, the lips oblong, 6 mm long, 2 mm broad, apically minutely dentate; stamens exserted to near the tips of the lips, filaments to ca. 5 mm long, puberulous; anther cells superposed, basally obtuse. Fruits elliptic, ca. 4 mm long, 1 mm broad, puberulous toward tip; seeds 2, oval, flattened, ca. 1 mm long, roughened. Found in clearings and along trails in deciduous forest of Pacific watershed in Alajuela and Guan- acaste provinces at elevations of 600 to 2000 m. Flowering collections have been made from late December through March. Dicliptera imbricata is recognized by its dense, spikelike thyrses, its small (7 mm long), obovate bracts subtending each flower, and its small (16 mm long), white corollas. It is most easily confused with D. unguiculata, which has spine-tipped bracts and rose-purple corollas. Dicliptera iopus Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 1 89. 1 898, and in Pitt., Prim, fl. costaric. 2: 305. 1900. Figure 4. Shrub to 3 m tall; internodes between leaf-bearing nodes 2-6 (9) cm long, 1.0-2.5 mm thick, strigose to glabrous. Leaves with petioles to 7 cm long, glabrous to sparingly puberulous; laminae ovate to elliptic, 3-15 cm long, 1-5.6 cm broad, apically acuminate, basally acute, margins entire, glabrous to sparingly strigose on both surfaces, cystoliths visible on both surfaces. Inflores- cences in axillary cymes, each cyme subtended by 2 el- liptic to ovate-oblong bracts, to ca. 3 cm long, 1 cm broad, apically acute, basally acute, glabrous to sparingly pilosulous; 2 bracts of unequal length subtending each flower, both linear-oblong, fused at base, the longer one 10-16 mm long, 4 mm broad at base, the shorter bract to 1 2 mm long, 4 mm wide, both apically acute, keeled, sparingly puberulous on both surfaces; 4 bracteoles, ob- lanceolate, to 8 mm long, 2 mm broad, puberulous to- ward tip, inner pair smaller, 3.5 mm long, 0.5 mm broad; peduncles to ca. 6.5 cm long, subhexangular, puberulous along ridges, pedicels of middle pair in a cyme longest, to ca. 4 mm long, sparingly puberulous. Flowers usually 3 per cyme, although some have 2 or 4, 4-5 cm long; calyx 5-merous, coalesced for ca. V* of its length, to 7 mm long, lobes linear, ca. 3.5 mm long, subtomentose; corolla lavender, inflated at about the middle of the tube, bilabiate, tube to 4 cm long, 2.5 mm broad at the base, 8 mm broad at the mouth, glandular-puberulous both inside and out, upper lip entire, acute, ca. 9 mm long, 5 mm broad, lower lip ca. 1 1 mm long, 8 mm broad at base, with 3 small, suborbiculate lobes at its apex, ca. 0.5 mm long; stamens exserted, filaments flattened, pi- lose, anther cells equal; style exserted, glabrous. Fruits clavate, ca. 1 5 mm long, apically apiculate, densely glan- dular-puberulous, 2 seeds per capsule, flattened, subor- bicular, ca. 3 mm in diameter. Found rather commonly over a wide range in Costa Rica in open meadows and cloud forests at elevations of 1300 to 3200 m on both the Carib- bean and Pacific slopes; also common in Panama and probably Nicaragua. Flowering collections have been made from September through March. Dicliptera iopus is recognized by its peduncled cymes usually in clusters of three, acute to obtusish floral bracts, subtomentose pedicels, and glandu- lar-puberulous corolla and capsule. It is easily con- fused with D. trifurca, which differs in its rounded and apiculate floral bracts, glabrous pedicels, and puberulous corollas and capsules. Dicliptera pallida Leonard, Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1211. 1938. Figure 5. Herb to 1.2 m tall; stems erect, branched, internodes between leaf-bearing nodes 4-10 cm long, 1.5-3.5 mm thick, subtomentose. Leaves with petioles to 1 .6 cm long, subtomentose to glabrescent; laminae ovate, 3.5-8 cm long, 2-3.5 cm broad, apically acuminate, basally acute and slightly tapering, margins undulate, sparingly pilose above and below, cystoliths visible, but not prominent. Inflorescences in sessile or subsessile, axillary cymes, peduncles to 7 mm long, subtomentose; flowers of the upper cymes mostly sessile, those of the lower, short- pedicelled, the pedicel of the middle flower longer than that of the lateral flowers, all pedicels tomentose; bracts subtending the cymes, leaflike, 2-3 cm long, 1-1.5 cm broad, those subtending the flowers unequal, elliptic to oblanceolate, the larger of the pair ca. 16 mm long, 5 mm broad, the shorter ca. 13 mm long, 2.5 mm broad, both bracts apically acuminate, whitish toward the base, hirtellous; bracteoles unequal, both narrowly lanceolate, the longer to ca. 1 2 mm long, 1.25 mm broad, the shorter slightly smaller, both ciliolate. Flowers 2.5-3.5 cm long; calyx to ca. 5 mm long, fused ca. 'A of length, segments lanceolate, ca. 3 mm long, 0.75 mm broad, minutely ciliolate; corollas red, puberulous, the tube ca. 2 cm long, 2 mm broad at the base, narrowing to 0.5 mm broad just above the ovary, expanding to 3 mm broad at the mouth, the lips ca. 8 mm long, the upper emarginate, the lower minutely 3-lobed; stamens exserted nearly to the tip of the upper lip. Fruits clavate, to 8 mm long, 5 mm broad, 1 mm thick, puberulous; seeds discoid, ca. 3 mm in diameter, puberulous. 38 FIELDIANA: BOTANY Found at middle to higher elevations in thickets and along streams on the slopes of the Cordilleras Central and Talamanca from 635 to 1700 m al- titude. This species is apparently endemic to Costa Rica. Flowering collections have been made from December through February. Dicliptera pallida is recognized by its axillary cymes with sessile to subsessile flowers, floral bracts and bracteoles with whitish bases, and compara- tively long (more than 2 cm) red corollas. It can be distinguished from the much more common and weedy D. unguiculata, which it most closely resembles, by these characteristics. Dicliptera podocephala Donnell Smith, Hot. Gaz. (Crawfordsville) 48: 299. 1909. Figure 5. Herb to subshrub, to 1.5m tall; stems erect, branching, internodes between leaf-bearing nodes 2-4 cm long, 1- 2 mm thick, glabrous. Leaves with petioles to 2.5 cm long, puberulous; laminae ovate, 2.5-10 cm long, 1-4 cm broad, apically acuminate, basally acute, slightly ta- pering, margins entire, cystoliths dense and prominent on both surfaces, glabrous. Inflorescences in peduncled axillary heads, 1 -several per axil, peduncles to 9.5 cm long, minutely puberulous mostly along 2 lines; bracts subtending the heads subulate, 3.5-5 mm long, 1 mm broad, glabrous; fertile flowers usually 5-7 per head, bracts subtending the flowers obovate to elliptic, 1 of each pair slightly larger than the other, the middle pair the largest, 10-14 mm long, 8-9 mm broad, apically rounded or obtuse, apiculate, narrowed at the base, subglabrous, margins ciliolate; bracteoles 4, linear-lanceolate, 5-8 mm long, puberulous. Flowers 14 mm long; calyx segments resembling bracteoles, slightly shorter; corolla pinkish lavender, 14 mm long, puberulous; stamens exserted. Fruits ovoid, 6 mm long, 4 mm broad, 1 mm thick, puberulous; seeds discoid, 2.5 mm in diameter, puber- ulous. Found in damp meadows, streamsides, and along trails in Cartago and Puntarenas provinces near 1000 m elevation and at 90 m elevation at La Selva in Heredia Province. Only two collections, Allen 5902 (F) and Opler 597 (F), have been ob- served other than the type collection. Flowering collections have been made in February and March. Dicliptera podocephala is recognized by its pe- duncled axillary heads with five to seven flowers per head and its obovate to elliptic bracts with rounded, apiculate tips. With these characteristics it can be distinguished from D. unguiculata, which it most closely resembles. Dicliptera skutchii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1213. 1938. Figure 5. Shrub to 3 m tall; erect, branching, internodes between leaf-bearing nodes 5-7 cm long, 3-5 mm thick, puber- ulous along 6 lines. Leaves with petioles 1-6.5 cm long, sparingly puberulous; laminae ovate-elliptic, 9-17 cm long, 3.5-7 cm broad, apically acuminate, basally acute, slightly tapering, margins entire, cystoliths relatively ob- scure above, more visible below, glabrous above and below. Inflorescences in axillary cymes on peduncles to 8 cm long, peduncles with appressed puberulence; ped- icels short, 1-2 mm long, hirtellous; bracts subtending the cymes leaflike, to ca. 8 mm long; bracts subtending the flowers lanceolate, glandular-puberulous, unequal, the longer 10 mm long, 3 mm broad, the shorter 7 mm long, 2.8 mm broad; 2 pairs of bracteoles subtending each flower, the outer pair lanceolate, 6.5 mm long, 1.5 mm broad, glandular-puberulous toward apex, the inner pair lanceolate-elliptic, 3.5 mm long, 1 mm broad, gla- brous. Flowers 3 per cyme, to 6 cm long; calyx to ca. 7 mm long, coalesced for ca. [h its length, the lobes subu- late, 3.5 mm long, 1 mm broad at the base, margins glandular-ciliolate; corolla purple, glandular-puberulous, the tube ca. 5 cm long, 3 mm broad at base, expanding to 10 mm broad above the middle and narrowing to 7 mm broad at the mouth, the lips to 10 mm long, 1 acute and finely notched at the tip, the other minutely 3-lobed, the lobes suborbicular, ca. 0.75 mm in diameter; stamens exserted to nearly the end of the lips. Fruits not seen. Found in cloud forests at elevations around 1000 m in the vicinity of El General. Only one collec- tion, Skutch 4140 (MO), in addition to the type specimen, Skutch 3090 (us), has been observed. Flower collection has been made in February. Dicliptera skutchii can be recognized by its pe- duncled cymes with short pedicels and its glan- dular-puberulous bracts, bracteoles, and corollas. By these characteristics it can be distinguished from D. iopus, which it most closely resembles. Dicliptera trifurca Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 1854: 172. 1855. Dicliptera trifurcata (Oerst.) Hemsl., Biol. Cen- tralbl. 2: 525. 1881. Diapedium trifurcatum (Oerst.) Kuntze, Revis. gen. pi. 2: 485. 1891. Figure 5. Herbs to subshrubs, to 2 m tall; stems branching, in- ternodes between leaf-bearing nodes 1.5-8 cm long, 1- 3 mm thick, glabrous. Leaves with petioles to 5.3 cm long, glabrous to sparingly puberulous along 2 lines; lam- inae ovate-elliptic, 2.5-15 (18.5) cm long, 1-4 (5.6) cm broad, apically acuminate, basally attenuate, margins undulate, glabrous above, glabrous to sparingly puber- ulous along costa and veins below, cystoliths more prom- inent below. Inflorescences in axillary cymes of 3 flowers each; peduncles to 8 cm long, mostly glabrous; bracts subtending the cymes leaflike, frequently exceeding the calyx; floral bracts oblong to oblong-spatulate, of un- equal lengths, the larger to 1 1 mm long, 2 mm broad, the smaller to 7 mm long, 1.5 mm broad, both apically BURGER: FLORA COSTARICENSIS 39 rounded and apiculate, glabrous; bracteoles oblong-lan- ceolate, to 5 mm long, 1 mm broad, sparingly puberu- lous. Flowers with pedicels to 3 mm long, glabrous to sparingly puberulous; calyx 6 mm long, segments subu- late, 4 mm long, 0.6 mm broad, puberulous; corolla red- violet or purplish, puberulous, 3.5-5.5 mm long, the tube to 4 cm long, 2 mm broad basally, expanding to 7 mm broad at the mouth, the lips oblong, to ca. 1.5 cm long, the upper lip 7 mm broad, apically acute, lower lip 3- lobed, the lobes subequal, ca. 0.75 mm long; stamens exserted to the tip of the upper lip, filaments sparingly puberulous; anther cells parallel, ca. 3 mm long, basally apiculate. Fruits stipitate, to 1 8 mm long, 4 mm broad, 2 mm thick, puberulous; seeds 4, discoid, ca. 4 mm in diameter, papillose. Found in higher elevation rain forests or cloud forests of the Cordilleras Central and Talamanca at elevations from 1300 to 2300 m. Flowering collections have been made from December through June; also found in adjacent Panama. Dicliptera trifurca is recognized by its pedun- cled, three-flowered cymes with glabrous pedicels, rounded and apiculate floral bracts, puberulous corollas more than 3 cm long, and puberulous cap- sules. These features distinguish it from D. iopus which it closely resembles. Dicliptera unguiculata Nees in Benth., Bot. voy. Sulphur 149. 1844. Figure 5. Herbs to 1 .5 m tall; steins erect or ascending, branched, inter nodes between leaf- bearing nodes 5.5-10 cm long, 1-2.5 mm thick, increasingly pilose toward tips, terete and glabrous below. Leaves soon deciduous, petioles to 9 cm long, pilosulous to subglabrous; laminae ovate to elliptic, 5-9 cm long, 3-5.5 cm broad, apically acuminate and often apiculate, basally acute to attenuate, margins entire to undulate, both surfaces of younger leaves pilose, older leaves subglabrous. Inflorescences composed of ax- illary umbels or in crowded terminal and axillary spike- like racemes to 9 cm long, 1.75 cm broad, bracts spat- ulate, unequal, the larger 10-1 2 mm long, 5.5 mm broad, the smaller 8-9 mm long, 3 mm broad, both apically awned, the awn ca. 2 mm long, both basally clawed, pilosulous, margins densely ciliate, 3-nerved; bracteoles linear-lanceolate, 3.5 mm long, 0.5 mm broad, ciliolate. Flowers sessile, to 15 mm long; calyx 2.5 mm long, coalesced a little more than Vi its length, lobes 1 mm long, minutely puberulous; corolla pink or lavender, puberulous, the tube 7 mm long, 1 mm broad at the base, 1.25 mm broad at the mouth, the lobes obovate, 8 mm long, 4 mm broad, 1 is apically obtuse, the other apically minutely 3-lobed. Fruits elliptic, 4 mm long, 2 mm broad, 0.75 mm thick, puberulous; seeds discoid, ca. 1.5 mm in diameter, puberulous. This is the most common species of Dicliptera in Costa Rica. It is found in moist thickets, open fields, and along roadsides at elevations of 900 to 1500 m. Flowering collections have been made between October and April. Its range extends north to Mexico, but it has not yet been collected in Panama. Dicliptera unguiculata is recognized by its inflo- rescences of axillary umbels and spikelike ra- cemes, with conspicuous obovate, awned bracts, short (to 1 5 mm long), pink or lavender corollas, and often leafless condition. By these character- istics it can be distinguished from D. pallida, a much less commonly occurring species, which it most closely resembles. It also resembles D. im- bricata, which is much more common than D. pallida, but more easily distinguished by the above characteristics. Dyschoriste Nees REFERENCE— C. E. Kobuski, Ann. Missouri Bot. Card. 15: 9. 1928. Herbs to suffrutescent plants, stems prostrate, ascend- ing or erect, glabrous or pubescent. Leaves sessile or petioled; laminae mostly entire. Inflorescences cymose, capitate, or spicate, terminal or axillary. Flowers sub- tended by foliaceous bracts and bracteoles; calyx deeply 5-parted, usually subulate-setaceous, ciliate; corolla 5-lobed, obscurely or distinctly bilabiate, tube erect, limb spreading; stamens 4, didynamous, anthers 2-celled, cells oblong, minutely calcarate to mucronate at the base. Fruit a capsule which is included in the persistent calyx, oblong-linear, glabrous, 2-4 seeded, separating with dif- ficulty at maturity into 2 valves; seeds flattened, subor- bicular, with mucilaginous hairs when wet. Dyschoriste is a widely distributed genus of about 100 species in both tropical and subtropical re- gions of the world. A few species are found in warm, temperate areas. Several species occur in Central America. Dyschoriste valeriana Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1215. 1938. Figure 5. Herbs, prostrate, erect or ascending, to 60 cm tall, frequently rooting at nodes of lower stem, stems qua- drangular; internodes between leaf-bearing nodes 1.5- 4.5 cm long, 0.75-1 .5 mm thick, puberulous to glabrous. Leaves petiolate to sessile, petioles to 1 cm long, glabrous to puberulous along 2 lines; laminae ovate, 2.5-5 (6) cm long, 1-2 (3) cm broad, apically acute to rounded or obtuse, basally attenuate, margins entire to undulate, cystoliths numerous, prominent above, obscure below, 40 FIELDIANA: BOTANY glabrous to subglabrous on both surfaces. Inflorescences of sessile axillary cymes, cymes of 2-5 flowers; 2 bracts subtending each cyme, bracts oblanceolate, 10-18 mm long, 2.5-6 mm broad, glabrous, margins minutely cil- iolate; floral bracts linear to linear-elliptic, to 10 mm long, 1 mm broad, bracteoles similar, but slightly small- er, margins of both minutely ciliolate. Flowers sessile, to 8-9 mm long; calyx 6-8 mm long at anthesis, 8-1 1 mm long at fructescence, coalesced for '/j-'/z of its length by thin, easily torn tissue, lobes subulate, ca. 5.5 mm long, 1 mm broad at base, single-nerved, sparingly pilose along rib, margins ciliolate; corolla blue or purplish, pu- berulous, the tube 6 mm long, 2 mm broad at base, 2.5 mm broad at the mouth, sub-bilabiate, lobes oblong, apically rounded, upper lip with 3 lobes to 3 mm long, 2 mm broad, lower lip with 2 lobes to 2 mm long, 1.5 mm broad; stamens included, longer filaments to 2 mm long, shorter filaments to 1 mm long, all glabrous, an- thers 1 mm long, minutely calcarate at the base of each cell, both cells joining to form an apiculate tip; ovary glabrous. Fruit oblong-lanceolate, 9 mm long, 2 mm broad, 1.75 mm thick, glabrous; seeds 4, ovate, flattened, 2.5 mm long, 2 mm broad, densely puberulous when wet. Found in disturbed habitats along trails and roadsides in deciduous forest areas of Guanacaste from near sea level to about 650 m elevation. Flowering collections made from December to April. One collection has been made from a similar habitat in Panama. Since many collections of this species have been made near Nicaragua, I expect that D. valeriana occurs there also. Dyschoriste valeriana is recognized by its square stem, small blue or purplish flowers, four stamens with two-celled anthers that have minutely cal- carate bases, and two- to four-seeded capsules. It closely resembles Hygrophila costata, which lacks calcarate anthers and has 1 2 to 16 seeds per cap- sule. Elytraria Michx. Herbs either caulescent or acaulescent. Leaves alter- nate or subopposite, basal or restricted to the terminal portion of the stem in a dense pattern, cystoliths lacking. Inflorescences compact, peduncles with imbricate, co- riaceous bracts. Flowers sessile with persistent perianth except the soon deciduous corolla; bracts single; brac- teoles 2; calyx 4-merous, segments narrow, entire or toothed apically; corollas 5-merous, bilabiate, white or blue, tube narrow, upper lip 1-lobed, lower lip 3-lobed; stamens 2, barely exserted, anthers 2-celled which are equal and parallel; ovary 2-celled; to 10 ovules in each cavity, style 1 . Fruits a capsule, basally constricted and apically acute; seeds borne on papilliform funicles. About 1 7 species have been described for this genus from southern North America plus New and Old World Tropics. Elytraria imbricate (Vahl) Pers., Syn. pi. 1: 23. 1805. Justicia imbricata Vahl, Eclogae ameri- canae 1: 1. 1796. Tubiflora squamosa (Jacq.) Ktze., Revis. gen. pi. 2: 500. 1891. is no longer a valid synonym. The genus Tubiflora was re- jected by the International Code of Botanical Nomenclature 1972. Figure 5. A weedy caulescent herb, occasionally acaulescent, to 50 cm tall; internodes between leaf- bearing nodes 0-15 mm long, 2-3 mm thick, glabrous to sparingly puberu- lous or pilose. Leaves alternate, sessile to petiolate, pet- ioles to 2 cm long, pilose; laminae ovate-lanceolate, 3.5- 13 cm long, 2—4.5 cm broad, apically acute, basally at- tenuate, narrowing to a winged petiole, margins undu- late, sparingly pilose above and below, more dense on costa and lateral veins. Numerous scapes borne at leaf axils, 5-28 cm long, branched or simple, covered with lightly appressed, sheathing, ovate, or subulate bracts. Inflorescences in terminal spikes, 1-several, 1.5-4 cm long, 4 mm broad; bracts oblong to elliptic, 4-6 mm long, 1.5-2 mm broad, awn-tipped and stiff with upper bracts bearing 4 hyaline wings, 2 rounded extending from base and 2 apical extending parallel to the apex; 2 brac- teoles per flower, subulate, to 3 mm long, costa ciliate. Flowers sessile, 5-8.5 mm long; calyx segments to 4.5 mm long, posterior segment elliptic, bidentate, 1 .25 mm broad, lateral segments subulate, 0.75 mm broad, an- terior segment elliptic, bidentate, 0.75 mm broad, all segments transparent, puberulous; corolla pale lavender or blue, glabrous, tube cylindrical, 4-7 mm long, 0.5 mm broad, lower lip spreading, to 4 mm long, 3-lobed, the middle lobe 2 mm broad, 2-lobed at apex, the lobes rounded, ca. 1 mm long, the lateral lobes similar, but slightly smaller, the upper lip erect, to ca. 2 mm long, 1 mm broad, apically 2-lobed. Fruits elliptic, ca. 3 mm long, 1 mm broad, glabrous; mature seeds light brown, rounded, triangular, ca. 0.5 mm long, glabrous. Found in dry pastures and along roadsides and trails of the Pacific watershed area at elevations of 50 to 1 1 70 m. Flowering collections have been made from November through March. This species ranges from the southwestern United States to western South America. Leonard (1951) reports that in Central America this plant is used as a remedy for dysentery, and the crushed leaves are said to cure pimples. Elytraria imbricata is recognized by its numer- ous spikes borne on axillary, bracted culms, its small bluish bilabiate corollas with awned, winged bracts, and its capsules lacking hooklike funicles. Superficially, this plant can be confused with Aphe- landra tonduzii, which differs in all of the above characteristics. Graptophyllum Nees Shrubs with branching, ascending stems. Leaves ses- sile to petiolate, laminae often colored and variegated, BURGER: FLORA COSTARICENSIS 41 margins entire or dentate. Flowers pedicellate, in short, nearly sessile cymes, terminal panicles or solitary in the axils of the leaves; calyx 5-merous; corollas red or purple, bilabiate, the tube inflated above, with 2, short, recurved lobes, the lower lip 3-lobed; stamens 2; staminodes 2. Fruits of stalked, clavate capsules; seeds 2-4. Ten species have been described for this genus, occurring in West Africa, New Guinea, Australia, and Polynesia. None is native to the New World. Graptophyllum pictum (L.) Griff., Not. pi. asiat., part 4: 139. 1854. Based on Justicia picta L., Sp. pi. 1: 21. 1763. G. hortense (L.) Nees in Wall., PI. asiat. rar. 3: 102. 1832. Figure 6. Shrubs, stems to 3 m tall; ink-modes between leaf- bearing nodes 4-7 cm long, 3-4 mm thick, glabrous. Leaves sessile to short-petiolate, petioles to 4 mm long, glabrous; laminae ovate to elliptic, 4.5-1 1 (20) cm long, 2-7 (9) cm broad, apically acuminate, basally acute to obtuse, slightly tapering, purplish or green, variously marked with yellow, margins entire, cystoliths dense and generally prominent above, obscure below, glabrous on both surfaces. Inflorescences in terminal racemes, the rachis glabrous or sparingly puberulous. Flowers with glabrous pedicels to 8 mm long, each subtended by 1 subulate bract, bract 3 mm long, 2 mm broad at base, minutely ciliolate, bracteoles 2, similar to bracts al- though slightly smaller; calyx segments subulate, ca. 3 mm long, 1 .25 mm broad at base, glabrous; corollas dark purple or crimson, glabrous outside, glandular-puberu- lous inside, 4.5-8 cm long, tube funnelform, ca. 3 cm long, 4 mm broad basally, 1 1 mm broad at the mouth, upper lip erect, oblong, ca. 1 6 mm long, 1 1 mm broad, apically 2-lobed, lobes rounded, 5 mm long, 4 mm broad, lower lip 3-lobed, lobes oblong, 1 5 mm long, narrowing to a rounded tip, middle lobe 7 mm broad, lateral lobes 6 mm broad; stamens exserted to just below the tip of the upper lip, filaments glabrous, anthers 4 mm long, 1 mm broad, cells slightly unequal at the base, acute api- cally and basally, staminodes to 3 mm long. Fruits not observed. This is a species that is cultivated as an orna- mental in the gardens of tropical America. Ac- cording to Bailey ( 1 949) it is probably native to New Guinea. Graptophyllum pictum is recognized by its mot- tled leaves, relatively large (to 8 cm long), bilabiate corollas, two stamens with two-celled anthers and two staminodes. It resembles various species of Odontonema, but can be most easily distinguished from them by its mottled leaves and larger corol- las. Habracanthus Nees Herbs or shrubs. Leaves usually with ovate to elliptic laminae. Flowers borne in loose to dense panicles; calyx 5-merous, segments narrowly lanceolate; corolla blue, red, or yellow, salverform, bilabiate, upper lip narrow, cuneate, lower lip usually oblong-cuneate, apically 3-lobed, lobes rounded; stamens 2, exserted, divergent, anthers 1 -celled, filaments attached below middle of an- ther; style exserted. Fruit a clavate capsule; seeds 4. This is a genus of about 40 species found in Mexico, Central America, and northern South America. Habracanthus silvaticus Nees in DC., Prodr. 11: 312. 1847. Figure 6. Shrub to 2 m tall, younger stems quadrangular; inter- nodes between leaf-bearing nodes 3-8 cm long, 1-2 mm thick, glabrous to strigose along 2 lines. Leaves with petioles 5-35 (75) mm long, glabrous to sparingly puber- ulous; laminae elliptic to ovate-elliptic, 6-13 cm long, 2.5-5 cm broad, apically acuminate, basally acute, gla- brous to sparingly puberulous along costa and lateral veins, margins undulate, cystoliths prominent on both surfaces. Inflorescences of loose, terminal thyrses to 10 cm long, 5 cm broad, lateral branches of thyrse dichot- omously branched, branches of inflorescence minutely strigose; bracts subulate, to 6 mm long, 1 mm broad at base, glabrous. Flowers with pedicels ca. 2 mm long; calyx whitish to purplish, segments subulate, 3-4 mm long, 0.5 mm broad at base, glabrous; corolla blue, bi- labiate, 12-18 mm long, glabrous, tube ca. 8 mm long, near cylindrical, ca. 1 mm broad, the lips ca. 10 mm long, the upper lip oblong, 2 mm broad, apically round- ed, lower lip elliptic, ca. 4 mm broad, apically 3-lobed, the lobes 0.5 mm long, middle lobe 1 mm broad, lateral lobes 0.5 mm broad; stamens exserted to ca. the tip of the upper lip. Fruit clavate, to 1 5 mm long, 3 mm broad, 2 mm thick, glabrous; seeds flattened, ovate, 2 mm long, 1.5 mm broad, puberulous. A relatively common plant in wet forests and adjacent clearings of higher elevations on Pacific and Caribbean slopes of the Cordillera Central from 975 to 2200 m elevation. Flowering collec- tions have been made from July through March. This species was first described in Chiapas, Mex- ico, and extends from there to Panama. Habracanthus silvaticus is recognized by its bluish calyx and bilabiate corolla, with two sta- mens that have one-celled anthers; also by its in- florescence in a loose, terminal thyrse. It resembles Razisea spicata, but can be distinguished from it in flower color and inflorescence. 42 FIELDIANA: BOTANY Hansteinia Oersted Herbs or shrubs. Leaves opposite, usually petiolate; laminae elliptic or ovate. Inflorescences in loose or com- pact panicles which are sometimes thyrsoid; flowers with 5-merous calyx, calyx segments narrow, equal; corollas red, orange, yellow, or white, cylindrical, narrow basally, abruptly expanded just above the calyx, somewhat obliquely enlarged and oblong, often constricted again at the mouth; upper lip erect, the lower lip barely distinct from the corolla tube, 3-lobed; stamens 2, attached at base of corolla tube, exserted, anthers 1 -celled, basally obtuse. Fruits a capsule, clavate, glabrous, 4-seeded. Fourteen species have been described for this genus, ranging from Mexico to Bolivia. In Costa Rica the species are all found at elevations above 1000 m. Generally these species are not often col- lected in Costa Rica, which may indicate infre- quent occurrence. This genus closely resembles the genera Habracanthus and Razisea. See the genus description of Razisea for a discussion of these differences. Key to Species of Hansteinia 1 a. Leaf bases rounded or subcordate 2a Ib. Leaf bases attenuate (except leaves subtending the inflorescence) 3a 2a. Corolla more than 20 mm long, glandular-puberulous; petioles mostly more than 2 cm long H. ventricosa 2b. Corolla less than 20 mm long, glabrous; petioles less than 5 mm long H. sessilifolia 3a. Capsules less than 12 mm long; leafy internodes 7-8.5 cm long H. gracilis 3b. Capsules more than 1 2 mm long; leafy internodes 6 cm or less long 4a 4a. Inflorescences in fairly open racemose panicles, branches relatively few, erect or ascending; bracts 2-5 mm long H. stricta 4b. Inflorescences in compact cymose panicles, branches numerous, spreading; bracts 5-10 mm long H. blepharorhachis Hansteinia blepharorhachis (Lindau) Durkee, comb. nov. Kolobochilus blepharorhachis Lin- dau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 189. 1898, and in Pitt., Prim. fl. costaric. 2: 309. 1900. Glockeria blepharorhachis (Lindau) Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1218. 1938. Razisea breviflora D. Gibson, Fieldiana, Bot. 34: 77. 1972. Figure 6. Herbs to suffrutescent plants to 2 m tall, stems subquadrangular to quadrangular in younger portions; internodes between leaf-bearing nodes 1.5-8 cm long, 1.5-2.5 mm thick, sparingly pilose mostly along 2 lines. Leaves with petioles 2-6 cm long, sparingly pilose; lam- inae elliptic-ovate, 7-18 cm long, 2-6.5 cm broad, api- cally acuminate, basally attenuate, margins undulate to crenulate, older leaves mostly glabrous, drying to very dark green or nearly black above, younger leaves with pilose hairs restricted to costa and veins, cystoliths abun- dant though minute and obscure, more visible beneath. Inflorescences in long, narrow, terminal thyrsoid pani- cles to 30 cm long, 3-5 cm broad; peduncle to 7 cm long, puberulous, the rachis pilose to subtomentose with brown to reddish brown hairs plus longer, scattered, glandular hairs, pedicels to 8 mm long, pubescence that of the rachis; bracts narrowly subulate, 5-10 mm long, 1.5-3 mm broad, bracteoles similar, but smaller, both puber- ulous plus a few longer glandular hairs. Flowers with 5- merous calyx, segments linear, 4.5-10 mm long, 0.75- 1 mm broad, minutely puberulous, plus much longer glandular hairs; corolla orange or orange-red, 1 5-22 mm long, the tube to ca. 14 mm long, 4 mm broad at base, expanding abruptly to 5-6 mm broad just above the calyx, glabrous outside, pilosulous glandular hairs inside, upper lip erect, oblong, 4-8 mm long, 3-4 mm broad, apically rounded, lower lip short, truncate, 3-lobed, lobes each ca. 1 mm long, suborbicular; stamens exserted to well above the upper lip. Fruits clavate, 13-15 mm long, 3.5-4 mm broad, 2.5-3 mm thick, glabrous; seeds flat- tened, ovate, 2.5-3.5 mm long, 2-2.5 mm broad, surface muricate. Plants of cloud forests at elevations of 1 500 to 2000 m. The locality of the type specimen ( War- scewicz s.n.) is specified only as Costa Rica and Veraguas (Panama). Flowering collections have been made in April, August, and October from the Cordilleras Tilaran, Central, and Talamanca. This species occurs rather frequently in light gaps. It occurs in habitats similar to those of Razisea spi- cata and is also hummingbird-pollinated. Hansteinia blepharorhachis is recognized by its compact and comparatively narrow, but much- branched, cymose panicles (thyrses) combined with leaf blades which are elliptic-ovate with attenuate bases. It is most similar to H. stricta, which differs BURGER: FLORA COSTARICENSIS 43 in its generally smaller, less compact, less branched, broader racemose panicles and smaller bracts (2.5 mm long vs. 5 to 10 mm). Hansteinia gracilis Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 143. 1855. Herb to suffrutescent plant to ca. 60 cm tall; steins subtetragonal, erect, simple or sparingly branched, in- ter nodes between leaf-bearing nodes 7-8.5 cm long, pu- berulous. Leaves with petioles to ca. 2.5 cm long; laminae elliptic-ovate, to 1 8 cm long, 7 cm broad, apically acu- minate, basally attenuate, margins subrepand-crenate, glabrous on both surfaces except minutely strigose on costa and veins. Inflorescences in terminal panicles to 1 5 cm long, rachis glandular-pubescent, hairs rust-col- ored. Flowers with 5-merous calyx, calyx segments red, 7-9 mm long at anthesis, 13.5-16 mm long at fruiting stage, glandular-villous; corolla red, 13.5-16 mm long, tomentose. Fruit 9-1 1 mm long, puberulous. Found in dense, damp woods on Mt. Jaris northeast of Santiago (Santiago de Puriscal) in San Jose Province, elevation 500 to 1000 m. I have seen no collections of this species. The above de- scription was adapted from the original, which was based on a collection in both flower and fruit col- lected in November. Hansteinia gracilis differs from other Costa Ri- can species in the genus by its shorter capsules (9 to 1 1 mm long vs. 1 2 mm or more) and its longer internode lengths (7 to 8.5 cm long vs. 6 cm or less). Hansteinia sessilifolia (Oerst.) Durkee, comb. nov. Glockeria sessilifolia Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 141. 1855. Figure 6. Herbs to suffrutescent plants to 2 m tall, older stems terete, younger stems subquadrangular, glabrous to pu- berulous along 2 lines; internodes between leaf-bearing nodes 4.5-5.5 cm long, 1.5-2.5 mm thick. Leaves sessile (in those subtending the inflorescence) to petiolate, pet- ioles to 4.5 cm long, puberulous; laminae ovate, 5.5-1 1 cm long, 2-6 cm broad, apically acuminate, basally rounded or subcordate in leaves at base of inflorescence to acute in lower leaves, margins entire to undulate, gla- brous or sparingly pilose on both surfaces, hairs mostly restricted to costa and veins, cystoliths abundant on both surfaces though minute and obscure. Inflorescences in loose, terminal, dichotomously branching, thyrsoid pan- icles to 38 cm long, 15 cm broad, sometimes with ad- ditional axillary panicles, rachises and branches puber- ulous to sparingly so, pedicels to 2.5 mm long, sparingly puberulous; bracts narrowly triangular, to 3 mm long, 0.5 mm broad, glabrous to very sparingly puberulous. Flowers with 5-merous calyx, calyx segments linear-su- bulate, to 4-6 mm long, 0.3-0.5 mm broad, sparingly glandular-puberulous; corolla red and yellow, 1 2-22 mm long, glabrous, the narrow basal portion 2.5-3 mm long, 1-2 mm in diameter, the upper portion expanded to 3- 5 mm diameter, then gradually narrowing to 2.5-4 mm diameter at the mouth, the upper lip spatulate, apically acute, 3.5 mm long, 1.5 mm broad at the base, 0.5 mm broad at the lip, lower lip short, recurved, ca. 1 mm long, 3 mm broad, lobes apically rounded; stamens exserted to 5 mm beyond upper lip, anthers 2-2.8 mm long; capsules clavate, 12-14 mm long, 2.3-3 mm broad, 1.8- 2.5 mm thick, glabrous; seeds flattened, elliptic 2-2.2 mm long, 1.2-1.4 mm broad, hispidulous. A plant of the wooded, southern slopes of Vol- can Barva (Barba), Volcan Poas, and the Monte- verde cloud forest area in Puntarenas Province. I have seen only two collections of this species, Leon 79 (F) and Dryer 1671 (F). Flowering collections have been made in September and October. Hansteinia sessilifolia is recognized by its loose, dichotomously branching inflorescences and its short corollas (12 to 22 mm long) with stamens slightly exserted. It most closely resembles H. ven- tricosa, but can be distinguished from it by the above characteristics. Hansteinia stricta (Leonard) D. Gibson, Field- iana, Bot. 34: 63. 1972. Glockeria stricta Leon- ard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1219. 1938. Figure 6. Herb to subshrub to 1.5 m tall, older stems terete, glabrous, younger stems quadrangular, sparingly puber- ulous or scattered brown puberulence along 2 lines; in- ternodes between leaf-bearing nodes 3-6 cm long, 1.5- 2 mm thick. Leaves with petioles 1.5-3.3 cm long, gla- brous to puberulous; laminae ovate, 6-12 cm long, 2.5- 5 cm broad, apically acuminate, basally attenuate, mar- gins entire to undulate, sparingly and minutely strigose on both surfaces, mostly restricted to costa and veins, minute cystoliths abundant and apparent on both sur- faces. Inflorescences in terminal and axillary racemose panicles to 27 cm long, 8 cm broad; rachis and branches minutely puberulous plus much longer (to 1 mm), scat- tered glandular hairs, pedicels subsessile to 8 mm long, pubescence that of the rachis; bracts and bracteoles subu- late, 2-5 mm long, 1-2 mm broad, minutely puberulous plus a few longer glandular hairs. Flowers with 5-merous calyx, calyx segments narrowly linear or subulate, 4.5- 8 mm long, 0.75-1 mm broad, minutely puberulous plus scattered glandular hairs; corolla orange-red and yellow, 15-19 mm long, 1 .5 mm broad at base expanding abruptly to 6 mm broad above the calyx and narrowing to 5 mm broad at the mouth, sparingly puberulous, upper lip erect, 4 mm long, 3 mm broad, apically acute, lower lip 3-lobed, lobes rounded, 0.75 mm long, 1 mm broad; stamens exserted well beyond the tip of the upper lip. Fruits 44 FIELDIANA: BOTANY clavate, 13-15 mm long, 3.5 mm broad, 2.5 mm thick, glabrous; seeds ovate, flattened, 2.5 mm long, 2 mm broad, muricate. Found in moist forests on the slopes of the Cor- dilleras Central and Talamanca at elevations of 1500 to 1900 m. Flowering collections have been made in November, December, and February. Hansteinia stricta is recognized by its ovate leaves with attenuate bases and its open paniculate inflorescences with relatively few branches and these strict, erect, or ascending. It most closely resembles H. blepharorhachis, which has cymose panicles (thyrses), with narrower, more compact branching and larger bracts (5 to 10 mm long vs. 2 to 5 mm). lections have been made in October, December, January, and February. There is variation in the degree of hairiness among specimens from over the range of Han- steinia ventricosa. Early flowering specimens in particular may be confused with Razisea spicata, but can be distinguished from it with their shorter corollas (2.6 to 3 cm vs. 4 to 6.2 cm long), which expand more abruptly, and by the more profuse branching of the inflorescences. Hansteinia ven- tricosa resembles H. stricta and H. blepharo- rhachis somewhat, but can be distinguished from them by its broader laminae, with rounded, sub- cordate, or obtuse bases, as opposed to their nar- rowed laminae with attenuate bases. Hansteinia ventricosa (Donnell Smith) D. Gibson, Fieldiana, Bot. 34: 63. 1972. Glockeria ventri- cosa Donnell Smith, Bot. Gaz. (Crawfordsville) 27:439. 1899. Figure 6. Herbs to subshrubs to 3 m tall, older stems terete, younger stems subquadrangular ; internodes between leaf- bearing nodes 2-7 (1 1) cm long, 1.5-3 mm thick, sub- tomentose, hairs ca. 1 mm long. Leaves with petioles to 5.8 cm long, pilose to subtomentose; laminae ovate to ovate-elliptic, 7-15 cm long, 3-12 cm broad, apically acuminate, hasally subcordate to rounded or obtuse, margins crenulate to undulate, both surfaces pilose, cys- toliths minute. Inflorescences in terminal and axillary thyrses to 29 cm long, 1 1 cm broad excluding corollas, rachises and branches pilose plus glandular hairs to pu- berulous, pedicels to 9 mm long, puberulous-glandular; bracts and bracteoles oblong to narrowly lanceolate, to 9 mm long, 2 mm broad, apically acute to acuminate, puberulous to pilose plus occasional glandular hairs. Flowers with 5-merous calyx, segments linear to 7 mm long at flowering and to 1 2 mm long at fruiting stage, 1 mm broad, puberulous plus occasional glandular hairs; corolla orange, 2.6-3 cm long, tube curved outward, sparingly glandular-pilose, 2 mm broad basally, con- tracting just above the ovary and then expanding abrupt- ly to 5 mm broad just above the calyx lobes, the upper lip erect, elliptic, to 5 mm long, 2 mm broad, apically bidentate, lower lip ca. 1.5 mm long, 3-lobed, the lobes ca. 1 mm long, rounded; stamens exserted to ca. 1.5 cm beyond the tip of the upper lip. Fruits clavate, 11-19 mm long, 3-4 mm broad, 3—4 mm thick, glabrous; seeds 4, oblong, flattened, basally oblique, 2 mm long, 1 .5 mm broad, coarsely roughened. This is a comparatively wide-ranging species found in shade or partial shade of moist, evergreen forests of the Cordilleras Tilaran, Central, and Ta- lamanca and of adjacent areas in Panama at ele- vations between 1000 and 2500 m. Flowering col- Henrya Nees REFERENCE— G. B. Happ, Monograph ofTetra- merium and Henrya. Ann. Missouri Bot. Gard. 24: 501-582. 1937. Herbaceous to suffrutescent perennials; stems erect, spreading or decumbent, younger stems often pubescent with glandular hairs, tetrangular to subterete, with pale, often exfoliating, epidermis. Leaves petiolate, often de- ciduous; laminae rotund-ovate to linear, entire, pubes- cent to glabrous on both surfaces, cystoliths present. In- florescences in terminal and axillary spikes with loosely or densely imbricated bracts; cauline bracts solitary; in- volucral bracts coalescent from base to near the apex on the posterior side and closely enclosing the fruit, green, mucronate at or near the apex. Flowers sessile or short- pedicellate; calyx 5-merous; corolla narrowly funnel- form, bilabiate, tube ca. '/3 the length of the corolla, an- terior lip 3-lobed, the posterior lip shallowly bilobed or entire; stamens 2, exserted; anthers 2-celled, nearly par- allel, one cell longer than the other. Fruit a capsule, seeds 2, flattened, hispid to hispidulous on one side, glabrous on the other. Happ had recognized 20 species for this genus, ranging from the northern states of Mexico and adjacent United States to Costa Rica. He consid- ered the position of the mucro on the involucral bract to be a very important character in species designations. This has been questioned by some, and it appears that there are many fewer species than he designated. Henrya scorpioides Nees in DC., Prodr. 1 1: 491. 1847. Tetramerium scorpioides (Nees) Hems- ley, Biol. Centralbl. 2: 526. 1882. BURGER: FLORA COSTARICENSIS 45 Herbs to suffrutescent plants to 1 m tall; older steins terete, internodes between leaf-bearing nodes 2-4 cm long, 1-1.5 mm thick, puberulous. Leaves with petioles 1-1 5 mm long, puberulous; laminae ovate, 1-6 cm long, 0.5-3 cm broad, apically acute to acuminate, basally obtuse to acute, margins entire, puberulous-glandular on both surfaces, often glabrate with age, cystoliths prom- inent on both surfaces. Inflorescences 2-10 cm long, loosely imbricated cauline bracts linear-elliptic, elliptic or linear-oblanceolate, 2-4 mm long, 1-2 mm broad; involucral bracts oblanceolate, 7-9 mm long, glandular- puberulous, the mucro just below the apex. Usually 2 flowers per bract, only 1 producing a fruit, calyx seg- ments lanceolate, ca. 1.5 mm long; corolla pale yellow to cream-colored, 10-15 mm long. Fruits clavate, 4.5- 6 mm long, 1.5-2.5 mm broad, ca. 1 mm thick, pu- berulous; seeds 1 or 2, flattened, elliptic, ca. 1.5 mm long, 1.25 mm broad, margins hispidulous. This species ranges from Mexico to Costa Rica. Habitats have been described only as "rocky soil" or "rocky thorn forests." Only one collection from Costa Rica, M. Quiros 671 (F), is known. It was collected in flower at Desamparados de San Ma- teo, Alajuela Province, in March at 100- to 500-m elevation. Henrya scorpioides is recognized by its spicate inflorescences, with loosely imbricated bracts, its coalesced involucral bracts that are conspicuously glandular-puberulous and are 7 to 9 mm long, and its cauline bracts that are conspicuously shorter than the involucral bracts (2 to 4 mm long). This may be confused with Tetrameriwn nervosum, which differs in its more compact spikes with larg- er bracts that are four-ranked and lack the co- alesced involucral bract. Herpetacanthus Nees Herbs or shrubs. Inflorescences spicate; bracts loosely imbricate. Flowers with a 5-merous calyx, calyx seg- ments equal; corolla 5-merous, bilabiate, upper lip api- cally bidentate, lower lip 3-lobed; stamens 4, didyna- mous, inserted or barely exserted, the longer of each pair 2-celled with one of the cells superposed, the shorter of each pair 1 -celled. Capsules clavate; seeds 4. This is a genus of about 1 0 species, ranging from Central America to Brazil. Most of the species described are from Brazil. This genus is distin- guished from other genera in the family with bi- labiate corollas by its didynamous stamens, the longer pair 2-celled, with one of the cells distinctly superposed and the shorter pair of stamens one- celled. Herpetacanthus panamensis Leonard, J. Wash. Acad. Sci. 32: 185. 1942. Standleyacanthus cos- taricanus Leonard, Ceiba 3: 143. 1952. Figure 7. Herbs or subshrubs to 1 m tall; stems ascending, spar- ingly branched, rooting at lower nodes in some, younger stems quadrangular, puberulous to pilose mostly along 2 lines, hairs with a purplish or reddish banding pattern, older stems terete, glabrous; internodes between leaf- bearing nodes 2.5-4 cm long, 0.75-1 .25 mm thick. Leaves with short petioles to 1 cm long, pubescence that of the stem; laminae ovate to ovate-elliptic, 4-14.5 cm long, 2.5-6 cm broad, apically acute to acuminate, basally acute to attenuate, margins undulate, glabrous to strigose on the costa and veins of both surfaces, cystoliths dense and prominent on both surfaces. Inflorescences in lax terminal and axillary spikes or spicate panicles, spikes to 3.5 cm long, 2.5 cm broad; bracts loosely imbricate, ovate to elliptic, 9-14 mm long, 4-7 mm broad, apically acute to acuminate, glabrous or subglabrous, margins ciliate; bracteoles linear to narrowly elliptic, 6-8 mm long, 0.5-1 mm broad when fruits are mature, glabrous, margins sparingly ciliate. Flowers with 5-merous calyx, segments subulate, 3-4 mm long, 0.5 mm broad, mi- nutely glandular-puberulous inside, glabrous outside, tipped by 1 or 2 hairs; corolla white, to 1 7 mm long, the tube 1 1 mm long, 2 mm broad at base, 3.5 mm broad at the mouth, glabrous, the lips ca. 5.5-8 mm long, pu- berulous toward the tip, upper lip ovate, 4 mm broad, apically minutely bidentate, lower lip 3-lobed, lobes 2- 3 mm long, middle lobe rounded, 2.5 mm broad, lateral lobes acute, ca. 1.5 mm broad, the stamens exserted or included, filaments of the longer pair 3 mm long, of the shorter pair 2 mm long, glabrous, anthers ca. 1 mm long, 0.3 mm broad. Fruits clavate, 9-10 mm long, stipe 3.5- 4 mm long, 0.5 mm thick, seed-bearing portion 3 mm broad, 1.5 mm thick, glabrous; seeds 4, flattened, ovate, 2.5 mm long, 2.0 mm broad, roughened. Collected in wet, shaded areas of forests, fre- quently along streams from near sea level to ca. 800 m in Panama and Costa Rica. In Costa Rica most collections have been made in Alajuela, Her- edia, and Limon provinces of the Caribbean wa- tershed area, but one, Allen 6297 (F, CR) is known from Esquinas forest of Puntarenas Province. Flowering collections in Costa Rica have been made in March, July, September, and October. The September collection, Allen 6297 (F), was from an area which has a more extended dry season than the others. One collection, Opler 119 (F), de- scribes the bracts as purplish. This specimen has, as have most others, dark purplish green bracts when dried; some, however, such as Schnell 810 (F), collected inland from Puerto Viejo in Limon Province, have yellowish green bracts. This vari- ation may be regional, or it may be an age differ- ence, since the Schnell collection was in an early flowering stage with no fruits. 46 FIELDIANA: BOTANY Herpetacanthus panamensis may be confused with various of the larger-bracted species of Jus- ticia, but can be easily distinguished from them, with its second pair of stamens that are single- celled (vs. two, two-celled stamens, with cells that are frequently superposed). If stamens are missing, the loosely imbricate bracts that are frequently purplish are the best distinction. Hygrophila R. Brown Erect or diffuse herbs. Leaves opposite. Flowers in axillary fascicles; bracts linear; calyx 5-merous, segments subequal; corolla tubes subcylindric, small, the limb 2-lipped; stamens 4, didynamous, or 1 perfect stamen and a staminode in each pair, the filaments in each pair, united at the base; anthers 2-celled, some mucronulate; posterior stigma lobe abortive. Fruit an oblong, non- stipitate capsule; seeds 4-18 or more. This genus is pantropical, with about 80 species. Hygrophila has some resemblance to the genus Dyschoriste, but can be distinguished from it in Central America by its white flowers (vs. blue or purple) and capsules with numerous seeds (14 to 48 vs. two to four). Hygrophila costata Nees, PI. hort. bonn. icon. 2: 7. t. 3, 1824, and in DC, Prodr. 11: 88. 1847. H. guianensis Nees, London J. Bot. 4: 634. 1 845. H. conferta Nees in Mart., Fl. bras. 9: 21. 1847. Figure 7. Herbs to 1.5 m tall, sometimes declining and rooting at lower nodes, steins obtusely quadrangular; internodes between leaf-bearing nodes 2-7 (10) cm long, 1-3 mm thick, glabrous. Leaves petiolate to subsessile, petioles to 2.5 cm long, glabrous; laminae lanceolate to elliptic, 3.5-18 cm long, 0.5-5 cm broad, apically acuminate, basally attenuate, margins entire to undulate, cystoliths very numerous on the upper epidermis, less numerous on the lower, glabrous above, sparingly puberulous be- low, mostly restricted to costa and veins. Flowers sessile, often in clusters of 3 or 4, bracts lanceolate-elliptic, 6- 9 mm long, 1.5-3 mm broad, sparingly puberulous, brac- teoles similar, 2-3 mm long, ca. 0.5 mm broad, calyx segments linear, equal, 5-9 mm long at anthesis, 6.5-12 mm long in fruit, sparingly puberulous; corolla white, sometimes streaked or spotted with purple, 5.5-9 mm long, tube ca. 1 .5 mm broad at base, expanding to 2 mm broad at the throat, puberulous, 2-lipped, lips ca. 3 mm long, upper lip 2-lobed, lower lip 3-lobed; stamens 4, didynamous, filaments of each pair united at the base, anthers 2-celled, ca. 1 mm long, cells basally mucron- ulate; fruits narrowly oblong, sessile, glabrous, 8-1 3 mm long, 6-8 seeds per locule. This is a wide-ranging plant found from Mexico to central Argentina in moist habitats. In Costa Rica it has been collected along streams, rivers, forested thickets, weedy roadsides, open pastures, and rain forests from sea level of the Osa Peninsula to mountain slopes of the Cordilleras Talamanca and Central at 1 500 m. Hygrophila guianensis and H. costata have been distinguished by leaf shape and size, length of ca- lyx, corolla, and capsule. I have brought H. gui- anensis into synonymy because the numerous measurements of all these characters in Costa Ri- can collections intergrade and overlap. Further- more, they show no habitat differences. This is a common and wide-ranging complex, which on the basis of these observations, appears to be one species. Hygrophila costata is recognized by its sessile flowers often in clusters of three or four, its in- conspicuous, white, bilabiate corollas with four didynamous stamens with two-celled, basally mu- cronulate anthers, and its oblong, sessile capsule with 1 2 to 16 seeds. Hypoestes Soland. REFERENCE— W. T. Thiselton-Dyer, Flora of Central Africa 5: 244. 1900. Herbs or shrubs. Leaves petiolate. Inflorescences in axillary and/or terminal spikes or panicles; floral leaves resembling reduced vegetative leaves often present; bracts 2 per flower, free or coalesced for '/3-% of their length. Flowers with 5-merous calyx, calyx lobes equal, subu- late, free or coalesced 'A-% of their length; corolla bila- biate, tube narrow, gradually expanding to the mouth, upper lip 2-lobed, lower lip 3-lobed; stamens 2, anthers 1 -celled, basally rounded. Fruits clavate; seeds 2 or 4. This is a genus of the Paleotropics, with 1 50 species extending from Africa to Australia. Most species are found in Madagascar. Hypoestes phyllostachya Baker, J. Linn. Soc., Bot. 22: 511. 1887. Figure 7. Herbs to 70 cm tall, young basal shoots especially velutinous; internodes between leaf-bearing nodes 2-6 cm long, 1.25-2.0 mm thick, subtetrangular, glandular plus appressed hairs to appressed hairs at angles only. Leaves petiolate, petioles to 2.5 cm long, appressed hairs on upper surface only; laminae ovate, 2-4.5 cm long, 1.5-2.5 cm broad, apically acute, basally obtuse to BURGER: FLORA COSTARICENSIS 47 rounded, margins entire, glabrous to sparingly puberu- lous on the midrib of both surfaces, most surfaces dotted with numerous pink spots to ca. 4 mm in size, cystoliths present on both surfaces. Flowers borne singly and in terminal and axillary spikes, often with a gradual tran- sition from solitary flowers to spikes, rachises glandular- puberulous; bracts 2 per flower, fused for up to half their length, oblong, 12-15 mm long, 1.4-1.5 mm broad, api- cally acute to obtuse, with one bract slightly longer than the other, velutinous plus glandular hairs, bracteoles su- bulate, apically tapered, 10-11 mm long, 1.2-1.3 mm broad, velutinous; flowers sessile. Flowers with a 5-mer- ous calyx, 6-7 mm long, segments equal, subulate, fused for ca. half their length; corolla magenta, 24-25 mm long, ca. 3 mm broad at the mouth, 0.7 mm broad basally, puberulous, upper lip 1 1-12.5 mm long, 3-lobed, lobes rounded, ca. 1 mm long, ca. 3.5 mm broad, lower lip 9.5-1 1 mm long, ca. 1.5 mm broad; stamens exserted, anthers 1-celled, ca. 1.7 mm long, 0.6 mm broad. Fruits clavate, 10-11 mm long, 1.4-1. 6 mm broad, 1.6-1. 7 mm thick, puberulous toward apex; seeds 4, oval, 1.6-1.8 mm long, 1.1-1.2 mm broad, papillate. This species is native to tropical Africa, but is often cultivated for its attractive foliage. Escapes from cultivation have been made along road banks in two cloud forest areas in Costa Rica, the Mon- teverde community at 1 500 to 2000 m elevation in Puntarenas Province and near San Miguel at about 1600 m elevation in Heredia Province. Flowering collections were made in April and De- cember. Hypoestes phyllostachya is most easily recog- nized by its pink-dotted leaves and magenta co- rolla, with a three-lobed upper lip and one-lobed lower lip. It is not easily confused with other Cen- tral American species of Acanthaceae. Justicia Linnaeus Herbs, shrubs, or vines. Leaves petiolate, laminae usu- ally ovate to elliptic and entire, cystoliths present and usually prominent on both surfaces of dried specimens. Inflorescences usually of spikes or panicles, occasionally in cymes or solitary; bracts various, small and distant, linear or subulate to large, conspicuous and imbricate. Flowers with calyx segments usually narrow and equal, some unequal, mostly 5(4)-merous; corollas bilabiate, usually white, pink, or purple, the tube usually narrow, funnelform, the upper lip 2-lobed, the lower lip 3-lobed; stamens 2, usually slightly exserted, but not exceeding the upper lip, anther cells 2, usually unequal, often su- perposed, with 1 or both cells apiculate or calcarate, the connective narrow to broad. Fruits a capsule, clavate, 4- seeded. This is the largest and most complex genus of the Acanthaceae, with approximately 420 species found mostly in the tropics, but extending well into temperate habitats. It is worldwide in distri- bution. The genus as treated here includes what, until recently, was recognized as Beloperone and Jacobinia. The Costa Rican species of Justicia are wide ranging, from near sea level to just above 2000 m elevation. Most species are found between 500 and 2000 m in moist habitats around openings or along trails in forests. Three species are found in dis- turbed habitats. About five species are cultivated. Justicia may be recognized by its two slightly exserted stamens with two-celled anthers that are frequently unequal, superposed, and basally apic- ulate and calcarate. In addition, the bilabiate, nar- row-tubed corollas and the lack of staminodes are useful features. Key to Species of Justicia la. Inflorescences in spikes or cymes 2a Ib. Inflorescences in panicles or thyrses (cymose panicles) 23a 2a. Inflorescences in axillary cymes 3a 2b. Inflorescences in spikes borne singly or in clusters 4a 3a. Leaves of a pair unequal; corolla violet or pale purple J. trichotoma 3b. Leaves of a pair equal; corolla orange-red to yellowish J. macrantha 4a. Leaves apically obtuse to rounded J. chamaephyton 4b. Leaves apically acuminate or acute 5a 5a. Bracts 4-19 mm long, imbricate 6a 5b. Bracts 1-3.5 mm long, lax (not imbricate) 20a 6a. Bracts spatulate; calyx segments 8-1 1 mm long 7a 6b. Bracts otherwise; calyx segments 4-7 mm long 8a 7a. Corolla 2.4-3.5 cm long; plant growing in open, disturbed habitats J. carthaginensis 7b. Corolla 1.1-1.2 cm long; plant growing in shade of moist forests J. costaricana 48 FIELDIANA: BOTANY 8a. Bracts reddish brown, 16-19 mm long; plants cultivated J. brandegeana 8b. Bracts green or brown, 4-1 1 mm long; plants not cultivated 9a 9a. Capsules 1 3-1 5 mm long; corollas 20-40 mm long 10a 9b. Capsules 5-10 mm long; corollas 7-19 mm long 1 la lOa. Corollas white; bracts small, inconspicuous, 4-6 mm long, linear-subulate J. urophylla lOb. Corollas rose-purple; bract larger, 10-11 mm long, oblanceolate . . . J. brenesii \ la. Leaf bases rounded to subcordate; leaves sessile to subsessile J. skutchii 1 Ib. Leaf bases acuminate, acute or obtuse; leaves with petioles mostly more than 5 mm long (except J. refractifolid) 1 2a 12a. Calyx glandular-puberulous; lower cell of anther acute or rounded 13a 12b. Calyx puberulous or glabrous; lower cell of anther spurred 17a 1 3a. Bracts of inflorescence 4-6 mm broad; peduncles lacking glandular hairs J. valerii 13b. Bracts of inflorescence 0.5-2.5 mm broad; peduncles with glandular hairs . 14a 14a. Corollas 15-19 mm long; calyx segments 5-7 mm long ..../. angustibracteata 14b. Corollas 7-1 1 mm long; calyx segments 2-4.5 mm long 1 5a 1 5a. Calyx segments (5) equal; spike lax J. tonduzii 15b. Calyx segments not equal (posterior calyx segment setaceous); spike compact 16a 1 6a. Bracts apically obtuse; leaves narrowly elliptical oblong J. parvibracteata 1 6b. Bracts apically acute; leaves lanceolate-elliptic J. pittieri 17a. Leaves no longer than 6.5 cm; posterior calyx segment much shorter than the rest . . J. candelariae 17b. Leaves to as long as 18 cm; calyx segments of equal lengths 18a 18a. Bracts obovate to elliptic, strigose; corolla 16-17 mm long /. orosiensis 18b. Bracts ovate, linear-spatulate, spatulate or orbicular, pilose or glandular-puberulous; corolla 7-1 2 mm long 1 9a 19a. Bracts linear-spatulate, spatulate or orbicular, glandular-puberulous; petioles glabrous J. costaricana 1 9b. Bracts ovate, pilose; petioles pilose to tomentose /. refractifolia 20a. Leaves with auriculate bases, sessile to subsessile /. ephemera 20b. Leaves basally attenuate (lacking auricles), petiolate 2 la 2 la. Corolla 12-19 mm long, anther cells superposed; calyx segments 4 J. metallica 21b. Corolla 30-44 mm long, anther cells subequal; calyx segments 5 22a 22a. Leaves elliptic; corollas narrowly funnelform (not fusiform when unopened) . . . J. tinctoria 22b. Leaves ovate; corollas fusiform when unopened J. spicigera 23a. Capsules 4-6 mm long; corollas 4-6 mm long J. comata 23b. Capsules 7-25 mm long; corollas 8-70 mm long 24a 24a. Plant a climbing vine; corollas 65-70 mm long J. sarapiquensis 24b. Erect herbs or shrubs; corollas 8-60 mm long 25a 25a. Shrubs 2-6 m tall; anther cells equal or subequal, but not superposed 30a 25b. Herbs mostly less than 1 .5 m tall; anther cells superposed 26a 26a. Posterior calyx segment shorter, ca. half the length of the others; corollas 25-35 mm long J. oerstedii 26b. Calyx segments of equal lengths; corollas 8-17 mm long 27a 27a. Inflorescence a compact spicate panicle resembling a single spike; bracts ovate to elliptic, 6- 8 mm long, 2.5-3.5 mm broad /. orosiensis 27b. Inflorescence a loose spicate panicle; bracts subulate to linear-subulate, 1 .5-2 mm long, 0.25- 1 mm broad 28a 28a. Leaves with auriculate bases, sessile to subsessile J. ephemera 28b. Leaves basally attenuate to rounded, lacking auricles, petiolate 29a 29a. Calyx segments 4, ca. 5 mm long; leaves ovate, 1.5-4 cm broad J. metallica 29b. Calyx segments 5, ca. 2 mm long; leaves lanceolate, 1-2 cm broad /. pectoralis BURGER: FLORA COSTARICENSIS 49 30a. 30b. 33a. Bracts subulate, 1-3 mm long; petioles short, 1 5 mm or less 3 la Bracts linear-lanceolate, elliptic or obovate, 5-14 mm long; petioles long to more than 50 mm long 33a 3 la. Flowers in axillary cymes; capsules 18-20 mm long /. macrantha 3 1 b. Flowers in loose spicate panicles; capsules 7-8 mm long 32a 32a. Leaves elliptic; corollas narrowly funnelform (not fusiform when unopened) . . . J. tinctoria 32b. Leaves ovate; corollas fusiform when unopened J. spicigera Inflorescences in large, terminal spicate panicles, corollas red with yellow tip; capsules ca. 1.5 cm long J. crenata 33b. Inflorescences in dense terminal or subterminal thyrses (cymose panicles), corollas yellow; capsules 2-2.5 cm long J. aurea Justicia angustibracteata Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1227. 1938. Rhy- tiglossa microphylla Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 159. 1 855, non Justicia microphylla Lam., 1 79 1 . Fig- ure 7. Herb to 1 m tall, young stem quadrangular; internodes between leaf-bearing nodes 1.5-4.5 cm long, 1.0 mm thick, puberulous along 2 lines. Leaves with puberulous to subglabrous petioles to 1 cm long; laminae ovate- lanceolate, 1.5 (2)-5 (6) cm long, 1-1.5 (2) cm broad, apically acuminate, basally acute, margins undulate, gla- brous to sparingly and minutely scabrous above, more so on costa and veins, puberulous below. Inflorescences in lax terminal spikes to 6 cm long, 1 cm broad excluding corollas; peduncle quadrangular, to 2 cm long, glandular- puberulous, rachis quadrangular, glandular-puberulous; bracts imbricate, oblanceolate, 10-11 mm long, 2-2.5 mm broad, ciliate, sparingly puberulous, bracteoles lin- ear, 6.5-8.5 mm long, 0.5-0.75 mm broad, glandular- puberulous. Flowers with 5-merous calyx, calyx seg- ments unequal, linear subulate, the posterior segment much smaller, ca. 3.5 mm long, 0.25 mm broad, the other segments 5-6.5 mm long, 0.5-0.75 mm broad, sparingly puberulous-glandular; corolla white with pur- plish markings, 15-19 mm long, 2-2.5 mm broad at base, ca. 3.5 mm broad at the throat, upper lip erect, 6- 7 mm long, 5-6 mm broad at base, narrowing to a keeled, rounded apex, lower lip 6.5-7 mm long, 6-7 mm broad at base, 3-lobed, the lobes ca. 2.5 mm long, semiorbic- ular, stamens exserted, but not beyond the upper lip, anthers strongly superposed, anthers with relatively wide separation, connective 0.5 mm long, anthers 1.0 mm long, 0.5 mm broad, apically and basally acute. Fruits clavate, 5-7.5 mm long, 2.25 mm broad, 1-1.5 mm thick, puberulous; seeds 2, flat, ovoid, dark brown, 1.5 mm long, 1.25 mm broad. Found along streams and on steep slopes through cloud forest areas of Cartago and San Jose prov- inces at elevations of 1800 to 2000 m. Flowering collections have been made in January, February, March, and June. Justicia angustibracteata is recognized by its ob- lanceolate bracts, unequally segmented, five-mer- ous calyx, and white corollas with purplish mark- ings. This species is not easily confused with other Justicia species when all of the above features are visible. It bears a superficial resemblance to J. par- vibracteata, but differs from it in its unequal calyx segments, its longer (10 to 1 1 mm vs. 4 to 6 mm), oblanceolate bracts (vs. elliptic) and its longer co- rollas (15 to 19 mm vs. 6.5 to 7.5 mm). Justicia aurea Schlecht., Linnaea 7: 393. 1832, non Justicia aurea (Rose) Lindau, Bull. Herb. Boissier. 5: 675. 1897. Justicia umbrosa Benth., PI. hartw. 79. 1 84 1 . Adhatoda umbrosa Nees in DC., Prodr. 11: 406. 1847. Jacobinia aurea Hemsl., Diagn. pi. nov. mexic. 35. 1879, non Hiern, 1877-1878. Ecbolium umbrosum Kuntze, Revis. gen. pi. 2: 981. 1891. Jacobinia umbrosa Blake, Contr. Gray Herb. 52: 103. 1917. Fig- ure 7. A shrub to 6 m tall, stem subquadrangular to qua- drangular in younger portions; internodes between leaf- bearing nodes 2-5 cm long, 4-7 mm thick, puberulous to tomentose. Leaves with petioles to 5.7 cm long, pu- bescence that of the stem; laminae ovate to ovate-elliptic, 11.5-30 (40) cm long, 5.5-12 (17) cm broad, apically acuminate, basally attenuate, margins entire, undulate or crenulate, glabrous to sparingly strigose above, hairs more dense on costa, glabrous to strigose below, cysto- liths dense and prominent on both surfaces. Inflores- cences densely thyrsoid at or near the ends of branches, to 18 cm long, 6 cm thick excluding corollas; bracts linear-lanceolate, 1 1-14 mm long, 1-2 mm broad; brac- teoles similar, but slightly smaller, both puberulous. Flowers with 5-merous calyx, segments lanceolate, 5.5- 7 mm long, 1.5 mm broad at base, puberulous; corolla yellow, bilabiate, cylindrical, to 5.5 cm long, 3 mm broad, puberulous-glandular outside, glabrous inside except for bearded throat, the lips ca. 2.5 cm long, upper lip erect, 7 mm broad, apically acute, lower lip 6 mm broad, api- cally 3-lobed, lobes ca. 1 mm long, apically rounded and slightly hooded, especially the middle lobe; stamens 50 FIELDIANA: BOTANY exserted to ca. the tip of the upper lip, anthers 3 mm long, glabrous, the cells unequal, basally acute, filaments stout, ca. 0.75 mm broad, glabrous; ovary 2 mm long, 1 mm thick, glabrous. Fruit clavate, puberulous, 2-2.5 cm long, ca. 5 mm broad, 4 mm thick; seeds 4, flattened, suborbicular, ca. 2 mm in diameter, surface roughened. This is a very attractive shrub, and it is fre- quently cultivated. In the wild it is a relatively wide-ranging species and is found from southern Mexico to Panama. In Costa Rica it has been col- lected in all provinces except Limon and Cartago, generally in and adjacent to rain and cloud forest from elevations of 100 to 2000 m. Flowering collections have been made in all months except May and August. Justicia aurea is recognized by its large, attrac- tive thyrsoid inflorescences with large, yellow co- rollas (more than 5 cm long), its shrubby habit, and its large leaves with long laminae (to 30 cm or more long). It is not easily confused with other species of Justicia. Justicia brandegeana Wassh. & Smith, Fl. Ilus. Catarinense ACAN:102. 1969; Beloperone guttata Brandegee, Univ. Calif. Publ. Bot. 4: 278. 1912; non Justicia guttata Wall., PI. asiat. rar. 1: 24. pi. 28. 1830. Figure 9. Herb to subshrub, 1-2 m tall, steins subquadrangular; in U- modes between leaf- bearing nodes 6-9.5 cm long, 1.5-2.5 mm thick, strigose mostly along 2 lines in older portions. Leaves with densely strigose petioles 0.5-1 cm long; laminae ovate-lanceolate, 3-3.5 cm long, 1-1.5 cm broad, margins entire, apically acute to slightly attenuate, apex blunt, basally acute, densely pilose on both surfaces, cystoliths visible on both surfaces although obscured by hairs. Inflorescences in terminal and axillary spikes to 6 cm long, 2 cm broad, excluding corollas; bracts reddish brown, imbricate, ovate, 16-19 mm long, 11-13 mm broad, apically acute, basally rounded to subcordate, puberulous, ciliolate, bracteoles ovate-elliptic, 12-14 mm long, 5-6 mm broad, apically acute, basally obtuse, pu- berulous with ciliolate margins, peduncles 5-10 mm long, densely strigose. Flowers with 5-merous calyx, segments subulate, ca. 5 mm long, posterior segment 1.25 mm broad basally, others ca. 0.75 mm broad, all densely puberulous; corolla white, 3 longitudinal rows of pur- plish dots in throat, 3.2 cm long, the tube narrow, 2 mm broad at base expanding to only 3.5 mm at the mouth, bilabiate, upper lip erect, 1 2 mm* long, 5 mm broad at base, apically acute, lower lip ca. 1 1 mm long, 2-3 mm broad, apex of 3 shallow, close, obtuse lobes; stamens exserted to the tip of the upper lip, anther cells unequal, the cells superposed ca. 3 mm long, basally spurred, filaments glabrous. Fruits clavate, 1 2 mm long, 4 mm broad, 3.5 mm thick, puberulous; seeds 4, dark brown, subglobose, 3 mm long, 2.5 mm broad, 0.5 mm thick, glabrous. This plant is native to Mexico and is commonly cultivated in much of Latin America. Justicia brandegeana is recognized by its at- tractive red bracts. These rather compact inflo- rescences of reddish bracts give the plant its com- mon name, the shrimp plant. In Costa Rica it is called camaroncillo. Justicia brenesii (Leonard) D. Gibson, Fieldiana, Bot. 34: 66. 1972. Beloperone brenesii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1201- 1202. 1938. Figure 8. Herbs to 1 m tall, erect or decumbent with branches frequent, occasionally rooting at lower nodes, younger stems quadrangular with dense brownish pilose hairs to 1 mm long at the angles, older stems terete and glabrous; intern (»d os between leaf-bearing nodes, 2-9 cm long, 1 .5- 2.5 mm thick. Leaves with petioles to 2.2 cm long, pilose; laminae ovate-elliptic, 3.5-6.5 cm long, 2-3.5 cm broad, apically short-acuminate, basally acute, margins entire with pilose hairs, both surfaces glabrous to sparingly pilose, cystoliths prominent and dense on both surfaces. Inflorescences in dense terminal or lateral spikes or spikelike thyrses to 5 cm long, 2.5 cm broad excluding corollas; bracts imbricate, oblanceolate, 10-11 mm long, 3.5-4 mm broad, pilose and ciliate; bracteoles of similar shape and length, but 1-2 mm broad, pilose and ciliate. Flowers with 5-merous calyx, segments subulate, all ca. 6 mm long, posterior segment 1.5 mm broad at base, lateral segments 1.25 mm broad, anterior segments 1 mm broad, all ciliolate; corolla rose-purple, to 2.5 cm long, 2.5 mm broad at base expanding to 5.5 mm broad at the mouth, glandular-puberulous, the lips ca. 10 mm long, the upper lip erect, apically acute and notched, lower lip 3-lobed, lobes ca. 3 mm long, apically obtuse; the stamens exserted to just below the tip of the upper lip, anther cells superposed, upper cells ca. 2 mm long, lower cells slightly longer, all cells basally calcarate. Fruits clavate, capsules ca. 13 mm long, 4 mm broad, 2.5 mm thick, puberulous, hairs retrorse, seeds not seen. Found in or adjacent to wet forest areas from 800 to 2200 m elevation. This is apparently an endemic species collected in Alajuela and Heredia provinces on both Caribbean and Pacific wa- tershed zones. Flowering collections have been made in September through April, except March. Justicia brenesii is recognized by its stems with densely pilose angles, its relatively broad, short, and crowded spikes or thyrsi with pilose and ciliate bracts, and its rose-purple corollas. It has some resemblance to J. carthaginensis, but differs from it in its generally smaller laminae, long-pilose stems, and pilose and ciliate bracts. Justicia candelariae (Oerst.) Leonard, Carnegie Inst. Wash. Publ. 461: 231. 1936. Rhytiglossa BURGER: FLORA COSTARICENSIS 51 candelariae Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjebenhavn 1854: 158. 1855. Dianthera candelariae Hemsl., Biol. Centralbl. 2:517. 1882. Figures. Herbs to 45 cm tall, steins frequently decumbent, often rooting at the lower nodes, quadrangular, pilose, densely so in younger portions; internodes between leaf-bearing nodes, 2.5 cm long, 1-1.5 mm thick. Leaves with short petioles 3-10 mm long, pilose; laminae ovate-elliptic to elliptic, 2.5-6.5 cm long, 1-2 cm broad, apically and basal ly acute, margins entire, glabrous to sparingly pilose above, sparingly pilose beneath, cystoliths visible on both surfaces, more prominent beneath. Inflorescences in ter- minal and axillary spikes to 7 cm long, 1.3 cm broad excluding corollas, peduncles to 2.2 cm long, pilose; bracts imbricate, spreading outward at an thesis, elliptic, 6.5-9 mm long, 3.5-5 mm broad, apically obtuse and mu- cronulate, basally acute, vein pattern prominent, pilose and ciliate; bracteoles narrowly elliptic, ca. 6 mm long, 1.5 mm broad, pilose and ciliate. Flowers with 5-merous calyx, calyx segments unequal, segments narrowly subu- late, posterior segment 2.5-3 mm long, 0.25 mm broad, all others 4-6 mm long, 0.5-0.75 mm broad, ciliolate; corollas white to pale purple, ca. 10 mm long, 2 mm broad at base expanding to 2.5 mm at the throat, gla- brous, upper lip acute, 3.5 mm long, 2.5 mm broad at base, lower lip 4.5 mm long, 3-lobed, lobes 2 mm long, middle lobe 2.75 mm broad, lateral lobes ca. 2 mm broad, all apically rounded; stamens barely exserted, an- ther cells unequal, ca. 0.5 mm long, upper cell slightly larger, lower cell calcarate. Fruits clavate, capsules 7 mm long, 2 mm broad, 1.5 mm thick, puberulous; seeds sub- orbicular, 1.2 mm in diameter, papillate. Herbs or suffrutescent plants to 1.5 m tall, stems te- trangular and grooved in younger portions, sparingly strigose mostly along grooves (sometimes along 2 lines) in younger stems, glabrous in older stems; internodes between leaf-bearing nodes, 2-7 cm long, 1-3 mm thick. Leaves with petioles 0.5-2.5 cm long, puberulous; lam- inae ovate to elliptic-ovate, 3-7 (14) cm long, 1.5-4(6.5) cm broad, apically acuminate, basally attenuate, margins entire, glabrous to strigose on the veins on both surfaces, cystoliths abundant and prominent on both surfaces. Inflorescences in terminal and axillary spikes, frequently in clusters of 2 or 3, spikes 2-5.5 cm long, 2-3 cm broad excluding corollas, peduncles short, frequently less than 1 cm long, puberulous; bracts imbricate, oblong-spatu- late to spatulate, sizes varying from 10 mm long, 4 mm broad near apex to 15(18) mm long, 7 (8) mm broad at the base of the inflorescence, apiculate, puberulous with glandular hairs toward the apex, margins ciliate; brac- teoles similar, but slightly smaller. Flowers sessile, calyx with 5 equal segments, segments linear-lanceolate, 8-15 mm long, 1-1.5 mm broad, puberulous, margins white, ciliate; corolla rose-purple, 24-35 mm long, sparsely and minutely puberulous outside, ca. 3 mm broad basally, expanding to ca. 5 mm broad at the mouth, tube bearded at 4 points inside just above the ovary, upper lip erect, 1 5-20 mm long, notched, lower lip 3-lobed, 1 5-20 mm long, middle lobe ca. 7.5 mm broad, lateral lobes ca. 5.5 mm broad, middle lobe personate; stamens exserted ca. 10 mm beyond the mouth of the corolla tube, filaments glabrous, anther cells 2.5-3 mm long, superposed, upper cell 1.5 mm above the lower, lower cell calcarate at base, upper cell blunt. Fruit clavate, capsules 13-17 mm long, 5-6 mm broad, 4 mm thick, apically apiculate, puber- ulous; seeds very dark brown, shiny, suborbicular, 3-3.5 mm in diameter. Collected in moist forests including oak forests at elevations of 1200 to 2000 m in provinces of Cartago and adjacent parts of San Jose. Flowering collections have been made in November, Decem- ber, January, and April. Justicia candelariae is recognized by its pilose stems, elliptic, subreticulate bracts with ciliate margins, relatively small, inconspicuous corollas ( 1 cm long or less), and its unequal calyx segments. It bears the same common Spanish name Zornia as Blechum brownei, which it resembles superfi- cally, but B. brownei is found at lower elevations, has a near regular corolla, and has four stamens instead of two. It also has some resemblance in its inflorescence to Justicia parvibracteata, which has much longer and narrower leaves, smaller bracts, and a setaceous posterior calyx segment. Justicia carthaginensis Jacq., Enum. syst. pi. 11. 1760, and in L., Syst. veg. ed. 16: 2: 153. 1817. Adhatoda carthaginensis (Jacq.) Nees in DC., Prodr. 11: 403. 1847. Figure 8. Found at lower elevations (0 to 100 m) of Gua- nacaste and Puntarenas provinces in and adjacent to tropical dry forests, frequently in secondary growth and along roadsides. This plant is found over a wide range from Mexico to South America. It is abundant where found and occurs most often in disturbed habitats. It is most frequently found at lower elevations, but has been collected from as high as 1300 m in some areas, although not above 100 m in Costa Rica. This species exhibits considerable variation in such features as corolla length as well as size, shape, and hairiness of bracts, leaves, and calyx. None of these variations is suf- ficiently consistent to warrant the designation of new species or varieties. Justicia carthaginensis is recognized by its dense spikes, conspicuous, large (13 to 17 mm long), broad, clavate capsules. Because it is weedy also, it has been confused with Blechum brownei, but can be easily distinguished from it in number of stamens (two vs. four), larger, bilabiate corolla, bract shape (oblong-spatulate vs. ovate), and much larger capsule (13 to 17 mm vs. 6 mm). 52 FIELDIANA: BOTANY Justicia chamaephyton D. Gibson, Fieldiana, Hot. 34: 67. 1972. Beloperone variegata Lindau, An- ales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim. fl. costaric. 2: 315. 1900. Figure 8. Herbs to 30 cm tall, stems ascending, rooting at lower nodes, younger portion quadrangular, densely puberu- lous, the hairs retrorse; internodes between leaf-bearing nodes, 2.5-4 cm long, 1.5-2.5 mm thick, puberulous. Leaves subsessile to petioles 1 cm long, petioles puber- ulous; laminae elliptic, 5-12 (15) cm long, 2.5-5 (6.5) cm broad, apically obtuse to rounded, basally attenuate, margins undulate, glabrous above, puberulous on costa and veins below, drying to a lighter green along the veins above, cystoliths abundant and conspicuous on both sur- faces. Inflorescences in loose terminal spikes to 8.5 cm long, 4 mm broad (excluding corollas), peduncle to 3 cm long, puberulous, rachis puberulous; bracts subulate, 2.5- 6 (7) mm long, 1 mm broad, puberulous, bracteoles sim- ilar, but ca. 2 mm long, 0.5 mm broad. Flowers 1 per bract; calyx 4-merous, segments subequal, 5-6 mm long, 0.5 mm broad, puberulous; corolla white or cream col- ored suffused with yellow-green markings, ca. 10 mm long, the tube narrow, 1 mm broad at base, 1.5 mm broad at the throat, puberulous, the upper lip 5-6 mm long, 3 mm broad, minutely bidentate at the tip, the lower lip 5 mm long, middle lobe ca. 2.5 mm broad, lateral lobes 2.5 mm long, 2 mm broad; stamens exserted to just beyond the tip of the upper lip, anther cells su- perposed, the lower calcarate. Fruits clavate, 10-1 1 mm long, puberulous; seeds flat, glabrous, 2.5 mm in di- ameter. Collected in and along the margins of moist pri- mary forests from the lowlands of Osa Peninsula and the Golfo Dulce area at near sea level to the Boruca area at ca. 300 m elevation, all in Puntar- enas Province. This species is also found in low- land areas of both the Pacific and Caribbean sides of Panama, ranging from the Costa Rican border to near Colombia. Justicia chamaephyton is recognized by its small size, slender, lax spikes, subulate bracts, and lam- inae with obtuse or rounded bases and apices. Su- perficially, J. chamaephyton resembles Aphelan- dra tonduzii, but in addition to its different flowers, it can be distinguished from it by its obtuse or rounded leaf bases. 3.2-7 cm long, 2-2.5 mm thick. Leaves sessile; laminae somewhat variable, ovate, elliptic or elliptic-oblong, 3.5- 10 cm long, 2.2-3.5 cm broad, apically acuminate, ba- sally acute, rounded or obtuse, margins entire to slightly undulate, glabrous to sparingly strigose along the costa, cystoliths dense on both surfaces, especially so above. Inflorescences in terminal and axillary spicate panicles, the spikes whorled at the rachis nodes, the flowers secund on the peduncle, the rachis and the branches grooved with the branches generally somewhat flattened; rachis and branches puberulous with a few longer glandular hairs; bracts subulate, ca. 2 mm long, 0.5 mm broad, keeled; bracteoles similar, slightly shorter. Flowers ses- sile with 5-merous calyx, segments equal, lanceolate, 1.5-2 mm long, 0.5 mm broad at the base, glabrous; corolla bilabiate, white, purplish, or white with purplish dots or streaks, 4-6 mm long, 0.8 mm broad at base expanding to 1.5 mm broad just above the base, gla- brous, tube 3 mm long, the upper lip 3 mm long, 1.8 mm broad at base, apically acute, the lower lip 4 mm long, 1.5 mm broad at base, expanding to 3 mm broad above, 3-lobed at the tip, the lobes about equal, 0.5 mm long, 1 mm broad, apically obtuse; stamens exserted ca. 2 mm beyond the corolla mouth, anther cells superposed, ca. 0.3 mm long, filaments pilose toward base, glabrous above. Fruits clavate, 4-5 mm long, 1.5 mm broad, 1 mm thick, puberulous; seeds 4, reddish brown, subor- bicular, flattened, ca. 1 mm in diameter, papillose. This is a species found in various disturbed hab- itats, stream banks, cutover hills, roadsides and along trails. It occurs throughout tropical America and is generally found in lower elevations below 500 m, although one collection (Standley & Val- erio 51873 F), has been made at La Hondura, San Jose Province at 1200 to 1500 m. It has been collected from all provinces in Costa Rica. Flow- ering collections have been made in all months except June, August, and September. Justicia comata is recognized by its spicate pan- icles that are both terminal and axillary, its small corollas (4 to 6 mm long) and capsules (4 to 5 mm long). It is most easily confused with J. pectoralis, which has only terminal spicate panicles, longer corollas (8 to 12 mm long), and capsules (7 to 8 mm long). Justicia costaricana Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1229. 1938. Figure 8. Justicia comata (L.) Lam., Encycl. 1: 632. 1783. Dianthera comata L., Syst. nat. 10: 850. 1759. Figure 8. Herbs to suffrutescent plants to 1 m tall, stems some- what grooved and hexagonal in younger portions, gla- brous to pilose; internodes between leaf-bearing nodes, Herbs to suffrutescent plants to 2 m tall, younger stems quadrangular, older stems terete; internodes between leaf- bearing nodes 2 (l)-7 cm long, 2 mm thick, puberulous along 2 lines toward the apex. Leaves with glabrous pet- ioles to 2 cm long; laminae ovate to elliptic, 5-15 cm long, 2-5.5 cm broad, apically acuminate, basally acute to obtuse, margins undulate, glabrous, cystoliths visible on both surfaces, but more prominent above. Inflores- BURGER: FLORA COSTARICENSIS 53 cence in loose, axillary spikes to 3 cm long, 1 cm broad, rachis glandular-pubcrulous. peduncle to ca. 1 cm long, puberulous along 2 lines; bracts linear-spatulate, spat- ulate, or orbicular, 8-10 mm long, 0.5 mm to 8 mm broad, glandular-puberulous, bracteoles linear, 8-10 mm long, 0.25-0.5 mm broad, glandular-puberulous; bracts and bracteoles usually spreading. Flowers with 5-merous calyx, segments lanceolate, equal, 5.5-6.5 mm long, 1- 1.25 mm broad, puberulous; corolla saccate, glandular- puberulous, pale green or greenish yellow with broad dark red or purplish streaks at the upper part of the tube, 1 1.5-12.5 mm long, 1.5 mm broad basally, expanding immediately to 4 mm broad just above base, bilabiate, upper lip ca. 6 mm long, 6.5 mm broad, apically bilobed, lobes acute, 0.5 mm long, 1 mm broad, lower lip 7-7.5 mm long, lateral lobes 4 mm long, 3.5 mm broad, middle lobe 4.5 mm long, 3.5 mm broad, all lobes apically rounded; stamens exserted to just below the upper lip, anther cells superposed, upper cells basally apiculate, 1.5 mm long, 0.75 mm broad, filaments glabrous. Fruit not seen. This is an endemic species of mid-altitudes, 1 000 to 1500 m. It grows in the shade of moist forests in both the Pacific and the Caribbean watersheds, including the slopes of the Cordilleras Central, Talamanca, and Tilaran. The collections of Palm- er 144 and 161 (NY) from Monte verde, Puntar- enas, have broader bracts, orbicular in shape, which are different from those of all other collections, but are otherwise identical. Flowering collections have been made from January through April. Justicia costaricana is recognized by its spread- ing, spatulate to orbicular bracts, spreading linear bracteoles, and greenish white, red-throated, sac- cate corollas. There is variation in the shape of the bracts which requires that other features be ex- amined before identification is confirmed. It may be confused with J. carthaginensis, but this species has longer corollas (25 to 30 mm) and grows in disturbed open habitats. Justicia crenata (Leonard) Durkee, Ann. Missouri Bot. Card., 65: 217. 1978. Jacobinia crenata Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1223. 1938. Figure 9. Shrub to 5 m tall, stem terete; internodes between leaf- bearing nodes 2.5—4 cm long, 2-3 mm thick, glabrous. Leaves with petioles to 5 cm long, glabrous; laminae elliptic, 9-24 cm long, 2.5-6 (9) cm broad, apically acu- minate, basally acute to attenuate, margins crenate, gla- brous, cystoliths visible on both surfaces. Inflorescences in large, terminal, spicate panicles to 27 cm long and 1 5 cm broad, peduncles and rachises puberulous; bracts el- liptic to obovate, apically rounded to acute, soon decid- uous, 5-10 mm long, 2-5 mm broad, sparingly glan- dular-puberulous; bracteoles similar, but smaller. Flowers with 5-merous calyx, the segments linear lanceolate, equal, to 7 mm long, 1.25 mm broad basally, sparingly glandular-puberulous, the margins minutely ciliolate; co- rolla bilabiate, red with yellow tip, to 5 cm long, pu- berulous glandular, the tube narrow, 3 mm broad at base narrowing to 2 mm broad at the mouth, the lips erect, each to 2.5 cm long, 6 mm broad, gradually narrowing from base to tip, the upper lip apically notched, the lower lip with 3 small lobes at the apex, the lobes each ca. 2 mm long, rounded and hooded apically; stamens exsert- ed to the tip of the lobes, the anther cells unequal, each to 2.5 mm long, each slightly spurred basally, the lower cell more so. Fruit clavate, 1.5 cm long, glabrous or with a few glandular hairs at the apex. Found in rain forests of Limon Province, rang- ing to cloud forests of higher elevations (to 1 500 m) of the Cordillera Central region. Also found in similar habitats of adjacent Panama. Flowering collections made in February and May through November, except July. Justicia crenata is recognized by its large, ter- minal panicles of short spikes, its rounded bracts, and its comparatively large, crenate leaves. It has been confused with J. urophylla, which differs in its dense spikes, linear, subulate bracts, entire to undulate leaf margins, and white corolla. Justicia ephemera Leonard, Contr. U.S. Natl. Herb. 31: 606. 1958. SufTrutescent plants to 2 m tall, stems upright, much- branched; internodes between leaf- bearing nodes 5-1 1 cm long, 2-3.5 mm thick, upper internodes tetrangular, glabrous to puberulous along 2 lines. Leaves sessile or nearly so; laminae elliptic, 5-17 cm long, 1.3-5.5 cm broad, apically long-acuminate, gradually narrowing to an auriculate base, margins crenulate, glabrous on both surfaces, cystoliths numerous, minute, visible on both surfaces. Inflorescences in terminal, lax, spicate panicles or single spikes in early flowering stages, to 10 cm long, peduncles 2-3 cm long, puberulous; floral bracts subu- late, 2-6 mm long, ca. 1 .0 mm broad. Flowers with 5- merous calyx, calyx segments subulate, 2.5 cm long, ca. 0.75 mm broad, sparingly ciliolate; corolla white, bila- biate, 10-13 mm long, 2 mm broad basally, glabrous, the lips equal, 8-9 mm long, the upper lip ovate, apically bidentate, 3.5-4 mm broad, lobes of lower lip ovate, apically obtuse, 2.5-3 mm broad, middle lobe grooved and laterally sculpted to form a nectar guide, slightly broader than lateral lobes; stamens exserted to ca. 'A-'/s the length of the corolla lobes, 3.5 mm long, glabrous, anthers moderately superposed, 1 mm long, apically and basally obtuse. Fruits not observed. This species is found in wet areas of rain forests in Colombia, Panama, and Costa Rica. One Costa Rican collection, (Allen 5223, us), is known from 54 FIELDIANA: BOTANY Puntarenas Province along the banks of the Rio Grande de Terraba, on the trail from Palmar Norte to Cana Blanca, at an elevation of 0 to 100 m. This flowering collection was made in late March. Flowering collections from Panama and Colombia were also made in late March or April. Justicia ephemera is recognized by its elliptic leaves with auriculate bases and small (10 to 13 mm long), white corollas with barely exserted sta- mens on broad, lax, spicate panicles. Because of its unusually shaped leaves, it is not easily con- fused with other Costa Rican species of Justicia. Justicia macrantha Benth., PI. hartw. 78: 1839. Figure 9. Shrub to 3 m tall, older stems terete, quadrangular toward apex; internodes between leaf-bearing nodes 2- 6 cm long, 1.5-3.5 mm thick, pilose to glabrate. Leaves with petioles to 1.5 cm long; laminae elliptic, 6-16 (19) cm long, 2.5-5.5 (6) cm broad, apically acuminate, ba- sally attenuate, margins crenulate, glabrous on both sur- faces except for pilose hairs in some, both surfaces fre- quently gland-dotted, glands more abundant below, cystoliths abundant on both surfaces. Inflorescences in axillary cymes; bracts and bracteoles subulate, 2-3 mm long, ca. 1 mm broad basally, glabrous, peduncles to 4.5 cm long, glabrous to sparingly pilose; pedicels 3-5 mm long, glabrous. Flowers with 5-merous calyx, segments subulate, 3-4 mm long, ca. 1 mm broad basally, keeled, glabrous, usually ciliolate; corolla orange red to yellow- ish, 4.5-5.5 cm long, deeply bilabiate, glabrous, the tube funnelform, basally 4 mm broad, 8 mm broad at the mouth, both lips ca. 2.8 cm long, the upper lip erect, 8 mm broad, apically dentate in 2 short lobes and hooded to enclose part of the style, the lower lip spreading, 1 cm broad, 3-lobed at the tip, the lobes about equal, 4 mm long, apically rounded; stamens exserted to just below the tip of the upper lip, the anthers pendulous, converg- ing apically and diverging basally, the cells 2.5 mm long, slightly unequal, basally rounded, the filaments stout, 1 mm broad, glabrous. Fruits clavate, 1 8-20 (22) mm long, 4 mm broad, 2.5 mm thick, puberulous; seeds oval, flattened, 3 mm long, 2 mm wide, roughened. This species is found through most of Central America in forests at altitudes above 500 m. In Costa Rica it has been collected in cloud forests as high as 1 750 m, including the Cordilleras Gua- nacaste, Tilaran (Monteverde), Central, and Tala- manca. It has been collected in all provinces except Limon. Flowering collections have been made from November through April, excluding March. Justicia macrantha is recognized by its axillary cymes with large colorful corollas, which are strongly bilabiate, and its pendulous anthers with cells lacking spurs. It has been confused with /. secunda, which has mostly terminal inflorescences which are thyrsoid (cymose panicles) or panicu- late, shorter corollas (2.5 to 4.3 cm long), upright anthers, and shorter capsules (ca. 1 cm long). Justicia metallica Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim, fl. costaric. 2: 311. 1900. Figure 9. Herb to 1 m tall, stem ascending, rooting at the lower nodes, younger stems quadrangular, the angles rounded, pilosulus in 2 lines, mixed with glandular hairs toward the apex, purplish; internodes between leaf-bearing nodes 1.5-6.5 cm long, 1-2 mm thick. Leaves with petioles 3- 10(15) mm long, pubescence of petioles that of the stem; laminae ovate, 3-9 (12) cm long, 1.5-4 (6) cm broad, apically acuminate to short acuminate, basally sparingly attenuate to rounded, margins entire, undulate or cre- nate, glabrous to sparingly strigose along the midrib and the costa, the cystoliths prominent on both surfaces, dried leaves greenish above, shiny metallic blue beneath. In- florescences of lax, terminal or axillary, spicate panicles, the spikes ca. 8 cm long, the first internode ca. 1 cm long, the peduncles and rachises glandular-puberulous, purplish; bracts and bracteoles linear subulate, to 1.5 mm long, 0.25 mm broad basally, glandular-puberulous. Flowers with a 4-merous calyx, calyx segments subulate, to 5 mm long, ca. 0.5 mm broad basally, glandular- puberulous; corollas pink to purplish, 12-14 mm long, sparingly puberulous, the tube cylindrical, to 9 mm long, 2 mm broad, the upper lip erect, to 4.5 mm long, 2 mm broad, apically minutely bidentate, the lower lip spread- ing, to 5.5 mm long, the lobes to 2 mm long, the middle lobe 2 mm broad, the lateral lobes 1.5 mm broad, all apically rounded; stamens exserted to just below the tip of the upper lip, the anther cells superposed, ca. 1 mm long, the lower cell spurred. Fruits clavate, capsules 7- 8 mm long, ca. 2 mm broad, 2 mm thick, sparingly puberulous; seeds reddish brown, suborbicular, 1.5 mm in diameter, roughened. Found along stream banks and in wet places of rain forests at elevations generally under 600 m. This species has been collected in much of Panama and in adjacent Costa Rica on the Pacific slope, mostly in the vicinity of the basin of the Rio Gen- eral and the Osa Peninsula. Flowering collections have been made in Costa Rica in January and February. Justicia metallica is recognized by its leaves with shiny blue metallic undersides, its purplish stems and rachises, its four-parted calyx, and its lax spi- cate panicles. It is not easily confused with other species of the genus. Justicia oerstedii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1230. 1938. Figure 9. BURGER: FLORA COSTARICENSIS 55 Herb to suffrutescent plant, 1.5 m tall, steins erect, terete to subquadrangular when young; internodes be- tween leaf-bearing nodes, 1.5-4.5 cm long, 0.75-1.25 mm thick, glabrous to puberulous. Leaves with petioles to 5 mm long, petioles glabrous; laminae elliptic to ovate, 3-10 (12) cm long, 1-2.5 (3) cm broad, apically acu- minate to acute, basally attenuate, margins entire to un- dulate, glabrous on both surfaces, cystoliths prominent on both surfaces. Inflorescences in terminal spicate pan- icles to 20 cm long, the spikes lax, the peduncles and rachises glandular-puberulous; bracts and bracteoles narrowly subulate, 1-1.5 mm long, ca. 0.5 mm broad basally, puberulous. Flowers with 5-merous calyx, the calyx segments subulate, 3—4 mm long, 0.5 mm broad, the posterior segment smaller, 1.5-2 mm long, 0.25 mm broad, all segments keeled, puberulous; corolla bright red, 2.5-3.5 cm long, puberulous, the tube narrow, 1 mm broad basally, 5-6 mm broad at the mouth, the upper lip oblong, to 9 mm long, 5 mm broad, apically minutely bidcntatc. the lower lip to 10.5 mm long, 3-lobed, the middle lobe 2.5 mm broad, the lateral lobes 1.5 mm broad, all lobes to 3 mm long, apically rounded; stamens exserted to about the tip of the upper lip, the anther cells superposed by a connective, 0.5 mm broad, the lower cell rounded, acute at the base. Fruits clavate, to 1 2 mm long, 2 mm broad, puberulous; seeds subor- bicular, flattened, ca. 1.5 mm in diameter, roughened. This species is abundant where found in rain forest and cloud forest areas in moist, shaded lo- cations along streams and trails at elevations of 500 to 1 800 m in Costa Rica and Panama. In many locations this is the most abundant herb in flower. In Costa Rica it is found mainly on the Pacific watershed areas of all the Cordilleras from Pan- ama to the Cordillera Guanacaste. It has not been collected in Limon Province. Flowering collec- tions have been made from December through April. Justicia oerstedii is recognized by its bright red flowers with corollas more than 2 cm long, its lax, spicate panicles with glandular-puberulous ra- chises, and its five-parted calyx with one segment much shorter than the others. It may be confused with J. macrantha, which differs in its longer co- rolla (more than 4 cm long), its axillary, cymose inflorescences, and its four-parted calyx. Justicia orosiensis Durkee, sp. nov. Figure 9. Herbae vel suffrutices, usque ad 1.2 m altae, caulibus erectis et difrusis. Folia petiolata, laminis ellipticis, 8- 23 cm longis, 2-7 cm latis, apice acuminatis, basi atten- uatis, supra glabris, inferne plerumque glabris, cystolithis nullis. Inflorescentiae paniculatae spiciformes, compac- tae, terminales, bracteis laxe imbricatis, obovatis vel el- lipticis, 6-8 mm longis, bracteolis similibus bractearum sed minoribus. Flores virello-albi ad lilacini, corollis 1 2- 14 mm longis, staminibus exsertis. Fructus clavati, 12- 14 mm longi, seminibus 4. Herbs to suffrutescent plants to 1.2m tall, stems erect, but diffuse, quadrangular when young and dark maroon, internodes between leaf-bearing nodes, 1.5-9.5 cm long, 1.5-3.5 mm thick, glabrous to sparingly puberulous. Leaves with glabrous to sparingly puberulous petioles 1- 2 cm long; laminae elliptic, 8-23 cm long, 2-7 cm broad, apically acuminate, basally attenuate, margins entire to slightly undulate, glabrous above, glabrous to sparingly puberulous along costa below, cystoliths obscure on both surfaces. Inflorescences in compact terminal, spicate panicles superficially resembling single spikes, 7-1 1 cm long, 1.5-2 cm broad excluding corollas, peduncles and rachises strigose; bracts and bracteoles obovate to ellip- tic, bracts loosely imbricate, 6-8 mm long, 2.5-3.5 mm broad, apically obtuse to rounded, basally attenuate, stri- gose mostly at base, bracteoles similar although slightly smaller. Flowers with 5-merous calyx, the calyx seg- ments subulate, 6-7 mm long, 1.25-1.5 mm broad, gla- brous; corolla greenish white to lilac, 16-17 mm long, secund, glabrous, the tube cylindrical, 6 mm long, 2 mm broad, the upper lip curved, oblong, partially enclosing the stamens, to 11 mm long, 6 mm broad, apically notched, the lower lip to 9.5 mm long, 3-lobed, the mid- dle lobe 4 mm broad, the lateral lobes 3 mm broad, all lobes to 7 mm long, apically rounded; stamens exserted to about the tip of the upper lip, the anther cells super- posed, 2.5 mm long, the lower cell calcarate, the upper cell with apiculate base, pollen is 4-porate. Fruits clavate, 12-14 mm long, 4—5 mm broad, 3.5—4 mm thick, gla- brous; seeds 4, suborbicular, ca. 3 mm in diameter. TYPE— Costa Rica; Cartago Prov., 10 km south of Tapanti, elevation 1 600 m, in primary forest high over Rio Grande de Orosi, Lent 954 (holo- type, F). Additional collections: Cartago Province, forested slopes above Platanillo, elevation 1000 m, Wilbur and Stone 10611 (DUKE); along stream near Tausito, elevation 1400 m, Lent 3818 (CR, F). Flowering collections were made in mid-May, late August, and February. Justicia orosiensis is recognized by its compact spicate panicles superficially resembling spikes and its secund, greenish white corollas. It may possibly be confused with the much more common and wide-ranging J. carthaginensis of disturbed habi- tats, which differs further in its dense spikes, larger rose-purple corollas (24 to 35 mm long vs. 16 to 17 mm long), and larger (25 to 35 mm long), ob- long-spatulate to spatulate bracts versus the 6- to 8-mm long, obovate to elliptic bracts of J. oro- siensis. Justicia parvibracteata Leonard, Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1231. 1938. Figure 10. 56 FIELDIANA: BOTANY Erect herb to 35 cm tall, stems quadrangular, terete in older portions; inter nodes between leaf-bearing nodes 1- 4 cm long, 1 mm thick, puberulous mostly along 2 lines. Leaves with puberulous petioles to 8 mm long; laminae narrowly elliptic-oblong, 3.5-7 cm long, 0.7-1.2 cm broad, margins entire to undulate, apically acute, basal 1 \ attenuate, cystoliths prominent on both surfaces, gla- brous. Inflorescences in terminal and axillary spikes or subpaniculate spikes to 6 cm long, 6 mm broad excluding corollas; bracts barely imbricate, elliptic to obovate, 4- 6 mm long, 1-2 mm broad, apically obtuse, apiculate, veins prominent, bracteoles narrowly elliptic, 3-4 mm long, 0.5-0.75 mm broad, sparingly ciliolate, peduncles 0.5-2.5 cm long, puberulous mostly along 2 lines, rachis glandular-puberulous. Flowers in fascicles of 2—4 per bract; calyx 5-merous, posterior segment setaceous, 1.5- 2.5 mm long, the remainder lanceolate, 3.5-4.5 mm long, ca. 0.5 mm broad, all glandular-puberulous, corolla white, 7.5-8 mm long, 1.5 mm broad basally, expanding to 2 mm broad at the mouth, sparingly puberulous, upper lip 3.5-4 mm long, 2 mm broad basally, apically acute, lower lip 4-4.5 mm long, 2 mm broad, 3-lobed, lobes 1 mm long, ca. 1.5 mm broad, apically rounded; stamens exserted to ca. Vi the length of the upper lip, anther cells superposed, ca. 0.5 mm long, lower one slightly smaller, rounded basally. Fruits clavate, 4.5-5.5 mm long, 1-1 .25 mm broad, 1-1.25 mm thick, sparingly puberulous, seeds reddish brown, ovate, 1 mm long, 1 mm wide, papillate. Found in wet areas along streams and river banks on the Caribbean slope of the Cordillera Central and on steep forested slopes of the Osa Peninsula, at altitudes of 450 to 1400 m. Flowering collec- tions have been made from January through March. Justicia parvibracteata is recognized by its nar- row spikes with small (4 to 6 mm long), elliptic to obovate bracts with narrow apices, short, white, inconspicuous corollas (less than 1 mm long), and its narrowly elliptic-oblong laminae. It has some resemblance to J. candelariae and J. valerii, but can be distinguished from these by its smaller bracts (4 to 6 mm long vs. 6 to 10 mm) and narrower leaves. Justicia pectoralis Jacq., Enum. syst. pi. 11. 1760. Figure 10. Herbs to 2 m tall, stems weak, erect or ascending, subquadrangular, grooved; internodes between leaf- bearing nodes 2-4.5 mm long, 1-2 mm thick, retrorse hairs mostly along 2 lines. Leaves with short strigose petioles to ca. 5 mm long; laminae narrowly to broadly lanceolate, 4-8 ( 1 0) cm long, 1 -2 (2.5) cm broad, margins entire to undulate, apically acuminate, basally acute to obtuse, cystoliths dense and prominent above, not vis- ible beneath, glabrous to strigose at the costa above, glabrous beneath. Inflorescences in terminal, lax, spicate panicles to 25 cm long, 1 5 cm broad, the lower inter- nodes of spicate branches ca. 2 cm long, rachises and peduncles glandular-puberulous; bracts and bracteoles subulate, 2 mm long, 1 mm broad basally, puberulous with a few glandular hairs. Flowers with a 5-merous calyx, segments narrowly subulate to 2 mm long, 0.5 mm wide, basally glandular puberulous; corollas white, lilac or purple, 8-12 mm long, puberulous outside, the throat transversely plicate, often spotted with dark pur- ple, the tube narrowly funnelform, basally 1 .25 mm broad, 2.5 mm broad at the mouth, the upper lip erect, trian- gular 3.5 mm long, 3 mm broad, apically acute, the lower lip spreading, to 5 mm long, 3-lobed, the lobes 1 mm long, the middle lobe 2 mm broad, the lateral lobes 1.5 mm broad, all apically obtuse; stamens exserted to about the tip of the upper lip, the anther cells slightly super- posed, basally apiculate, the filaments glabrous. Fruits clavate, capsules 7-8 mm long, 1.5-2 mm broad, ca. 1 mm thick, puberulous; seeds orbicular, flattened, 1.5 mm in diameter. This is a wide-ranging species of roadsides and waste places in tropical America. In Costa Rica it occurs from near sea level to 900 m altitude and has been collected in all provinces except Alajuela. Flowering collections have been made from Jan- uary through May. Justicia pectoralis is recognized by its terminal spicate panicles, with corollas 8 to 12 mm long, puberulous capsules 7 to 8 mm long, and lanceo- late laminae 4 to 8 mm long. It is most easily confused with J. comata, but can be distinguished from it by its terminal spicate panicles (vs. ter- minal and axillary spicate panicles), longer corol- las (8 to 12 mm vs. 4 to 6 mm long), and longer, puberulous capsules (7 to 8 mm long vs. papillose capsules, 4 to 5 mm long). Justicia pittieri Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 1 89. 1 898, and in Pitt., Prim, fl. costaric. 2: 312. 1900. Figure 10. Herbs to 50 cm tall, stems ascending, younger stems quadrangular, grooved, retrorse hairs mostly along 2 lines; internodes between leaf-bearing nodes 5.5-7 cm long, 1.5-2 mm thick. Leaves petiolate, petioles short, 1-10 mm long, strigose; laminae lanceolate-elliptic, 5-12 cm long, 2-3 cm broad, margins entire, apically acuminate, basally acute, glabrous to sparingly strigose, cystoliths dense and prominent on both surfaces. Inflorescences in terminal and axillary spikes, to 1 5 cm long, 1 cm broad, excluding corollas, rachises and peduncles with retrorse pubescence mostly along 2 lines, plus some glandular hairs, peduncle to 1 1 cm long, bracts imbricate, spread- ing at anthesis, elliptic, 4.5-5.5 mm long, 1.5-2.2 mm broad, apically and basally acute, subglabrous, glandu- lar-puberulous, ciliate, bracteoles elliptic, 3.5-4 mm long, 0.5-0.75 mm broad, glandular-puberulous, ciliate. Flow- BURGER: FLORA COSTARICENSIS 57 ers in fascicles of 2-3 per bract, calyx 5-merous, posterior segment setaceous, 2 mm long, remaining segments su- bulate, 3.5-4.5 mm long, 0.5-0.75 mm broad, glandular- puberulous; corollas white with purple dots, 7-9 mm long, 1 mm broad at the base expanding to 2.25 mm broad at the mouth, puberulous with a few glandular hairs, the throat transversely plicate, the plaits purplish, upper lip 5 mm long, 2 mm broad basally, apically bi- dentate, lower lip 3.5 mm long, the lobes orbicular, ca. 1 mm in diameter; stamens exserted ca. '/z the length of the upper lip, anther cells superposed, ca. 0.5 mm long, basally obtuse, lower cell slightly smaller. Fruits not seen. Only two collections of this species have been observed: the type, Pittier 8642 (CR) from lowland rain forests of the upper Rio Yorkin between Costa Rica and Panama and Skutch 4652 (F, MO) at 600 to 850 m altitude in the vicinity of Pejibaye (Pe- jivalle), Cartago Province. These flowering collec- tions were made in January and March. Justicia pittieri is recognized by its inflores- cences with elliptic bracts (4.5 to 5.5 mm long), which spread at anthesis, flowers in fascicles of two to three per bract, unequal calyx segments, and stems with retrorse hairs mostly along two lines. Justicia pittieri may be confused with J. ton- duzii, but differs from it in its unequal calyx seg- ments, elliptic bracts (vs. oblanceolate), and gla- brous to sparingly strigose leaves (vs. pilose). lobes 2-2.5 mm long, apically obtuse, the middle lobe 2-2.5 mm broad, the lateral lobes 1.5-2 mm broad, the stamens exserted to ca. the tip of the upper lip, the fil- aments attached to the tube just below the mouth, pu- bescence of minute glandular hairs, the anther cells su- perposed, the upper cell mucronate, the lower cell spurred. Fruits clavate, capsules 7.5-10 mm long, 2 mm broad, 2 mm thick, puberulous; seeds ovate, ca. 1.5 mm long, 1.5 mm broad, flattened, roughened. Found at lower elevations frequently in moist, disturbed areas that are shaded in both Costa Rica and Panama. It has been collected only in the Ca- ribbean lowlands of Costa Rica. Flowering collec- tions have been made in November, December, and July. Justicia refractifolia is recognized by its axillary spikes with small, imbricate, ovate bracts which are brown when dried and have a prominent re- ticulate venation and ciliate margins; its small, relatively inconspicuous flowers (ca. 1 5 mm long) with superposed anther cells that are spurred only on the lower cell; and its rather large (11 to 18 cm long), elliptic-obovate leaves. Justicia refractifolia has some resemblance to J. refulgens, but can be distinguished from it by its unbranched spikes (vs. trifurcate spikes), ovate ciliate bracts (vs. lanceo- late without cilia), and longer bracts (8 to 10 mm vs. 6 mm long). Justicia refractifolia (Kuntze) Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1232. 1938. Ec- bolium refractifolium Kuntze, Revis. gen. pi. 2: 487. 1 89 1 . Justicia pseudopolystachia Cuf., Arch. Bot. Sist. 10: 49. 1934. Figure 10. Herbs to suffrutescent plants, 1 m tall, stems simple, erect, subquadrangular toward the apex, subterete below; internodes between leaf- bearing nodes 4-10 cm long, 1.75-2.5 mm thick, pilose to tomentose. Leaves with pilose to tomentose petioles to 3 mm long; laminae el- liptic to elliptic-obovate, 1 1-18 (21)cm long, 3-6.5 (8.5) cm broad, margins entire to undulate, apically acumi- nate, basally acute to attenuate, glabrous to strigose on the costa and veins above, pilose to sparingly strigose beneath, cystoliths prominent to obscure above, obscure beneath. Inflorescences in terminal and axillary spikes, borne singly and in clusters, to 1 5 cm long, 1 cm broad, the rachis pilose to strigose; bracts ovate, 8-10 mm long, 3.5-5 mm broad, imbricate, apically acute, basally ob- tuse, drying brown, the veins prominent, sparingly pilose on both surfaces, ciliate; bracteoles lanceolate, slightly falcate, 6.5-9 mm long, 1.5-2 mm broad, puberulous, ciliolate. Flowers with 5-merous calyx, segments nar- rowly subulate, 5.5-6 mm long, 0.5-0.75 mm broad, puberulous, ciliolate; corolla white to purple, the tube cylindrical, 7-8 mm long, 1.5 mm broad, puberulous, the upper lip erect, ca. 4—4.5 mm long, 2 mm broad, apically bidentate, lower lip 4-5 mm long, 3-lobed, the Justicia sarapiquensis McDade, Syst. Bot. 7: 489- 493. 1982. Figure 10. Suffrutescent vine climbing to 15 m; internodes be- tween leaf-bearing nodes 1.5-2.5 cm long, 2-2.5 mm thick, subquadrangular, sparingly puberulous, older stems terete. Leaves with sparingly puberulous petioles 5-10 mm long; laminae ovate, 4.5-7.7 cm long, 2.2-3.5 cm broad apically acuminate, basally obtuse to rounded or slightly cordate, margins entire, glabrous on both sur- faces, cystoliths prominent on both surfaces. Inflores- cences in lax, terminal panicles of up to 15 flowers, pe- duncles and branches sparingly puberulous; bracts subtending the flowers borne singly or in pairs, ovate, magenta, 15-21 mm long, 14-16 mm broad, apically obtuse, glabrous, bracts on lower parts of peduncle small- er; bracteoles 2 per flower, lanceolate, magenta, 16-19 mm long, 6-10 mm broad, apically acute, glabrous. Flowers with 5-merous calyx, calyx lobes equal, 1 5-22 mm long, 2-4 mm broad, white, apically acute and apic- ulate, glabrous; corolla white, 6.5-7 cm long, outside with fairly dense white pilose hairs 1-1.5 mm long, the tube 38—40 mm long, 5-6 mm broad at base, narrowed to 2.5-3 mm broad above the ovary, expanding to 5-6 mm broad at the throat, the upper lip erect, 23-26 mm long, apically acute, the lower lip slightly reflexed at an- thesis, 27-29 mm long, apically 3-lobed, each lobe 3-4 mm long; stamens extending to just below the upper lip, anther cells slightly superposed, ca. 4 mm long. Fruits 58 FIELDIANA: BOTANY clavate, dark brown, 15-20 mm long, 5-6 mm broad, 3-4 mm thick, glabrous, floral bracts, bracteoles, and calyx are strongly reflexed when fruits are mature; seeds 4, orbicular, flattened, 3—4 mm in diameter, glabrous. This species grows in shaded areas of the rain forest lowlands of northern Costa Rica at eleva- tions of 100 to 575 m. It has been collected in only two sites, near Quesada (Villa Quesada), San Carlos, and the La Selva Research Station, both in Heredia Province. Flowering collections have been made in March. Justicia sarapiquensis is recognized by its viny habit, its large magenta bracts and bracteoles, its white calyx, and its white, pilose corolla. It is not easily confused with other species of Justicia from Costa Rica; however, the Panamanian species, /. graciliflora (Standley) D. Gibson, has a viny habit, but lacks the large, showy bracts and bracteoles. Justicia skutchii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1234. 1938. Figure 10. Shrub to 1 m tall, stems quadrangular, older portions terete; internodes between leaf-bearing nodes 1-3.5 cm long, 1.5-2 mm thick, densely pilose. Leaves subsessile; laminae ovate-elliptic, 3.5-13 cm long, 3.5-5 cm broad, margins entire to slightly undulate, apically acuminate, basally rounded or subcordate, cystoliths prominent on both surfaces, sparingly pilose above, pilose beneath. Inflorescences in terminal spicate panicles, spikes lax, to 9.5 cm long, 8 mm broad excluding corollas, peduncle and rachis pilose-glandular, bracts elliptic, 3.5-4 mm long, 1 mm broad, sparingly puberulous, ciliolate, brac- teoles linear, 3-3.5 mm long, ca. 0.25 mm broad, spar- ingly glandular-puberulous. Flowers in clusters of 2-3 per bract, calyx 5-merous, posterior segment setaceous, 3.5-4 mm long, puberulous, other segments subulate- linear, 3.4-5 mm long, 0.5 mm broad, puberulous; co- rolla white and purple, glabrous, 7.5-9 mm long, tube cylindrical 4.5-5 mm long, 2 mm broad, upper lip acute, ca. 3 mm long, 2 mm broad, lower lip ca. 3 mm long, lobes 0.5 mm long, rounded, lateral lobes 1 .25 mm broad, middle lobe 2 mm broad; stamens exserted to ca. '/2 the length of the upper lip, cells unequal, 0.5 mm long, ba- sally acute. Fruits clavate, capsules 8.5-9 mm long, 2 mm broad, 2 mm thick, puberulous, seeds reddish brown, ovate, 1.75 mm long, 1.5 mm broad. Found in moist habitats on rocks along a stream in a forest in the vicinity of General Viejo (El General), San Jose Province, at 9 1 5 m elevation. Known only from the type collection, Skutch 3012 (MO). Justicia skutchii is recognized by its lax, spicate panicles with small (3.5 to 4 mm long), elliptic bracts, dense yellowish pilose hairs on the stem, and basally rounded to subcordate leaves. It re- sembles J. tonduzii, but can be distinguished from it by its leaves with rounded to subcordate bases (vs. obtuse or narrowed bases) and its longer cap- sules (8.5-9 mm long vs. 5 mm). Justicia spicigera Schlecht., Linnea 7: 395. 1832. Jacobinia spicigera (Schlecht.) L. H. Bailey, Stand. Cycl. Hort. 1715. 1915. Figure 11. Erect or scandent shrub, 1-1.5 m tall; internodes be- tween leaf-bearing nodes 1-5 cm long, 1-2 mm thick, quadrangular, glabrous to puberulous along 2 lines. Leaves with petioles 5-10 mm long, petioles mostly puberulous along 2 lines; laminae ovate, 6.5-17 cm long, 3.5-9 cm broad, apically acuminate, basally attenuate, margins en- tire to undulate, cystoliths abundant and prominent on both surfaces, dried leaves often purplish black. Inflo- rescences in terminal and axillary spicate panicles, flow- ers secund, peduncles and rachises puberulous; bracts and bracteoles subulate, 1-1.5 mm long, glabrous. Flow- ers with a 5-merous calyx, calyx segments subulate, 2.5- 3 mm long, 0.75-1 mm broad basally, glabrous; corollas orange, 3.5-4.4 cm long, appearing fusiform before opening, glabrous, the upper lip erect, ca. 1 5 mm long, 6.5 mm broad, apically acute, the lower lip ca. 15 mm long, 5 mm broad, frequently rolled up, lobes small, suborbicular, 1.5 mm long, 1.5 mm broad; stamens exserted approximately to the tip of the upper lip, the anther cells subequal, ca. 2.5 mm long, basally apiculate. Fruits not seen. Found along stream banks in thickets and for- ests of Mexico and Central America, this plant is often cultivated for use as laundry bluing and in some areas as a dye. Only one collection of this plant, Skutch 4809 (F, MO, NY), has been made in Costa Rica, in the Rio General basin in March. It had abundant flowers, but no fruits. Justicia spicigera is recognized by its frequently darkened, ovate leaves, axillary spicate panicles with secund flowers, and orange corollas (3.5 to 4.4 cm long) which are fusiform when unopened. It is easily confused with J. tinctoria, which is also cultivated for the same purposes, but can be dis- tinguished from it by its fusiform corollas (vs. nar- rowly funnelform) and its larger, ovate leaves (6.5 to 1 7 cm long vs. elliptic leaves 4.5 to 14 cm long). Justicia tinctoria (Oerst.) D. Gibson, Fieldiana, Bot. 34(6): 74. 1972. Sericographis tinctoria Oerst., Vidensk. Meddel. Dansk Naturhist. For- en. Kjobenhavn 1854: 150. 1855. Jacobinia tinctoria (Oerst.) Hemsley, Biol. Centralbl. 2: 552. 1882. Figure 11. BURGER: FLORA COSTARICENSIS 59 Shrubs to 2 m tall, younger steins subquadrangular, the angles rounded; internodes between leaf-bearing nodes 1.5-5 cm long, 1.5-2.5 mm thick, glabrous. Leaves with glabrous petioles to 1.2 cm long; laminae elliptic, 4.5- 14 cm long, 1.5-4.5 cm broad, margins entire to un- dulate, apically acute to acuminate, basally attenuate, cystoliths visible on both surfaces, but more prominent below. Inflorescences in lax axillary spikes or spicate panicles with secund flowers, occasionally branched, spikes to 4 cm long, peduncles to 2 cm long, glabrous to puberulous; rachis subquadrangular, internodes 4-5 mm long; bracts and bractoles subulate, 1-2 mm long, 0.5- 1 mm broad at base. Flowers with 5-merous calyx, calyx segments subulate, 2.25-3 mm long, 1 mm broad at base, glabrous; corolla orange or red, 3-4 cm long, the tube slightly funnelform, 3 mm broad basally, 5 mm broad at the mouth, the lips subequal, ca. 16 mm long, the upper lip erect, 7 mm broad basally, apically acute and keeled, the lower lip oblong, erect or slightly spreading, 4.5 mm broad, apically 3-lobed, the lobes equal, 2 mm long, 1 mm broad, apically obtuse and keeled; stamens exserted to about the tips of the lips, the anther cells subequal, 2.5 mm long, obliquely attached to the con- nective, mucronulate, filaments glabrous. Fruits not seen. First described from Costa Rica, this species is found in dry thickets of Central America and is often cultivated. When steeped in hot wtaer, its leaves produce a bluing solution, azul de mata, used in laundering white clothes. Flowering col- lections have been made in January, March, April, and June. Justicia tinctoria is recognized by its short ax- illary spikes with secund flowers, red or orange, narrowly funnelform, corollas (3 to 4 cm long), and short calyx segments (2.25 to 3 mm long). It is easily confused with J. spicigera, but can be distinguished from it by its smaller elliptic leaves (4.5 to 14 cm long vs. larger ovate leaves 7-17 cm long), and narrowly funnelform corollas (vs. fu- siform corollas). Justicia tonduzii Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim, fl. costaric. 2: 313. 1900. Figure 1 1. Herbs to 25 cm tall, stem ascending, branching, youn- ger stems terete to subquadrangular, pilose hairs to 3 mm long; internodes between leaf-bearing nodes, 2-4 cm long, 1-1.5 mm thick. Leaves with pilose petioles to 5 mm long; laminae ovate to ovate-elliptic, 5-8 cm long, 2-3.2 cm broad, margins undulate, apically acuminate, basally obtuse to rounded, pilose on both surfaces, cys- toliths dense and prominent on both surfaces. Inflores- censes in lax terminal and axillary spikes to 7 cm long, rachiscs and peduncles pilose plus shorter glandular hairs, bracts not imbricate at anthesis. oblanceolate, 2.5-3 mm broad, apically acute, pilose, bracteoles linear ca. 2.5 mm long, 0.5 mm broad, pilose. Flowers with 5-merous ca- lyx, calyx segments equal, subulate, 2-2.5 mm long, glan- dular-puberulous; corollas purplish, ca. 5-7 mm long, tube cylindrical, ca. 1.5 mm broad, glabrous, the throat longitudinally furrowed, upper lip 2.5 mm long, 1.5 mm broad, apically acute, upper lip 2.5 mm long, 1.5 mm broad, apically 3-lobed, lobes ca. 0.5 mm long, apically rounded; stamens exserted ca. '/2 the length of the upper lip, anther cells superposed, upper cell larger, basally acute, lower cell basally rounded. Fruits clavate, 6 mm long, 2 mm broad, puberulous, seeds 4, orbicular, 1 mm in diameter. Only one collection of this species has been ob- served, Pittier 16073 (us), from the forests of Vic- toria Farm, plains of Zent, 200 m elevation, Li- mon Province. This flowering collection was made in January. Justicia tonduzii is recognized by its lax inflo- rescences, with minute, oblanceolate bracts (2.5 to 3 mm long), equal calyx segments, and long pilose hairs of stem, inflorescence, rachis, and leaves. Justicia tonduzii may be confused with J. pittieri, but differs from it in its equal calyx segments, oblanceolate bracts (vs. elliptic), and long pilose hairs. Justicia trichotoma (Kuntze) Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1236. 1938. Ec- bolium trichotomum Kuntze, Revis. gen. pi. 2: 488. 1891. Justicia asymmetrica Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim. fl. costaric. 2: 310. 1900. Figure 1 1 . Herbs or small shrubs to 2 m tall, stem erect, subqua- drangular, lower portion terete; internodes between leaf- bearing nodes 1-5 cm long, 1-2 mm thick, strigose. Leaves with strigose petioles to 2 cm long; laminae ovate to ovate-lanceolate, 1 of each pair conspicuously larger than the other, the larger 5-9 cm long, 2-4.5 cm broad, the smaller 2-5.5 cm long, 1-2 cm broad, all apically acu- minate, basally attenuate, margins undulate, sparingly strigose to glabrous above, similar below but more dense, cystoliths often inconspicuous, more visible on lower surface. Inflorescences in small, axillary cymes, pedun- cles to 2.7 cm long, strigose; bracts and bracteoles subu- late, to 1.5 mm long, 0.5 mm broad basally, strigose. Flowers sessile, calyx 5-merous, segments narrowly su- bulate, to 6 mm long, 1 mm broad, puberulous; corolla white, pink or violet, 16-20 mm long, puberulous, the tube funnelform, 1 .5 mm in diameter basally, expanding to 4 mm in diameter at the mouth, upper lip to 6 mm long, 4 mm broad, apically acute with a rounded apex, lower lip to 7 mm long, 3-lobed, lobes 3 mm long, api- cally obtuse, middle lobe 2.5 mm broad, lateral lobes 2 mm broad; stamens exserted to about the tip of the upper lip, anther cells superposed, lower cells calcarate. Fruits clavate, to 1 1 mm long, 2.5 mm broad, 1 mm thick, puberulous; seeds 4, dark brown, suborbicular, ca. 2 mm in diameter, roughened. 60 FIELDIANA: BOTANY Found in wet habitats adjacent to streams, in clearings of rain forests and wet thickets in Pan- ama and Costa Rica, mostly at elevations below 1000 m. In Costa Rica collections have been made only in the Caribbean watershed. Flowering col- lections have been made in February, March, April, and June. Justicia trichotoma is not easily confused with other species in the genus, because of the unequal size of the leaf pairs. Justicia urophylla (Lindau) D. Gibson, Fieldiana, Bot. 34: 74. 1972. Beloperone urophylla Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 1 89. 1898, and in Pitt., Prim. fl. costaric. 2: 316. 1900. Figure 11. Shrub to 2 m tall, stems subterete; internodes between leaf-bearing nodes 2-7 cm long, 1.5-2.5 mm thick, gla- brous to sparingly puberulous in 2 lines. Leaves with petioles to 3 cm long, pubescence that of the stem; lam- inae ovate-elliptic, 4.5-13 cm long, 2-5 cm broad, mar- gins entire to undulate, apically acuminate, basally at- tenuate, glabrous, cystoliths dense, plainly visible on both surfaces. Inflorescences in 1 -several dense terminal spikes to 6 cm long, ca. 6 mm broad excluding corollas, the peduncles to 2 cm long, puberulous mostly along 2 lines, the rachis glabrous to puberulous; bracts imbricate, lin- ear-subulate 4-6 mm long, ca. 1 mm broad basally, pu- berulous, ciliolate, bracteoles triangular, ca. 1 mm long, 0.5 mm broad basally, puberulous. Flowers imbedded in the rachis, calyx segments 5, equal, subulate, 2-3 mm long, ca. 0.5 mm broad, puberulous; corolla white, 3-4 cm long, puberulous, the tube slightly funnelform, 2 mm broad basally, narrower just above the ovary, ca. 3 mm broad at the mouth, the lips 1 1-16 mm long, the upper lip erect, apically bidentate, the lower lip 3-lobed, the lobes 7-10 mm long, the middle lobe 4 mm broad, the lateral lobes 3 mm broad, all apically rounded; stamens exserted ca. 2 mm below tips of the lips, anther cells superposed, basally acute, ca. 2 mm long, connective ca. 1 mm long, the filaments glabrous. Fruits clavate, cap- sules 13-15 mm long, 4 mm broad, 3 mm thick, gla- brous, seeds light brown, suborbicular, ca. 3 mm in di- ameter, glabrous. Found in rain forests of Costa Rica and Panama from sea level to about 1 100 m elevation. In Costa Rica numerous collections have been made, but all are from the vicinity of San Ramon in Alajuela Province. Flowering collections have been made in Costa Rica in February and March. Justicia urophylla is recognized by its dense, ter- minal spikes witih short, inconspicuous bracts (4 to 6 mm long), its white corollas (more than 3 cm long), and its equally five-parted calyx with short segments (2 to 3 mm long). It may be confused with J. secunda, which is found in adjacent regions of Panama, but has not been collected in Costa Rica. It can be distinguished from J. secunda by its dense spikes (vs. a looser, paniculate to thyrsoid inflorescence), white corollas (vs. red or purplish red), and its leaves with basally attenuate laminae (vs. acute or obtuse). Justicia valerii Leonard, Publ. Field Mus. Nat. Hist, Bot. Ser. 18: 1236. 1938. Figure 11. Decumbent herb to 30 cm tall, stems subquadrangular to near terete in older portions; internodes between leaf- bearing nodes 1.5-7 cm long, ca. 1 mm thick, puberu- lence retrorse. Leaves petiolate, petioles short, to 5 mm long, puberulence retrorse to pilose; laminae elliptic- ovate, 3.5-10 (13) cm long, 1.5-4 (5) cm broad, margins entire to crenulate, apically acuminate, basally attenuate, cystoliths dense and prominent on both surfaces, gla- brous. Inflorescences in terminal and axillary spikes to 7 cm long, ca. 1 cm broad excluding corollas; bracts imbricate, ovate-elliptic, 6-10 mm long, 4-6 mm broad, apically obtuse to rounded apiculate, basally obtuse to rounded, strigose hairs mostly restricted to midrib, veins very prominent, forming a coarse reticulation, ciliolate; bracteoles lanceolate, 7-8 mm long, 1 .25-1.5 mm broad, apically acute to acuminate, apiculate, basally acute, veins very prominent, pubescence that of the bracts, ciliolate, peduncles 1-3 cm long, puberulence retrorse. Flowers with a 5-merous calyx, posterior segment setaceous, ca. 2 mm long, other segments subulate, 5.5-6.5 mm long, 0.5-0.75 mm broad at base, glandular-puberulous, co- rollas white or bluish white, 9-1 1 mm long, 2 mm broad at base expanding to 3.5 mm broad at the mouth, spar- ingly puberulous, upper lip ca. 4.5 mm long, 2.5 mm broad at base, apically acute, lower lip 5 mm long, 3- lobed, lobes 2.5 mm long, middle lobe 2.5 mm broad, lateral lobes 2 mm broad, all apically rounded; stamens exserted ca. 1 mm beyond the mouth, anther cells su- perposed, upper cell ca. 0.75 mm long, lower cell ca. 0.5 mm long, both cells basally rounded. Fruits clavate, 7- 8 mm long, 2 mm broad, 1.5 mm thick, puberulous; seeds reddish brown, flattened, suborbicular, ca. 1.25 mm in diameter, papillate. Found in pastures, along trails, and in clearings of premontane moist to premontane rain forest from 750 to 2000 m elevation in both the Carib- bean and Pacific watersheds of the Cordilleras Ti- laran, Central, and Talamanca. Flowering collec- tions have been made from December through March. Justicia valerii is recognized by its stems with short retrorse hairs, spikes with imbricate bracts which are ovate-elliptic with a coarse reticulation pattern, and unequal calyx segments. It is easily confused with J. candelariae and differs from it in its stems with short retrorse hairs (vs. pilose stems), BURGER: FLORA COSTARICENSIS 61 longer laminae (3.5 to 10 cm vs. 2.5 to 6.5 cm), and glandular-puberulous calyx (vs. ciliolate). I outeridium S. Watson REFERENCE— A. Richardson, Revision of Lou- teridium (Acanthaceae). Tulane Studies in Zool- ogy and Botany 17: 63-76. 1972. Shrubs or soft- wooded trees, at least some species deciduous. Leaves petiolate, laminae ovate, elliptic, ob- lanceolate or obovate with entire or crenate margins, cystoliths present and usually prominent in dried spec- imens. Inflorescences in terminal, cymose, bracteate panicles, with a pair of cymes at each node, bracts and bracteoles small. Flowers witih a 3-merous calyx, co- rollas large, red, light green or yellow, the tube expanding abruptly just above the ovary, bilabiate, 2 lobes above, 3 lobes below, throat prominently gibbous; stamens 2 or 4, exserted from an arched position in the bud, anthers 2-celled, cells equal. Fruits a sessile capsule with 2 car- pels; retinacula present, seeds numerous. This is a genus of the New World tropics, with nine woody species found in Mexico through Pan- ama. Collections of L. costaricense have been made in Darien Province of Panama. This province is adjacent to Colombia, and it is likely to be found in that country as well. Louteridium is not easily confused with other genera of the Acanthaceae since it is the only genus with a large, three-merous calyx. This, plus the large flowers and sessile, many-seeded capsules, makes it relatively easy to identify. In Costa Rica this species occurs in moist forests from 500 to 1000 m elevation. It is found in hab- itats such as on rotted logs or humus-covered rocks in streams and along weedy, shaded roadsides. It is known from Vueltas (Las Vueltas), Alajuela Province, near Siquirres in Limon Province and San Vito of Puntarenas Province. Numerous col- lections have been made in lowland rain forests of most provinces in Panama. Flowering collec- tions were made in March and August. Louteridium costaricense is recognized by its large three-merous calyx with narrowly elliptic segments and its large, showy corolla with two stamens. It is most similar to L. tamaulipense of Mexico, but differs from it in its persistent bracts (vs. caducous) and glabrous inflorescence (vs. glan- dular-puberulous). Megaskepasma Lindau Shrub. Leaves petiolate; laminae large, elliptic, cys- toliths lacking. Inflorescences in loose terminal spikes, the bracts and bractlets large and conspicuously colored. Flowers with 5-merous calyx of equal segments; corolla bilabiate, the upper lip bifid, the lower lip 3-lobed; sta- mens 2, the anthers 2-celled; stigma subcapitate. Fruit a clavate capsule, 4-seeded, the seeds borne on hooklike funicles. This is a monotypic genus and is endemic to the Neotropics. It is recognized by its large, showy bracts and its large leaves, which when dried, are shown to lack cystoliths. Louteridium costaricense Radlk. & Donnell Smith, Hot. Gaz. (Crawfordsville) 37: 422. 1904. Figure 12. Shrubs to 1.5 m tall; stems tetragonal, inter nodes be- tween leaf-bearing nodes 1.5-10 cm long, 3-4.5 mm thick, glabrous. Leaves with petioles to 5 cm long, gla- brous; laminae elliptic, 2 1-32 cm long, 9.5-15 cm broad, apically acuminate, basally attenuate, margins entire, both surfaces glabrous, cystoliths dense and prominent on both surfaces. Inflorescences in narrow terminal pani- cles, peduncles to ca. 15 cm long, pedicels 1.5-3 cm long, glabrous, bracts and bracteoles ovate-lanceolate. 8-12 mm long, 3-5 mm broad, glabrous. Flowers with 3-mer- ous calyx, segments loosely enclosing the bud, charta- ceous, narrowly elliptic, apically acute, subequal. 2-4.5 cm long, 1-2 cm broad, cystoliths numerous, venation conspicuous; corolla pale green or yellow, 3.5 cm long, glabrous, lobes apically rounded, 8-9 mm long, 6-7 mm broad, throat prominently gibbous; stamens 2, exserted. anthers ca. 10 mm long. Fruit clavate, 3-3.5 cm long, 7 mm broad, 7 mm thick, glabrous. Megaskepasma erythrochlamys Lindau, Bull. Herb. Boissier 5: 666. 1897. Figure 13. Shrub to 4 m tall; stems subquadrangular. internodes between leaf-bearing nodes 2.5-4 cm long, 6-8 mm thick, puberulous. Leaves with petioles to 4 cm long, petioles puberulous; laminae elliptic, 13-28 cm long, 3.5-13 cm broad, apically acuminate with a rounded, apiculate tip, basally acute, margins entire to crenulate, sparingly stri- gose on the costa and lateral veins of both surfaces, cys- toliths none. Inflorescences in lax, clustered spikes to 20 cm long, 4 cm broad excluding corollas, peduncle to 3.5 cm long, brown puberulous, rachis terete, brown puber- ulous, bracts deep reddish purple drying to reddish brown, solitary, ovate to elliptic, 3.5-4.7 cm long, 1.5-2 cm broad, apically acute, basally acute, sparingly strigulose on both surfaces, mostly restricted to the costa, margins entire; bracteoles elliptic to 20 mm long, 5 mm broad, apically acute, basally attenuate, sparingly strigulose. Flowers with 5-merous calyx, calyx segments equal, lan- ceolate, ca. 10 mm long, 2 mm broad basally, densely strigose; corolla white, tube cylindrical, 20-25 mm long, 2-3 mm broad basally, pilosulous, the upper lip lanceo- 62 FIELDIANA: BOTANY late to 35 mm long, 5 mm broad basally, apically 2-lobed, the lobes subulate, 1.5 mm long, 0.5 mm broad basally, the lower lip elliptic, 35 mm long, 8 mm broad, pilo- sulous outside, apically 3-lobed, the lobes lanceolate, 7 mm long, 2 mm broad basally; stamens exserted ca. 3 cm beyond throat of the corolla, the filaments glabrous, the anthers 4.5 mm long, 1.5 mm broad, basally mu- cronate; ovary 2 mm long, glabrous. Fruits clavate, 30- 35 mm long, 8-10 mm broad, 6-7 mm thick, apically attenuate and mucronate, glabrous; seeds 4, sub- orbiculate, flattened, ca. 7 mm in diameter, glabrous. A native of Venezuela, this plant is cultivated throughout much of the Neotropics. Megaskepasma erythrochlamys is recognized by its shrubby habit, its large leaves which lack cys- toliths, and its flowers with large (3.5-4.7 cm long) reddish purple bracts, and its large (50-60 mm long) white, bilabiate corollas with two exserted stamens having two-celled anthers with equal cells. It most closely resembles the genus Justicia, but differs in its leaves, which lack cystoliths, and its much larger bracts. Mendoncia Veil. Tall, suffrutescent vines. Leaves opposite, petiolate, laminae pinnately veined, margins mostly entire, cys- toliths lacking. Flowers axillary, 1 to several or occa- sionally many in each axil, each flower pedicellate and subtended by 2 flat or keeled bracts, connate prior to anthesis and variably so at anthesis; calyx reduced to an annular, often membranous structure, usually glabrous; corolla 5-merous, the tube funnelform or tubular, usually expanded at the base, the lobes equal or nearly so, re- flexed or spreading, the throat often oblique; stamens 4, didynamous, the filaments short, the anthers linear-lan- ceolate, 2-celled, usually flattened, apically acute, glan- dular-puberulous, basally lobed, the lobes more or less unequal; ovary oblique. Fruit a compressed drupe, often apically oblique, the endocarp pulpy; seeds 1 or 2. This is a genus of approximately 60 species from Central and tropical South America, tropical Af- rica, and Madagascar. In Mendoncia gracilis, M. lindavii, and M. littoralis (of Colombia and Pan- ama), the bracts are completely joined before an- thesis, enclosing the flower in a clear, colorless liquid until it opens. While this has not been re- ported for other species of Mendoncia, the connate habit of the bracts before anthesis suggests that other species may have fluid-enclosed flowers as well. Mendoncia is recognized by its viny habit, its paired, connate bracts partially enclosing five- merous flowers with four didynamous stamens, and its drupaceous fruits. Key to Species of Mendoncia la. Corolla 1 .5 cm long; flowers in axillary umbels M. brenesii Ib. Corolla 2.5 cm long or more; flowers axillary, borne singly or in 2's or 3's 2a 2a. Young stems narrowly winged; hairs on upper leaf surface not arising from stellate bases M. retusa 2b. Young stems not winged; hairs on upper leaf surface arising from stellate bases 3a 3a. Corollas less than 3.5 cm long; bracts less than 2 cm long 4a 3b. Corollas more than 3.5 cm long; bracts more than 2.5 cm long 5a 4a. Petioles 2-4 cm long; bracts obovate to elliptic and basally subcordate M. costaricana 4b. Petioles 1-2 cm long; bracts elliptic to ovate and basally rounded M. gracilis 5a. Bracts oblong-lanceolate; widest below the middle; hairs on stem evenly distributed, somewhat appressed M. landavii 5b. Bracts oblong to oblong-elliptic; widest at the middle; hairs on stem mostly at the angles and spreading . M. tonduzii Mendoncia brenesii Standley & Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18(4): 1239. 1938. Figure 12. Suffrutescent vine; stems scandent, subquadrangular, grooved; internodes between leaf-bearing nodes 4.5-12 cm long, 1.5-3 mm thick, glabrous to sparingly and mi- nutely strigose. Leaves with petioles to 2.6 cm long, pet- ioles sparingly strigose; laminae ovate-elliptic, 6-11.5 cm long, 3-7.5 cm broad, apically acuminate with a mucro 1-2 mm long, basally rounded, margins entire, glabrous above and below except for sparingly minutely strigose on the costa. Inflorescences in axillary umbels of 5-10 flowers radiating from short, flat, rounded, ax- illary spurs, pedicels ca. 1 cm long, sparingly strigose; bracts ovate, 8-12 mm long, 5-7 mm broad, apically BURGER: FLORA COSTARICENSIS 63 acuminate, hasally rounded, sparingly strigose, bracts frequently fused to ca. V* their length. Flowers with a membranous calyx; corolla white, ca. 1 5 mm long, gla- brous, the tube ca. 8 mm long, 3 mm broad, slightly expanded basally, the lobes obcordate, ca. 7 mm broad; stamens included, the anthers 3 mm long, apically cal- carate, basally obtuse. Few collections of this species have been made, but these have been in shady areas of cloud forests of the Cordillera Central, Alajuela Province, at elevations of 1000 to 1500 m. It has also been collected in adjacent Panama at 1 500 to 2000 m. Flowering collections have been made from June through August. Mendoncia brenesii is most easily recognized by its axillary umbels of small, white flowers. It is not easily confused with other Mendoncia species if its axillary umbels are present. Mendoncia costaricana Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 113. 1855. Figure 12. Herbaceous vine; stems subquadrangular, grooved; in- ter nodes between leaf-bearing nodes 5.5-14 cm long, 1.5-2.5 mm thick, strigose. Leaves with petioles to 3.8 cm long, petioles strigose; laminae elliptic to elliptic- ovate, 7.7-13.5 cm long, 3-6.7 cm broad, apically abruptly acuminate and sometimes falcate, the apiculate tip to ca. 2 mm long, basally acute to near rounded, margins entire, sparingly scabrous above with hairs aris- ing from stellate bases, sparingly strigose beneath. Flow- ers axillary, borne singly or in 2's or 3's, pedicels ca. 2 mm long, strigose; bracts obovate to elliptic, 15-18 mm long, 8-10 mm broad, apically obtuse and apiculate, basally subcordate, strigose outside, glabrous within; co- rollas white with dark spots at the throat, to 3 cm long, 8 mm broad at the throat, 4-5 mm broad, ovary gla- brous. Fruit flattened, ovoid with double margins, ca. 20 mm long, 1 3 mm broad, 4 mm thick, puberulous. The seven collections of this species in Costa Rica indicate that it is wide ranging in rain forests at elevations of 500 to 1000 m in the Pacific wa- tershed area of San Jose Province and the Cor- dillera Central portions of Alajuela, Cartage, and Puntarenas provinces. It is also found in adjacent Chiriqui Province of Panama. Flowering collec- tions have been made in July, August, and Sep- tember. Mendoncia costaricana is recognized by its comparatively short (3 cm long), white corollas borne singly or in pairs of axillary flowers, its stri- gose stems, and its relatively long petioles (2 to 4 cm long). It may be confused with M. gracilis, but can be distinguished from it by its longer petioles (2 to 4 cm long vs. 1 to 2 cm) and its obovate to elliptic bracts, which are basally subcordate. Mendoncia gracilis Turrill, Bull. Misc. Inform. 418. 1919. Suffrutescent vine; young stems quadrangular, strigose to sericeous, hairs appressed upward; internodes between leaf-bearing nodes, 4.5-10.5 cm long, 1.2-2.5 mm thick. Leaves with strigose to sericeous petioles to 2 cm long, mostly ca. 1 .5 cm long; laminae elliptic to elliptic ovate, 8.5-12.5 cm long, 4-7 cm broad, apically acuminate, often with a mucro to 2 mm long, basally rounded to obtuse, margins entire, the veins prominent, usually 4 pairs, sparingly strigose above, hairs emerging from stel- late bases, strigose below. Inflorescences axillary, the pedicels borne singly to 3 or 4, 1.2-1.5 cm long, strigose to sericeous; bracts elliptic to ovate, entire, 15-19 mm long, 6-9 mm broad, apiculate, basally rounded, strigose to sericeous outside. Flowers with the calyx reduced to an entire membranous cup ca. 1 mm deep, glabrous; corolla white, funnelform, ca. 3 cm long, basally 3 mm broad expanding to 6 mm broad at the throat, the throat streaked with reddish brown, oblique, the lobes 5-6 mm long, 4-5 mm broad, shallowly emarginate, the tube glandular puberulous inside below the stamens, remain- der of corolla glabrous; stamens included. Fruits ovoid, ca. 1.7 cm long, purple-black and fleshy at maturity. This species, originally described from along trails and openings in the rain forests of the Pacific slope and intermountain areas of Colombia, has also been collected in similar habitats in central Panama. One collection, Durkee 83-4 (F, GRI), has been made in the Golfo Dulce area, Puntarenas Province of Costa Rica, in mature rain forest along the new road to Jimenez from the Interamerican Highway near the Rio Olla. The flowering collec- tion was made in January. Mendoncia gracilis can be recognized by its white corollas about 3 cm long borne singly to three or four in leaf axils, elliptic bracts ( 1 5 to 1 9 mm long), basally rounded with upwardly appressed seri- ceous hairs; young stems and leaves also have up- wardly appressed hairs, and petioles are 1 to 2 cm long. It may be confused with M. costaricana, which differs in its longer petioles (2 to 4 cm long) and its obovate to elliptic bracts, which are basally subcordate. Mendoncia lindavii Rusby, Mem. Torrey Bot. Club 4: 241. 1895. Figure 12. Suffrutescent vine; younger stems quadrangular, in- ternodes between leaf-bearing nodes, 3.5-17 cm long, 1.5-3 mm thick, wooly hairs rust-colored. Leaves peti- olate, petioles to 3 cm long, wooly with rust-colored hairs; laminae elliptic, 5.8-11.5 cm long, 3.3-6.4 cm 64 FIELDIANA: BOTANY broad, apically acuminate, hasally obtuse to rounded, margins entire, sericeous above with hairs rising from stellate bases, sericeous beneath, but hairs more dense and prostrate. Flowers axillary, pedicellate, borne singly or in pairs, pedicels to 3 cm long, pubescence that of the petiole; mature bracts oblong-lanceolate, subfalcate, 3.2- 4 cm long, 0.8-1.2 cm broad, apically attenuate, basally rounded, wooly with rust-colored hairs; corolla red, pink, or white, ftmnelform, 4-5 cm long, 6 mm broad basally expanding to 8 mm broad at the throat, glabrous, the 5 equal lobes to 6 mm long, 4.5 mm broad, apically retuse to rounded; stamens attached at about the mid- point of the corolla tube, anthers 10-13 mm long, 1.5 mm broad, filaments 2 mm long; ovary velutinous, ca. 4 mm long, 2.5 mm broad, style to 5 cm long, stigma forked. Fruits flattened, obovate, to 20 mm long, 1 1 mm broad, 8 mm thick, apically oblique and terminated by a portion of the persistent style, dark brown (dull purple when fresh) and densely puberulous. Ranging from Belize and Guatemala to Bolivia, this species has been collected in rain forest areas of Costa Rica at elevations of 500 to 1000 m on the Pacific Watershed only in Puntarenas Province near Panama and San Jose Province in the vicinity of General Viejo (El General). Flowering collec- tions have been made in June and January. Mendoncia lindavii is recognized by its large red, pink, or white corollas (4 to 5 cm long), its large (3.2 to 4 cm long), oblong-lanceolate, subfalcate bracts with conspicuous, reddish brown wooly hairs, and its large (to 20 mm long) dark brown or purplish, obovate fruits. It may be confused with M. retusa, which also has large flowers, but can be distinguished from it by its wooly stems (vs. strigose), its nonwinged stems (vs. narrowly winged), and its wooly, oblong-lanceolate bracts (vs. mostly glabrous, oblong). Mendoncia retusa Turrill, Bull. Misc. Inform. 423. 1919. Figure 12. Suffrutescent vines with narrowly winged, quadran- gular stems; internodes between leaf-bearing nodes 4- 14 cm long, 1-2.5 mm thick, sparsely strigose to gla- brous. Leaves petiolate, petioles to 5.2 cm long, sparsely strigose to puberulous; laminae elliptic, 7-15 cm long, 2.5-7.5 cm broad, apically acuminate and mucronulate, basally acute, obtuse or rounded, margins undulate, sparsely strigose on costa beneath. Flowers axillary, ped- icellate, borne singly, pedicels 1.75-3 cm long, sparingly strigose to glabrous; bracts green, elliptic to elliptic-ob- long, 1.5-3.0 cm long, 1-2 cm broad, apically retuse or rounded, mucronate, basally rounded, mostly glabrous; corolla white with a purplish throat, 4-5 cm long, gla- brous, lobes equal, 5-6 mm long, apically obtuse. Fruit irregularly obovate to 19 mm long, 9 mm broad, 8 mm thick, puberulous. Found in lowland rain forests ranging from Mexico (Chiapas) through Panama, this species is one of the most commonly found in Central Amer- ica, although, again, few collections have been made in Costa Rica. However, these collections are from both the Caribbean and Pacific areas at elevations of 15 to 100 m. Flowering collections have been made in June and October. Mendoncia retusa is recognized by its narrowly winged, quadrangular stems, its large (4-5 cm long), white corollas with purplish throats, and its elliptic to elliptic-oblong bracts, which are apically retuse or rounded and mucronate. The bracts are some- what variable, and in those specimens with more oblong bracts, there may be confusion with M. lindavii, which differs in its rust-colored, wooly hairs, its lack of winged stems, and its red, pink, or white corollas. Mendoncia tonduzii Turrill, Bull. Misc. Inform. 413. 1919. Figure 12. Suftrutescent vines with quadrangular stems; inter- nodes between leaf-bearing nodes 9-12 cm long, 2-3 mm thick, densely pilose, hairs tawny and mostly restricted to the angles. Leaves petiolate, petioles to 2 cm long, densely pilose tawny hairs; laminae elliptic to elliptic- ovate, 7-13 cm long, 5-8.3 cm broad, apically acumi- nate, often with an apicule to 2 mm long, basally acute to rounded, margins entire, pilose hairs with stellate bas- es above, more densely pilose below. Flowers solitary or in pairs in the axils of the upper leaves, pedicellate, ped- icels 2-4 cm long, pilose; bracts oblong to oblong-elliptic; ca. 4 cm long, 1.5 cm broad, apically rounded and mu- cronate, basally rounded, densely pilose, hairs tawny; corolla white, ca. 5 cm long, 8 mm broad at the oblique throat, the lobes equal, emarginate, spreading, ca. 8 mm long, 5-6 mm broad, glabrous. Fruit purplish black at maturity, ca. 2 cm long, 1 cm broad, 6 mm thick, densely puberulous. This uncommon species has been collected in forests at 500 to 1000 m elevation of the Tilaran area, Guanacaste Province, Brenes 12675 (F), and Tucurrique, Cartage Province, Tonduz 12946 (us, holotype). Three collections have also been made in similar habitats in Code and Colon provinces of Panama. Mendoncia tonduzii is recognized by its dense, tawny pilose hairs, its oblong to oblong-ovate bracts, 4 cm long, apically rounded and mucro- nate, and laminae with pilose hairs on stellate bas- es. This distinguishes it from the more common M. lindavii, which it most closely resembles. Nelsonia R. Br. Herbs, stems softly villous. Leaves sessile; laminae with entire margins. Inflorescences of dense, terminal BURGER: FLORA COSTARICENSIS 65 and axillary spikes; bracts imbricate, 1 per flower. Flow- ers with 4-merous calyx; corolla bilabiate; stamens 2, anthers 2-celled, the cells parallel, each cell globose; stig- ma 2-lobed. Fruits a 2-chambered capsule, capsules ob- long; seeds borne on papilliform funicles. There is only one species described for this ge- nus, which was first collected in Java. The genus was named for David Nelson, a gardner who ac- companied Cook on his last voyage. Nelsonia canescens (Lam.) Spreng., Syst. veg. ed. 16. 1:42. 1825. Figure 13. Herbs, decumbent to ascending with narrow, branch- ing stems to 40 cm long; internodes between leaf-bearing nodes 1-5 cm long, 1-2 mm thick, villous to wooly. Leaves sessile; laminae ovate, 1-8 cm long, 0.5-3 cm broad, apically acute to obtuse, basally acute to obtuse, margins entire, cystoliths lacking, villous above and be- neath. Inflorescences in dense spikes to 6.5 cm long, 5.5 mm broad excluding corollas, pedunculate or sessile, mostly axillary, flowers sessile to pedicels ca. 1 mm long; bracts imbricate, 1 per flower, ovate, 4-6 mm long, 4- 5 mm broad, apically acuminate, velutinous. Flowers with 4-merous calyx, segments ca. 4 mm long, unequal, posterior segment ovate, 2.5 mm broad, anterior seg- ment lanceolate, 1.5 mm broad, lateral segments lan- ceolate, 1 mm broad; corolla deep purple to blue, bila- biate, the tube ca. 3 mm long, upper lip 2-lobed, ca. 1 mm long, lower lip 3-lobed ca. 1 mm long. Fruit sessile, ca. 5 mm long, ca. 10 seeds per locule. Found in disturbed habitats, especially in moist, sandy areas along roadsides, trails, and streambeds, this species occurs in Mexico, Central America, northern South America, tropical Africa, India to Indo-China, Malay Peninsula, and Australia. Col- lected in Costa Rica at elevations of 150 to 1500 m in Alejuela, Guanacaste, and San Jose prov- inces. Flowering collections have been made in February through May. Nelsonia canescens is recognized by its decum- bent stems with numerous, compact spikes, ve- lutinous, imbricate bracts subtending minute flow- ers with bilabiate bluish corollas, and two stamens with two equal-celled anthers. Its capsules are ses- sile, and its seeds lack the hooklike funicles which serve to eject the seeds of most Acanthaceae. Cer- tain specimens may resemble Elytraria imbricata, which has larger leaves (6 to 20 cm vs. 1 to 8 cm long) and stiff, awned bracts (vs. soft, awnless bracts). Odontonema Nees REFERENCE— V. M. Baum and J. L. Reveal, A new proposal to conserve 8037 Odontonema (Acanthaceae). Taxon 31: 757-759. 1982. Herbs or shrubs. Leaves sessile to petiolate; laminae often large with cystoliths visible on both surfaces of dried specimens. Inflorescences in terminal panicles, thyrses, or verticillate inflorescences, bracts inconspic- uous. Flowers with 5-merous, equally segmented calyx; corolla bilabiate, the upper lip 2-lobed, the lower lip 3-lobed; stamens 2, exserted or extending to the mouth, the anthers 2-celled with the cells parallel and basally blunt, staminodes 2. Fruits in clavate capsules, seeds 4 on hooklike funicles. This is a genus of 29 species, all endemic to the Neotropics. Odontonema tub! for me (Bertol.) Kuntze, Re vis. gen. pi. 2: 493. 1891. Justicia tubiformis Bertol., Novi Comment. Acad. Sci. Inst. Bononiensis 4: 405. 1840. ThyrsacanthusflagellumOerst.,Vi- densk. Meddel. Dansk Naturhist. Foren. Kj0- benhavn 1854: 146. 1855. Odontonema flagel- lum (Oerst.) Kuntze, Revis. gen. pi. 2: 494. 1891. Thyrsacanthus strictus Nees in DC., Prodr. 11: 324. 1947. Odontonema strictum (Nees) Kuntze, Revis. gen. pi. 2: 494. 1891. Figure 13. Suffrutescent herbs to shrubs 2.5 m tall; internodes between leaf-bearing nodes 1.5-9 cm long, 1 .5-3.25 mm thick, puberulous. Leaves sessile to petiolate, petioles to 3 cm long, puberulous to subglabrous; laminae elliptic to broadly elliptic-ovate, 1 1-36 cm long, 3.5-1 1 cm broad, apically acuminate, basally attenuate, margins crenate to entire (often on the same plant), cystoliths more vis- ible on upper surface, obscure beneath, glabrous. Inflo- rescences verticellate to 35 cm long, 1.5 cm broad ex- cluding corollas or fruits, peduncles 1.5-7 cm long, puberulous, rachis puberulous to tomentose, flowers pedicellate with 1 to 10 pedicels per cluster, pedicels puberulous; bracts and bracteoles inconspicuous, subu- late, 3-10 mm long, 1-2 mm broad, puberulous. Flowers with 5-merous calyx, calyx lobes equal, subulate, 2-4 mm long, 0.6-1 mm broad, minutely puberulous; corolla pink or red, funnelform, 15-30 mm long, 4-5.5 mm broad at the throat, 1-3 mm broad at the base, glabrous, upper lip 7-8 mm long, 4.5-5 mm broad, bilobed, lobes apically acute, 3.5-4 mm long, lower lip 7-8 mm broad, lobes elliptic, apically rounded, 3-4 mm broad, all lobes with glandular hairs inside; stamens exserted, anthers ca. 2.5 mm long, filaments glabrous, staminodes 1-3.5 mm long. Fruits clavate, 2-3 cm long, 4 mm broad, 3-4 mm thick, glabrous, seeds 4, ovate with an oblique base, flattened, 5-6 mm long, 3.5-4.5 mm broad, 1 mm thick, surface densely rugose. This species is found in light gaps, clearings, and along streams and trails in rain forests of lower elevations to rain forest transition at higher ele- vations of Mexico, Central America, and Panama. In Costa Rica it has been collected in both the Atlantic and Pacific watersheds from all provinces except Guanacaste at elevations of 50 to 1200 m. 66 FIELDIANA: BOTANY Flowering collections have been made in July and October through April. This is a wide-ranging species which is subject to much variation in leaf, flower, and fruit size, as well as pubescence of the rachis and number of flowers per cymule. Many of the early species were erected on these bases, but additional collections show intergradation in all of these characters. The only character that appears unique to Odontonema Jlagellum (Oerst.) Kuntze is the narrowed, flagellate tip of the inflo- rescence which I believe is an early flowering stage of O. tubiforme. Odontonema tubiforme is recognized by its ver- ticillate inflorescences with near-equal, funnel- form corollas that are pink or red with two exserted stamens with two-celled anthers and two short staminodes. It is most frequently confused with Razisea species, but can be distinguished from them by its staminodes, two-celled anthers, and more broadly funnelform corollas. Poikilacanthus Lindau Herbs or shrubs. Leaves of a pair unequal, petiolate. Flowers borne in small terminal or axillary, bracted spikes or heads. Flowers with 5-merous calyx; corolla narrow, bilabiate; stamens 2, anthers 2-cclled. the cells unequally attached, the cells rounded or ending in a short spur; pollen elliptic, polyporate. Fruits a clavate, 2-chambered capsule with 4 seeds, each borne on hooklike retinacula. subulate, 7-9 mm long, 1.3-1.5 mm broad, ciliolate to- ward apex; corolla lavender or purplish, very narrowly funnelform, 6.5-7.8 cm long, 4-5 mm broad at the throat, 1.5 mm broad basally, glandular-puberulous, upper lip 30-35 mm long, 7-9 mm broad, apical K minutely 2-lobed, lower lip 22-27 mm long, 9-12 mm broad at ca. midpoint, apically 3-lobed, lobes ca. 2 mm long, 3 mm broad; stamens exserted to ca. the tip of the upper lip or just beyond, anther cells ca. 2.25 mm long, lobes obtuse. Fruits clavate, 12-15 mm long, 3.5-5 mm broad, 3.1-3.5 mm thick, sparingly puberulous. Found in gaps or clearings, along trails and streams in cloud forest areas of Mexico (Chiapas), Honduras, Guatemala, Nicaragua, and Costa Rica. Collected in Costa Rica in cloud forest areas of the Cordilleras Tilaran, Central, and Talamanca at elevations of 1000 to 1750 m, mostly on the Pacific watershed, but occasionally on the Carib- bean as well. Flowering collections have been made in February, March, and September through No- vember. Poikilacanthus macranthus is recognized by its unequal leaf pairs, its lavender, narrowly funnel- form bilabiate corolla (6.5 to 7.8 cm long), two exserted stamens with widely superposed anther cells, and elliptic, polyporate pollen. It is most easily confused with species of Justicia, but can usually be distinguished from these in its unequal leaf pairs (excluding Justicia trichotoma), its large, narrowly funnelform corollas, and its polyporate pollen (vs. two-, rarely three-, or four-, porate). This is a Neotropical genus of about 12 species found in Central America and tropical South America. It is similar to the genus Justicia, but differs from it in its unequal leaf pairs and its pollen morphology. Poikilacanthus macranthus Lindau, Bull. Herb. Boissier 3: 481. 1895. Figure 13. Shrubs to 3 m tall; internodes between leaf-bearing nodes 2-6.5 cm long, 1-2.25 mm thick, quadrangular, puberulence of appressed hairs varying from a few hairs restricted to the angles to dense. Leaves of a pair unequal, petioles 0.5-2.6 cm long, pubescence that of the stem; laminae ovate to ovate-elliptic, 3-7 (8.5) cm long 1.5- 3.5 (4) cm broad, apically acuminate, basally obtuse to rounded and often oblique, margins entire, appressed hairs restricted to costa and veins on both surfaces to pilose, cystoliths dense and prominent on both surfaces. Inflorescences in terminal, sessile, headlike clusters of 2-4 flowers; bracts oblong to spatulate, 6-9 mm long, 1.5-3 mm broad, ciliate, bracteoles similar, but smaller. Flowers with 5-merous calyx, calyx segments equal, Pseuderanthemum Radlk. Herbs or small shrubs. Leaves commonly petiolate; laminae elliptic, ovate, or lanceolate, cystoliths generally visible on both surfaces of dried specimens. Inflores- cences in terminal or axillary racemes, spikes or panicles; 1 bract and 2 bracteoles subtending each flower or flower cluster, bracts and bracteoles inconspicuous. Flowers with 4 or 5 subulate calyx segments; corolla white, blue, pur- plish or pink, salverform, the tube narrow, the limb spreading, the 5 lobes subequal; stamens 2, included or exserted, inserted at or above the middle of the corolla tube; anther 2-celled, the cells basally obtuse or acute; staminodes 2, minute. Fruits in clavate capsules, seeds 2 or 4 on hooklike retinacula. This is a genus of about 1 20 species from the New and Old World tropics. The flowers of this genus resemble those of the genus Chamaeran- themum which differs in its four stamens that are included, with the posterior pair either one-celled or sterile. BURGER: FLORA COSTARICENSIS 67 Key to Species of Pseuderanthemum la. Inflorescence a cymose panicle (thyrse); rachis of inflorescence glabrous P. atropurpureum Ib. Inflorescence a raceme, spike, or spicate panicle; rachis glandular-puberulous or puberulous . . 2a 2a. Petioles conspicuously winged, laminae bases long-attenuate; stamens exserted . . . P. cuspidatum 2b. Petioles not winged, laminae bases acute or attenuate, but not long-attenuate; stamens included 3a 3a. Flowers in opposite fascicles of 2-3 or more; laminae mostly more than 9 cm long . . . . P. pittieri 3b. Flowers borne singly and opposite, or if in pairs, alternate; laminae mostly less than 9 cm long . . . 4a 4a. Flowers borne singly and opposite; capsules 16-19 mm long, puberulous P. praecox 4b. Flowers borne singly or in pairs, mostly alternate; capsules 1 1-15 mm long, glabrous . . . . P. standleyi Pseuderanthemum atropurpureum (Bull) Bailey, Gentes Herb. 1: 130. 1923. Eranthemum atro- purpureum Bull. Card. Chron. 1 : 6 19. 1 875. Fig- ure 13. Shrubs or small trees to 4 m tall; internodes between leaf-bearing nodes 2.5-5 cm long, 2-4 mm thick, te- trangular. glabrous. Leaves petiolate, petioles 0.5-1 .8 cm long, glabrous except for a tuft of hair at the stipular ring; laminae elliptic to ovate, 4.5-14.5 cm long, 4.5- 6.5 cm broad, apically acute, some apiculate, basally attenuate, margins entire, greenish beneath and dark pur- ple above, glabrous on both surfaces, cystoliths dense and near punctiform on both surfaces. Inflorescences in terminal and axillary cymose panicles with the flowers in subsessile fascicles of mostly 3 flowers; pedicels, pe- duncles and rachises glabrous, pedicels to 5 mm long, peduncles to 7 cm long, rachises to 1 3 cm long, bracts elongate, lower bracts ca. 5-7 mm long, 2-3 mm broad, upper bracts progressively smaller, all ciliolate. Flowers with 5-merous calyx, the lobes equal, subulate, 2-3 mm long, 0.75-1 mm broad, puberulous; corolla white with magenta spots in the throat, the tube cylindrical, 10-12 mm long, 2-2.5 mm broad basally, the limb 20-22 mm broad, the lobes elliptic, 8-10 mm long, 5-8 mm broad, apically rounded, ciliolate; stamens exserted, filaments attached just below the mouth, glabrous anther cells equally attached, ca. 1.5 mm long, basally acute, stam- inodes 1.5-2 mm long. Fruits not observed. This species, frequently cultivated in tropical America, is believed to be a native of Polynesia. It sometimes escapes from cultivation. Flowering collections have been made in January and No- vember. Pseuderanthemum atropurpureum is rec- ognized by its cymose panicles and purplish leaves, which are frequently variegated with green, white, and pink. These features make it easily distin- guishable from other species of the genus. Pseuderanthemum cuspidatum (Nees) Radlk., Sit- zungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Munchen 13: 286. 1883. Eranthemum cuspidatum Nees in Bentham, PI. hartw. 148. 1839. Figure 13. Herbs to 60 cm tall; stems erect; internodes between leaf-bearing nodes 1.3-7 cm long, 0.5-3.5 mm thick, quadrangular, puberulous along 2 lines. Leaves petiolate, petioles subsessile to ca. 3.5 cm long, glabrous to spar- ingly puberulous along 2 lines; laminae ovate, 8-19 cm long, 3.2-8.2 cm broad, apically acuminate, basally long- attenuate, margins entire, glabrous to very sparingly sca- brous on both surfaces, cystoliths dense and prominent on both surfaces. Inflorescences in lax terminal and ax- illary spikes frequently branched from the base, to 16 cm long, 4 mm broad excluding corollas, the rachis glan- dular-puberulous; lowermost bracts subtending the in- florescence, suborbicular, to 2 cm long, 1.5 cm broad, floral bracts subulate, 2-3 mm long, 0.75-1 mm broad, ciliolate; bracteoles similar, though smaller; flowers ses- sile, paired or in pseudowhorls of up to 5 flowers. Flowers with a 5-merous calyx, calyx lobes equal, subulate, 2.5- 3.5 mm long, 0.5-0.75 mm broad, glabrous to sparingly puberulous plus occasional glandular hairs; corolla pur- plish, glabrous, tube cylindrical 1.5-2.8 cm long, 0.8-1 mm broad, lobes of the limb elliptic, 8.5-9 mm long, 4.5-5 mm broad, apically obtuse; stamens barely exsert- ed, filament ca. 2.5 mm long, staminodes 0.5 mm long, anther cells 1.25 mm long. Fruits clavate, 11-15 mm long, 2-2.5 mm broad, 1.5-2 mm thick, glabrous, seeds 4, ovate, 2.5-3 mm long, 2-2.5 mm broad, rugose. Found in shady, moist, forested areas from southern Mexico to northwestern South America, this species is wide ranging and not uncommon as suggested by the relatively numerous collections. In Costa Rica it is found in both major watersheds from elevations of about 700 to 1575 m. It has not been collected in Guanacaste, Heredia, or Li- mon provinces. Flowering collections have been made in May and August through December. Pseuderanthemum cuspidatum is recognized by its often-branched, terminal and axillary spikes with lax flowers in pairs or pseudowhorls, glabrous to subglabrous five-merous calyx, and its long, conspicuously winged petioles (or long-attenuate laminae). It is most easily confused with P. pittieri, which differs in its more pronounced glandular- puberulous calyx, its elliptic laminae with more abruptly attenuate bases, and its shorter corolla tubes (7.5 to 1 1 mm long vs. 15 to 28 mm long) with stamens included. 68 FIELDIANA: BOTANY Pseuderanthemum pittieri Leonard, Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1245. 1938. Figure 14. Herbs or small shrubs to 80 cm tall, stems erect; in- ter nodes between leaf-bearing nodes 1-7 cm long, 2-4 mm thick, subquadrangular, puberulous. Leaves with short petioles to 6 mm long, glabrous to sparingly puber- ulous; laminae elliptic, 8.5-19 cm long, 3-8.5 cm broad, apically acuminate, basally attenuate or acute, margins undulate, cystoliths moderately dense and visible on both surfaces, glabrous above, purplish puberulence on costa and main veins beneath. Inflorescences in terminal and axillary spikelike racemes, frequently branched; flowers opposite, frequently in fascicles of 2-3, pedicels short, ca. 0.5 mm long, puberulous-glandular; bracts subulate, 2.5-3 mm long, 1-1.5 broad basally, puberulous plus occasional glandular hairs. Flowers with a 5-merous ca- lyx, calyx lobes equal, long-subulate, 3-4 mm long, 0.4- 0.5 mm broad basally, glandular- puberulous: corolla white, pink, or lavender, the tube subcylindrical, 7.5-1 1 mm long, 1-1.5 mm broad basally and up to ca. mid- point where the stamens are attached and then expanding to 1.5-2 mm broad at the limb, tube glandular-puber- ulous, lobes of the limb 10-11 mm long, 4-5 mm broad, apically obtuse; stamens included, attached at midpoint of corolla tube, filament ca. 2 mm long, staminodes 0.5 mm long, anther cells 1 .25 mm long. Fruits clavate, ca. 1 8 mm long, 2.8 mm broad, 2.2 mm thick, apically acute, puberulous; seeds 4, oval, flattened, ca. 3 mm long, 2.5 mm broad, tubercled. Found growing in shade of moist lowland forest areas of primary and secondary forests in the Golfo Dulce area and the Osa Peninsula at elevations of 50 to 200 m. Flowering collections have been made in December and January. Pseuderanthemum pittieri is recognized by its comparatively large, elliptic leaves with short-at- tenuate lamina bases, spikelike racemes, glandu- lar-puberulous calyx lobes and corollas, and short, subcylindric corolla tubes (7.5-1 1 mm) with in- cluded stamens. It is most easily confused with the more widespread P. cuspidatum, which differs in its conspicuously winged petioles (long-attenuate laminae), true spikes, glabrous to subglabrous ca- lyx lobes and corollas, and longer cylindric corolla tubes (15-28 mm) with exserted stamens. Pseuderanthemum praecox (Bentham) Leonard, J. Wash. Acad. Sci. 31: 99. 1941. Eranthemum praecox Benth., PI. hartw. 291. 1848. Figure 14. Herbs to 30 cm tall; Intel-nodes between leaf-bearing nodes 1.8-5 cm long, 1-1.5 mm thick, quadrangular, puberulous mostly along 2 lines. Leaves with petioles 4- 10 mm long, glabrous to sparingly puberulous; laminae elliptic to elliptic-ovate, 1 .5-8 cm long, 0.8-2.5 cm broad, apically acuminate, basally attenuate, margins undulate, cystoliths dense and visible on both surfaces, sparingly strigose above, strigose on costa and main veins below. Inflorescences in short terminal and axillary spikes 1- 6.5 cm long, flowers opposite, borne singly, bracts subu- late, 1.5-3 mm long, 0.5-0.75 mm broad, sparingly pu- berulous, bracteoles similar though smaller; rachises glandular-puberulous. Flowers with a 5-merous calyx, calyx lobes long-subulate, 2.5-3.5 mm long at early an- thesis, 5-6 mm long in fruit, 0.5-0.75 mm broad, glan- dular-puberulous to mostly puberulous; corolla laven- der, pink or bright rose, glabrous outside, the tube cylindrical, 16-19 mm long, ca. 1.5 mm broad, lobes elliptic, 8-12 mm long, 3.5-5 mm broad, apically acute; stamens included, filaments 1.75-2 mm long, stami- nodes ca. 0.5 mm long, anthers 1.5-2 mm long. Fruits clavate, 15-18 mm long, ca. 3 mm broad, 4 mm thick, puberulous plus occasional glandular hairs on some; seeds suborbicular, ca. 2 mm in diameter, rugose. In Costa Rica this species is found in or adjacent to premontane or montane rain forests of the Cor- dilleras Tilaran and Central at elevations of 1450 to 2000 m, where flowering collections have been made in October, November, February, and April. This species has a range extending to northern and central Mexico. Pseuderanthemum praecox is recognized by its comparatively small, elliptic laminae that are mostly 2 to 6 cm long, its lax spicate inflorescences with flowers opposite and borne singly, and its lavender to bright rose corollas with cylindrical tubes (16 to 19 mm long) and stamens included. It most clearly resembles P. standleyi, which dif- fers in its inflorescences with flowers solitary or in pairs mostly alternating on the rachis, slightly shorter corolla tube (ca. 1 5 mm long vs. 1 6 to 19 mm long), its staminodes that are slightly longer than the filaments (vs. much shorter), and its shorter, glabrous capsules (10 to 15 mm vs. 15 to 1 8 mm long and puberulous). Pseuderanthemum standleyi Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:1 246. 1 938. Figure 14. Herbs to 1 m tall; Intel-nodes between leaf-bearing nodes 8-20 mm long, 0.9-1.1 mm thick, puberulous mostly along 2 lines, terete. Leaves with petioles 3-6 mm long, puberulous; laminae ovate to lanceolate, 3-8 (9.5) cm long, 1.2-5 cm broad, apically acuminate, ba- sally attenuate, margins entire to undulate, glabrous to pilose above, puberulous beneath, mostly at the costa and veins, the cystoliths dense, moderately prominent on both surfaces. Inflorescences in terminal spicate pan- icles and in axillary spikes, the spikes lax, to 9 cm long, the rachises glandular puberulous; flowers solitary or in pairs, mostly alternate; bracts and bracteoles subulate, 1.5-3 mm long, ca. 0.5 mm broad basally, puberulous- glandular. Flowers with 5-merous calyx, calyx segments linear-subulate, 5-6 mm long, puberulous-glandular; co- rolla purplish, the tube cylindrical ca. 15 mm long, 1 mm broad, sparingly and minutely puberulous, lobes of BURGER: FLORA COSTARICENSIS 69 the limb elliptic, ca. 7 mm long, 3.5 mm broad, apically obtuse; stamens included, the staminodes slightly longer than the filaments. Fruits clavate, 1 1-15 mm long, 2.5- 3 mm broad, 1.5-2 mm thick, glabrous; seeds 4, oblique, ca. 2.5 mm long, 2 mm broad, tuberculate. Found in moist forests and thickets of the Pacific slopes of the Cordilleras Central and Talamanca, adjacent Panama, and up to west-central Mexico. Flowering collections have been made in Decem- ber and May. Pseuderanthemum standleyi is recognized by its lax spicate inflorescences with flowers solitary or in pairs, mostly alternate on the rachis, its purplish corolla with a tube about 15 mm long, and its staminodes, which equal or exceed the length of the filaments. It most closely resembles P. praecox, which has the flowers borne singly and opposite, longer corollas ( 1 6 to 1 9 mm), much shorter stam- inodes, and longer, puberulous capsules (15 to 18 mm long vs. glabrous, 10 to 15 mm long). Razisea Oerst. Shrubs or herbs to 2.5 m tall. Leaves petiolate except those subtending the inflorescence which are frequently sessile or subsessile; laminae elliptic to ovate or lanceo- late, cystoliths dense and prominent on both surfaces of dried specimens. Inflorescences racemose to thyrsoid, mostly terminal, sometimes branched; flowers sessile or pedicellate; bracts and bracteoles small and inconspic- uous. Flowers with 5-merous calyx, calyx lobes equal; corolla bilabiate, the upper lip narrow, the lower lip broad and shallowly trilobate; stamens 2, exserted, the fila- ments flattened, the anthers 1 -celled; style exserted, the stigma 2-parted. Fruit a capsule, clavate, 4-seeded. This is a Central American genus of three species. Two of the species are endemic to Costa Rica, and the third, R. spicata, is wide ranging over all of Central America and Panama. Razisea is very similar to the genus Kalbreyeriella of Panama and Colombia, which differs in its beaked corollas prior to expansion and its calyx lobes which are two to three times longer than the bracts and bracteoles. Razisea is also very similar to the genus Hanstei- nia, which differs in its corolla tube, which is usu- ally abruptly expanded above the ovary, and in its paniculate inflorescences. Razisea is also dif- ficult to distinguish from the genus Habracanthns. The Costa Rican species of Habracanthus has a blue corolla, but many of the South American species have corollas of the same color as Razisea species (i.e., red or yellow). It is quite possible that a more comprehensive study of these genera will bring them into synonymy. Razisea citrina is known only from two collec- tions, and R. wilburii, while relatively abundant where found, is apparently restricted to the Carib- bean lowlands of Costa Rica. Key to Species of Razisea la. Corolla lemon yellow; inflorescence a broad thyrse (cymose panicles) with pedunculate, compound cymes R. citrina Ib. Corolla bright red; inflorescence racemose with opposite sessile fascicles of 1-10 flowers 2a 2a. Corolla 4—6 cm long R. spicata 2b. Corolla 2.5-3.3 cm long R. wilburii Razisea citrina D. Gibson, Fieldiana, Bot. 34: 78. 1972. Figure 14. Suftrutescent plants to 2 m tall. Inter nodes between leaf-bearing nodes, 2.5-4.5 cm long, 1.5-3 mm thick, glabrous. Leaves petiolate, petioles 1.5-5.5 cm long, gla- brous, leaves subtending inflorescence sessile or subses- sile; laminae lanceolate, lanceolate-ovate or elliptic, 10- 19.5 cm long, 5-9 cm broad, apically acuminate, basally acute to attenuate, glabrous on both surfaces, cystoliths dense and prominent on both surfaces. Inflorescences terminal, in broad thyrses (cymose panicles) 2-3-flow- ered; pedicels 5-15 mm long, glabrous; bracts subulate, 3-5 mm long, glabrous, bracteoles similar but smaller. Flowers with 5-merous calyx, calyx lobes long-subulate 7-9 mm long, glabrous; corolla lemon yellow, glabrous, 3.5-4.5 cm long, the tube ascending and curved outward, basally 3 mm broad, expanding to 5-11 mm broad at the mouth, upper lip erect, linear, minutely fimbriate, lower lip when open, forming an oblique angle from the upper lip, shallowly trilobate, the lobes ca. equal; sta- mens exserted 12-15 mm beyond the tip of the upper lip, anthers ca. 5 mm long. Fruits clavate, ca. 20-25 mm long, 4-4.5 mm broad, 2.5 mm thick, glabrous, seeds 4, ovate, 2-2.5 mm long, 1.5-2 mm broad, papillate. I have seen only two specimens of this species: the holotype, Jimenez 2475 (F), collected in Oc- tober in cloud forest area between Agua Buena 70 FIELDIANA: BOTANY and San Vito (San Vito de Java), Puntarenas Prov- ince, 1200 m altitude, and Weaver et al. 1775 (DUKE), collected in August at 1000 m altitude, where it was described as a common shrub along a stream in the forest. Both specimens were in flower. Razisea citrina is recognized by its broad ter- minal thyrses (cymose panicles) with lemon-yel- low corollas. It is easily distinguished from other species of Razisea by these two features. Razisea spicata Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjebenhavn 1854: 142. 1855. Kolobochilus leiorhachis Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and in Pitt., Prim. fl. costaric. 2: 308. 1900. Figure 14. Shrub to 2.5 m tall; internodes between leaf-bearing nodes 2-9 cm long, 2-4 mm thick, quadrangular, gla- brous to infrequently, but densely, pilose. Leaves peti- olate, petioles 1.0-3.5 (8.5) cm long, glabrous; laminae elliptic to ovate, 9.5-29 cm long, 3.5-8.5 cm broad, api- cally acuminate, basally attenuate, acute, obtuse, or rounded, margins entire to slightly crenate, glabrous above, glabrous to sparingly puberulous along the costa and veins beneath, cystoliths dense and prominent on both surfaces. Inflorescences in terminal racemes to 30 cm long, rachis puberulous to subglabrous, flowers nu- merous, solitary or in fascicles of 2-1 0 at opposite nodes, pedicels 5-10 mm long, puberulous, bracts subulate, 5- 10 mm long, 1-2 mm broad, sparingly puberulous, brac- teoles similar, but smaller. Flowers with 5-merous calyx, calyx lobes linear, 4.5-9 mm long, ca. 0.5 mm broad, puberulous; corolla bright red, funnelform, ascending, slightly curved to one side, 4-6.2 cm long, glabrous, ca. 9 mm broad at the mouth, 1.5-2 mm broad basally, upper lip erect, linear, to ca. 8 mm long, 4 mm broad basally, apically rounded, the lower lip with 3 small, acute lobes at the tip, the lobes equal, ca. 1 mm long, the lower lip, when open, forming an oblique angle from the upper lip; stamens exserted 10-20 mm beyond the tip of the upper lip, the filaments to ca. 4.5 cm long, glabrous, anther cell 3-4 mm long, the bases rounded. Fruits clavate, 20-22 mm long, 3.5-4 mm broad, 3.0- 3.5 mm thick, glabrous; seeds suborbicular, ca. 2.5 mm in diameter, tuberculate. Found growing at the edges of light gaps and clearings in lowland rain forests and cloud forests, this species extends from Colombia to Guatemala. It has been collected from all provinces in Costa Rica at elevations of 50 to 1800 m. Flowering collections have been made from June through March. Razisea spicata is recognized by its terminal ra- cemes with pedicellate flowers either solitary or in fascicles of two to 1 0 at opposite nodes, its con- spicuous bright red corollas, which are 4 to 6.2 cm long, funnelform, ascending, and slightly curved outward from the rachis, and its two exserted sta- mens with single-celled anthers. This species is widespread and shows variation in number of flowers per node, length of corolla, length of calyx lobes (4.5 to 9 mm), and pubescence of young stems. It may be distinguished from the similar R. citrina, which has lemon yellow corollas and less variation in calyx lobe length (4.5 to 9 mm vs. 7 to 9 mm). Razisea wilburii McDade, Syst. Bot. 7: 489. 1982. Figure 14. Shrubs to 1.5 m tall; internodes between leaf-bearing nodes 1.5-5.5 cm long, 2-3 mm thick, puberulous to pilose mostly along 2 lines, trichomes white to reddish brown in a transverse banding pattern. Leaves petiolate, petioles 1.5-1 1.5 cm long, pubescence that of the stem; laminae ovate to elliptic or obovate, 12-23.5 cm long, 5-10 cm broad, apically acuminate, basally acute to ob- tuse, margins entire to crenate, glabrous above, mod- erately puberulous on costa and veins beneath, cystoliths dense and prominent on both surfaces. Inflorescences in terminal racemes borne singly or to 3, to ca. 30 cm long; flowers numerous, borne singly or in fascicles of 2-3 at opposite nodes, pedicels 2.5-6 mm long, sparingly pu- berulous; bracts subulate, 5-6.5 mm long, ca. 1 mm broad basally, ciliolate, bracteoles similar, but smaller. Flowers with 5-merous calyx, calyx lobes linear-subu- late, 4-5 mm long, 0.5-0.75 mm broad basally, glabrous to sparingly puberulous; corolla bright red, funnelform, ascending, slightly curved, 2.7-3.3 cm long, ca. 1.5 mm broad basally, sparingly puberulous, upper lip erect, ovate, 4-6 mm long, 2-3 mm broad, the lower lip truncate, shallowly trilobate, the lobes ca. 0.5 mm long, stamens exserted 3-8 mm beyond tip of the upper lip, anthers 3- 3.5 mm long. Fruits clavate, 16-22 mm long, 3-4 mm broad, 3-3.5 mm thick; seeds 4, suborbicular, 2-3 mm in diameter, tuberculate. This species is known mainly from the type lo- cality, the Organization for Tropical Study Field Station, La Selva, on the Rio Puerto Viejo near the town of Puerto Viejo de Sarapiqui. Numerous collections have been made at this locality, and its is assumed that the species occurs elsewhere in the Caribbean lowlands of northern Costa Rica. Flowering collections have been made in January, June, September, and October. Razisea wilburii is recognized by its terminal racemes with pedicellate flowers either solitary or in fascicles of two to three at opposite nodes, its conspicuous bright red corollas which are 2.7 to 3.3 cm long, and its calyx lobe length of 4 to 5 mm. It is most easily confused with R. spicata, BURGER: FLORA COSTARICENSIS 71 which differs in its longer corolla (4 to 6.2 cm long) and mostly longer calyx lobes (4.5 to 9 mm vs. 4 to 5 mm long). Ruellia L. REFERENCES— R. W. Long, A Biosystematic Ap- proach to Generic Delimitation in Ruellia (Acan- thaceae). Taxon 22(5/6): 543-555. 1973. R. W. Long, Biosystematic Observation on Aphragmia inundata (Acanthaceae) from Mexico. Rhodora 78: 17-24. 1976. Perennial herbs or shrubs. Leaves mostly petiolate; laminae elliptic, ovate, ovate-lanceolate, oblong-spatu- late, apically acuminate or obtuse, basally attenuate or acute, the margins entire or undulate, cystoliths present and usually prominent on both surfaces. Flowers often large and showy, borne either singly or in various inflo- rescence forms which are either terminal or axillary or both, regular, sometimes curved, pedicellate or sessile; bracts 1 or absent; bracteoles 2 or absent. Flowers with a 5-merous calyx with mostly equal segments; corolla bluish, white, yellow, or red, funnelform, salverform, or sometimes saccate, the tubes usually narrow and the expanded portion often campanulatc. with 5 spreading, obtuse lobes; stamens 4, didynamous, the anthers 2-celled, basally muticous; stigma lobes unequal. Capsule oblong or clavate, the dissepiment remains attached to the cap- sule wall at maturity; seeds flattened and, when moist, rimmed by sticky, mucilaginous hairs, appearing gla- brous when dry. This is the second largest genus of the Acantha- ceae, with approximately 250 species distributed worldwide. Most species are found in the tropics, but a few extend into temperate regions. Key to Species of Ruellia la. Bracts, calyx, young leaves, and branches of inflorescence all with numerous glandular hairs; calyx segments unequal in length 2a Ib. Bracts, calyx, young leaves, and branches of inflorescence not all with numerous glandular hairs; calyx segments equal in length 5a 2a. Capsules clavate; calyx with 1 segment conspicuously longer than the others . . R. inundata 2b. Capsules elliptic or cylindric; calyx segments unequal, but none conspicuously longer than the others 3a 3a. Leaves and bracts gland-dotted R. golfodulcensis 3b. Leaves and bracts lacking gland-dots 4a 4a. Stamens slightly exserted; calyx segments 9-1 1 mm long R. paniculata 4b. Stamens included; calyx segments 1 6-20 mm long R. nudiflora 5a. Corolla 6 cm or longer; capsules 22 mm long or greater 6a 5b. Corolla 5.5 cm long or less; capsules 20 mm long or less lOa 6a. Inflorescences spicate; laminae and bracts gland-dotted; calyx lobes 3.5-4 mm long . . 7a 6b. Inflorescences paniculate or flowers solitary to clusters of 5 in the axils of uppermost leaves; laminae and bracts eglandular; calyx lobes 9-35 mm long 8a 7a. Young stems puberulous; laminae puberulous only on costa and veins beneath R. tubiflora var. tetrastichantha 7b. Young stems subtomentose; laminae pilose on both surfaces . . . R. tubiflora var. hirsuta 8a. Plants cultivated; inflorescences paniculate; bracts 4-10 mm long R. malacosperma 8b. Plants native; flowers axillary and solitary to clusters of 5; bracts 18-40 mm long .... 9a 9a. Flowers sessile, stamens included; bracts sericeous plus glandular hairs R. praeclara 9b. Flowers on pedicels 2-4 mm long, stamens exserted to just beyond the mouth; bracts glan- dular-puberulous R. palustris lOa. Leaves and bracts gland-dotted R. golfodulcensis lOb. Leaves and bracts lacking gland-dots 1 la 1 la. Plants cultivated for large showy flowers; capsules 25-30 mm long R. malacosperma 1 Ib. Plants native; capsules 6.5-20 mm long 12a 1 2a. Capsules elliptic; younger stems with glandular hairs R. nudiflora var. puberula 1 2b. Capsules clavate; younger stems lacking glandular hairs 1 3a 1 3a. Flowers in compact cymes (often appearing fascicled) at the ends of long unbranched peduncles; corollas puberulous-glandular R. biolleyi 72 FIELDIANA: BOTANY 13b. Flowers in spikes, spicate panicles, dichotomously branching panicles, or flowers axillary and solitary to clusters of 2-4; corollas eglandular 1 4a 14a. Flowers axillary and solitary to clusters of 2-4 and some of these occasionally on peduncles to no more than 5 cm long 1 5a 14b. Flowers in spikes, spicate panicles or dichotomously branching panicles 18a 1 5a. Corollas 4.5-7 cm. long; capsules 14-20 mm long R. jussieuoides 1 5b. Corollas 2-3.5 cm long; capsules 6.5-1 3 mm long 1 6a 16a. Capsules 6.5-8 mm long; leaves sessile or with petioles to 4 mm long R. geminiflora 16b. Capsules 1 1-13 mm long; leaves with petioles 5-15 mm long 17a 1 7a. Corolla puberulous; uppermost bracts 2-4 cm long; capsule minutely puberulous R. metallica 1 7b. Corolla glabrous; bracts less than 1 5 mm long; capsules glabrous R. barbillana 18a. Corollas 2.8-3.2 cm long; capsules 15-20 mm long; calyx in fruit 6-10 mm long R. stemonacanthoides 18b. Corollas 2.2-2.8 cm long; capsules 12-15 mm long; calyx in fruit 2-5 mm long 19a 1 9a. Capsule and calyx lobes puberulous plus glandular hairs; peduncle angled, but not winged R. cooperi 1 9b. Capsule and calyx lobes puberulous to glabrous, but eglandular; peduncle angled and slightly winged 20a 20a. Peduncles mostly less than 10 cm long, glabrous; younger stems glabrous; bracts elliptic- oblong R. tonduzii 20b. Peduncles mostly more than 10 cm long, sparingly puberulous; younger stems puberulous; bracts oblong-spatulate R. pittieri Ruellia barbillana Cufodontis, Arch. Bot. Sist. 10: 47. 1934. Herbs to 40 cm tall. Stem quadrangular, bearing nu- merous punctiform cystoliths, retrorse hairs at the angles. Leaves on petioles 1-1.5 cm long; laminae ovate-lanceo- late, to 10 cm long, 2.5 cm broad, apically acuminate, hasally attenuate, margins subrepand or undulate, gla- brous above, glabrous to appressed-hirsute on the costa and veins beneath, cystoliths visible on both surfaces. Inflorescences of clustered cymes borne in the axils of the upper leaves, bracts and bracteoles small. Flowers sessile, with 5-merous calyx, calyx 5 mm long, segments subulate; corolla pale purple, ca. 3 cm long, glabrous, 2 mm broad hasally, 6 mm broad at the throat, the limb spreading, the lobes ovate, rounded, ca. 8 mm long, 6 mm broad; stamens included. Fruits glabrous, seeds 6. This species is known only from the type col- lection which was made on the bank of Rio Bar- billa near Finca Waldeck in Limon Province. The above description was adapted from the original. Ruellia barbillana is recognized by its inflores- cences of clustered cymes borne in the axils of upper leaves, its sessile flowers with bracts less than 15 mm long, its purplish, glabrous corollas about 3 cm long, and its glabrous capsules. It is most easily confused with the more common and widespread R. metallica, which differs in its longer bracts (2 to 4 cm long) and its puberulous corollas and capsules. Ruellia biolleyi Lindau, in Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and in Pitt., Prim, fl. costaric. 2: 301. 1900. Ruellia longissima D. Gibson, Fieldiana, Bot. 34: 79. 1972. Figure 15. Herbs to 1 m tall, stems erect; internodes between leaf- bearing nodes 1-14 cm, 1-3.5 mm thick, quadrangular, yellow pilosulous. Leaves petiolate, petioles 8-22 mm long, yellow pilosulous; laminae elliptic, 3.5-14 cm long, 2-5 cm broad, apically acuminate, basally attenuate, margins undulate, glabrous above, puberulous (often yel- low) along costa and main veins beneath, cystoliths dense and prominent on both surfaces. Inflorescences in axil- lary cymes, peduncles 10-23 cm long, quadrangular, slightly winged, sparingly puberulous; flowers sessile to pedicels ca. 2 mm long, often in compact fascicles, lower bracts leaflike, elliptic to spatulate, to 1 5 mm long, upper bracts similar, but smaller. Flowers with 5-merous calyx, calyx segments long-subulate, 4-5 mm long, glandular- puberulous; corollas violet or purplish, 20-24 mm long, unexpanded portion of tube ca. 1 0 mm long, ca. 1 mm broad basally, 5.5-7 mm broad at mouth, puberulous plus some glandular hairs outside; stamens included, an- thers basally rounded. Fruits clavate, 13-17 mm long, 3.4-4.0 mm broad, 1 .4-1 .5 mm thick, glandular-puberu- lous, more so at the stipe; seeds 4-8, orbicular, 1.8-2 mm in diameter. Found in lowland rain forest areas of Panama and Costa Rica. In Costa Rica it has been collected in mature or recently cut lowland rain forests from near sea level to 300 m elevation in Heredia and BURGER: FLORA COSTARICENSIS 73 Limon provinces. Flowering collections have been made in Costa Rica in January, February, March, July, and August. Ruellia biolleyi is recognized by its long, pedun- cled ( 1 0 to 23 cm long) inflorescences bearing com- pact, near-sessile flowers in compact cymes. The flowers often appear fascicled at the ends of these long peduncles. Further, it is recognized by its vi- olet or purplish, puberulous-glandular corollas (20 to 24 mm long) with included stamens, glandular- puberulous calyx segments (4 to 5 mm long), and 4- to 8 -seeded glandular-puberulous capsules, which are 1 3 to 17 mm long. It resembles R. coop- eri, which differs in the more extensive branching of its inflorescence, thus lacking the often-fascicled flowers, a white corolla (which may turn purplish with age) that is slightly longer (2.5 to 2.8 cm long), and a puberulous calyx and fruit (vs. glandular- puberulous). I have placed R. longissima D. Gibson in syn- onymy here because all of its characteristics lie within the range of those described for the more widespread and variable R. biolleyi. Ruellia cooperi Leonard, Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1251. 1938. Figure 15. Herbs to ca. 1.5m tall, stems erect; inter nodes between leaf-bearing nodes 3-12 cm long, 2-2.8 mm thick, te- trangular, subglabrous. Leaves on puberulous petioles 1- 3.5 cm long; laminae elliptic, 5-16 cm long, 1.6-6 cm broad, apically acuminate, basally attenuate, margins undulate, glabrous above, puberulous on veins beneath, cystoliths dense and prominent on both surfaces. Inflo- rescences of peduncled, axillary, dichotomously branch- ing panicles, the peduncles 8-30 cm, but mostly more than 15 cm long, sharply quadrangular, glabrous, the branches frequently curving; flowers borne singly or in pairs at the axils of the branches or at their terminus; bracts elliptic to spatulate, lower bracts leaf] ike. 20-25 mm long, 6-8 mm broad, upper bracts much smaller, all glabrous. Flowers with 5-merous calyx, lobes 2-5 mm long, puberulous plus occasional glandular hairs; corolla white or pale lavender, 2.3-2.5 cm long, sparingly pu- berulous, unexpanded portion of tube ca. 1 cm long, 7-8 mm broad at the mouth, 1.5 mm broad basally; stamens included. Fruits clavate, 13-14 mm long, 3-3.6 mm broad, ca. 2 mm thick, puberulous plus occasional glandular hairs. Found at lower altitudes of Panama and Costa Rica. Only one collection, Burger, Visconti and Gentry 10643 (GRI) from near Quepos, Puntarenas Province, has been seen. This was collected in Feb- ruary on shaded slopes above a stream in rain forest. Ruellia cooperi is recognized by its inflores- cences of dichotomously branching panicles on long peduncles (8 to 30 cm long), capsules which are puberulous with occasional glandular hairs, and white to pale lavender corollas which are spar- ingly puberulous. It most closely resembles R. biolleyi, which differs in its inflorescences, with cymose flowers that are short-pedicelled and often fascicled, and its violet or purplish corollas which are puberulous with some glandular hairs. Ruellia geminiflora Kunth in H.B.K., Nov. gen. sp. 2:240. 1817. Figure 15. Ascending herbs to ca. 50 cm tall; internodes between leaf-bearing nodes 1.5-5.5 cm long, 1-1.5 mm thick, quadrangular, appressed pilose mostly at the angles. Leaves sessile to petioles 4 mm long; laminae ovate to ovate-elliptic, 4-7 cm long, 0.9-2.2 cm broad, apically acute, basally attenuate, margins entire to crenulate, ma- ture leaves pilose beneath, young leaves and those sur- rounding the flowers densely puberulous to tomentose, cystoliths densely distributed over both surfaces where visible. Pedicels axillary, borne singly or in pairs, ca. 1 mm long. Flowers with 5-merous calyx, lobes equal, subulate, 5-7 mm long, 0.8-1 mm broad basally, puberu- lous, ciliate; corolla mauve, funnelform, 2. 1-3.5 cm long, puberulous, unexpanded portion of tube 8-13 mm long, ca. 1 mm broad, 6-9 mm broad at the mouth, lobes equal, suborbicular, 6-10 mm broad; stamens included, the filaments glabrous; ovary and style puberulous. Fruit clavate, 6.5-8 mm long, 3.3-3.7 mm broad, 2-2.8 mm thick, puberulous; seeds 4, orbicular, flattened, 2-3 mm in diameter. This species is found in open grassland areas of Central America, the West Indies, and northern South America. In Costa Rica it has been collected in the Pacific watershed areas of Alajuela and Guanacaste provinces often in secondary succes- sion following burning. Collections in Costa Rica have been made at altitudes of 150 to 500 m. Flowering collections have been made in January, February, March, May, and June. Ruellia geminiflora is recognized by its rela- tively small, subsessile leaves (4 to 7 cm long), its axillary, short-pedicelled flowers borne singly or in pairs with mauve corollas (2.1 to 3.5 cm long), and its short, stout, clavate, puberulous capsules (6.5 to 8 mm long). It may be confused with R. metallica, which has larger, petiolate, often lus- trous leaves (to 15 cm long), inflorescences with sessile, axillary flowers in fascicles of three to four, linear to linear-elliptic bracts (2 to 4 cm long) which intergrade with leaves basally, and its clavate cap- sules (11 to 13 mm long). 74 FIELDIANA: BOTANY Ruellia golfodulcensis Durkee, sp. nov. Figure 15. Herbae suffruticosae, usque ad 1 m altae, caulibus erectis, glabris. Folia petiolata, laminis ellipticis, 7-15 cm longis, 1.4-4 cm latis, apice acuminatis, basi longe attenuatis, subglabris et glanduloso-punctatis, cystolithis supra prominentibus. Inflorescentiae terminales, race- mosae, angustae, vel in specimenibus junioribus flores axillares solitarii. Flores corollis albis infundibulifor- mibus, ca. 4 cm longis, staminibus inclusis. Fructus non visi. Suflrutescent plant to 1 m tall, stems erect; internodes between leaf-bearing nodes 1 .7-8 cm long, 1-2 mm thick, tetrangular, glabrous, uppermost internodes gland-dot- ted. Leaves on glabrous petioles 5-1 5 mm long; laminae elliptic, 7-15 cm long, 1.4—4 cm broad, apically acu- minate, hasally long-attenuate, margins entire, sparingly ciliolate, subglabrous and gland-dotted on both surfaces, gland-dots more prominent in younger leaves, cystoliths visible on both surfaces but more prominent above. In- florescences of narrow, terminal racemes in older spec- imens, with axillary flowers borne singly in younger spec- imens; bracts lanceolate, 1 5-26 mm long, 3-6 mm broad, gland-dotted, margins ciliolate. Flowers on glandular- puberulous pedicels 3-5 mm long; calyx 5-merous, lobes linear, subequal, 13-20 mm long, 0.75-1.0 mm broad, glandular-puberulous; corolla white, funnelform, ca. 4 cm long, sparingly glandular-puberulous, tube 2.8 cm long, unexpanded portion of tube 1.5 cm long, lobes rounded, 17 mm long, 6 mm broad; stamens included. Fruits not observed. TYPE— Costa Rica; Puntarenas Prov., Golfo Dulce and Rio Grande de Ten-aba, elevation 1 600 m, undershrub in forest, December 1947, Skutch 5304 (holotype, us). Additional Collections: Pun- tarenas Province, Golfo Dulce area, vicinity of Esquinas Experiment Station, elevation sea level, common, April 26, 1949, Allen 5266 (us). Flow- ering collections were made in April and Decem- ber. Ruellia golfodulcensis is recognized by its short- pedicellate flowers with a conspicuously glandu- lar-puberulous calyx with subequal, linear lobes ( 1 3 to 20 mm long); white, glandular-puberulous corollas (ca. 4 cm long) borne singly at leaf or bract axils; leaves and bracts which are gland-dotted. It most closely resembles the much wider-ranging R. metallica of similar habitats, which differs in its leaves which lack glandular dots, shorter calyx lobes (5 to 7 mm long) which are subulate, and shorter corolla (2.3 to 2.7 cm long) which is puberulous only. Ruellia inundata Kunth in H.B.K., Nov. gen. sp. 2:239. 1817. Figure 15. Herbs sometimes suffrutescent, to 1.5 m tall; inter- nodes between leaf-bearing nodes 2-7.5 cm long, 2.1- 2.8 mm thick, subquadrangular, pilose. Leaves on pilose petioles 1-8.5 cm long; laminae ovate to ovate-lanceo- late, 4-15 cm long, 1.2-8 cm broad, apically acuminate, basally attenuate, margins denticulate, pilose above and beneath, young leaves glandular-puberulous, cystoliths common, but more visible above. Inflorescences of as- cending axillary dichasia which often form dense pani- cles, pedicels 8-30 mm long, glandular-pubescent; floral bracts paired, lanceolate, often slightly unequal, 3-5 mm long, 0.8-1.4 mm broad, puberulous-glandular, lower bracts similar, but larger. Flowers with 5-merous calyx, segments linear, 10-14 mm long, 1 segment conspicu- ously longer (ca. 2 mm) than the others, glandular-pu- berulous; corolla pink or mauve, narrowly funnelform, the tube slightly curved, 2.2-2.5 cm long, 1-1.2 mm broad basally, 3-4 mm broad at the mouth, glabrous to sparingly puberulous, the 5 lobes equal, rounded 4-5.5 mm long, 3-3.5 mm broad; stamens included. Fruits clavate, 8-10 mm long, 2.4-3.1 mm broad, 1.2-2 mm thick, glabrous; seeds 2-4, flattened, orbicular, 2.0-2.5 mm in diameter. This is a common species growing in open or partially shaded areas that have been recently dis- turbed, such as roadsides and trails. Its range ex- tends from Mexico to much of South America. In Costa Rica it has been collected only on the Pacific watershed at altitudes of 50 to 900 m in Alajuela, Guanacaste, Puntarenas, and San Jose provinces. Flowering collections have been made from No- vember through March. Ruellia inundata is recognized by its ascending axillary dichasia that often form dense panicles; pink or mauve, narrowly funnelform corollas (2.2 to 2.5 cm long, excluding lobes); small, paired, lanceolate bracts (3 to 5 mm long) which are often slightly unequal in length; and a glabrous, clavate capsule (8 to 10 mm long). Bracts, calyx, pedicels, and young leaves all have numerous glandular hairs which give fresh specimens a fetid odor. It is most easily confused with R. paniculata, which can be distinguished by its cylindric capsules (10 to 13 mm long) and slightly exserted stamens. Long (1976) presents evidence suggesting that R. inundata belongs in the genus Aphragmia on the basis of hybridization experiments as well as morphological comparisons with R. tuberosa, the type for the genus, and R. caroliniensis. However, Long does not mention comparisons between R. inundata and R. paniculata with which it is often confused because of morphological similarities. Ruellia paniculata is found in similar habitats and extends over the same wide range from Mexico to Brazil. Ruellia paniculata differs from R. inundata in ways that suggest closer affinity to R. tuberosa, with its bluish corolla and cylindrical capsule (vs. red or pink corolla and clavate capsule), suggesting BURGER: FLORA COSTARICENSIS 75 that R. paniculata does belong in the genus. Until the relationships among R. inundata, R. panicu- lata, and other species clearly regarded as within the genus Ruellia are more thoroughly investigat- ed, I have decided to recognize R. inundata as Ruellia. Ruellia jussieuoides Schlecht., Linnaea 6: 370. 1830. Ruellia achimeniflora (Oerst.) Hemsl., Biol. Centralbl. 2: 503. 1882. Arrhostoxylum achimeniflorum Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 1854: 131. 1881. Figure 15. Herbs or suffnitescent plants to 2.5 m tall; internodes between leaf-bearing nodes 3-14 cm long, 2-4 mm thick, tetrangular, puberulous mostly at the angles. Leaves pet- iolate, petioles 1-3 cm long, puberulous; laminae, ellip- tic, elliptic-ovate or lanceolate, 5-14.5 cm long, 2-5.5 cm broad, apically long-acuminate, basally attenuate, margins entire to crenulate, sparingly pilose to pilose above, pilose beneath, veins and veinlets conspicuous above in dried specimens, cystoliths dense and promi- nent on both surfaces. Inflorescences axillary, flowers sessile and solitary to clusters of 2-3 at the ends of puber- ulous peduncles 0.25-5 cm long; bracts lanceolate-ellip- tic, 2-4 cm long, 5-10 mm broad, puberulous, bracteoles similar though smaller. Flowers with 5-merous calyx, calyx lobes subequal, long-subulate, 1 lobe often slightly longer than the rest, 6-20 mm long, 0.8-1 mm broad basally, ciliolate, puberulous; corolla lavender or purple, funnelform, frequently curved, puberulous outside, the tube 3.3-5.5 cm long, ca. 1 mm broad basally, expanding to 5-10 mm broad at the mouth, the lobes subequal, suborbicular, 10-18 mm in diameter; stamens included, anthers ca. 3 mm long. Fruits clavate, 1 4-20 mm long, 3.5-5 mm broad, ca. 2.6 mm thick, puberulous plus minute glandular hairs; seeds 10-14, elliptic, flattened, ca. 2.5 mm long, 2 mm broad, puberulous, when wet, margins covered with dense, mucilaginous hairs. Ranging from southern Mexico to Costa Rica, this species is found from 300 to 1 500 m elevation in Costa Rica where it has been collected mainly in openings and along trails in rain forests around San Jose and Cartago provinces, although several collections have been seen from the vicinity of San Isidro of Puntarenas Province. Flowering collec- tions have been made from November through April. Ruellia jussieuoides is recognized by its axillary inflorescences with sessile flowers solitary to clus- ters of two to three at the ends of peduncles 0.25 to 5 cm long, corollas lavender or purple with tubes 3.3 to 5.5 cm long, stamens included, fruits 14 to 20 mm long, puberulous. It resembles R. tubiflora var. tetrastichantha, which differs by its flowers borne in terminal or lateral spikes resulting in a conical rachis, bracts and calyx lobes that are gland-dotted, and white corollas which are 7 to 8 cm long. Ruellia malacosperma also resembles R. jussieuoides, but differs with its dichotomously branched cymose inflorescences and its pedicellate flowers. I have placed Ruellia achimeniflora in synonymy with R. jussieuoides because I can find no characters that differ consistently between these two taxa. Ruellia jussieuoides is much wider rang- ing, and it appears that R. achimeniflora is a pop- ulation which differs mainly in wider leaves, but this variation is inconsistent. Ruellia malacosperma Greenm., Proc. Amer. Acad. Arts 34: 572. 1899. Herbs to 60 cm tall; internodes between leaf-bearing nodes 1.5-8.5 cm long, 1.8-2.2 mm thick, quadrangular, sparingly pilose. Leaves on glabrous petioles 0.5-2 cm long; laminae narrowly elliptic 5-15 cm long, 1-3.4 cm broad, apically acute or obtusish, margins undulate to entire, glabrous above and beneath, cystoliths visible on both surfaces. Inflorescences of ascending axillary di- chasia that branch once or twice, peduncles quadrangular to 9 cm long, glabrous pedicels 5-1 5 mm long, sparingly glandular-puberulous; bracts linear 4-10 mm long, 1- 1.5 mm broad, glabrous. Flowers with 5-merous calyx with lobes long-subulate, 10-12 mm long, ca. 1.25 mm broad basally, glandular-puberulous; corolla mauve or purple, 4.5-5.5 cm long, cylindrical portion ca. '/a the length, 2.3-3 mm broad, the throat funnelform, ca. 3 cm broad at the mouth, the lobes rounded, ca. 10 mm long, 1.5-1.8 mm broad; the stamens included. Fruits nar- rowly clavate, 2.5-3 cm long, 6 mm broad, glabrous; seeds 8-20, discoid, ca. 3 mm in diameter, silky with white, appressed hairs when dry. This species is native to Mexico, but is culti- vated throughout much of tropical America, in- cluding Costa Rica, and especially the central val- ley area. The only Costa Rican collection I have examined was from Parque Bolivar of La Hon- dura, San Jose Province. It was collected in flower in June. Ruellia melacosperma is recognized by its sim- ple panicles with one or two dichotomous branch- es on peduncles to 9 cm long, its large, showy, mauve or purple corollas (4.5 to 5.5 cm long), and its large, narrowly clavate, glabrous capsules (2.5 to 3 cm long). It has been confused with R. ste- monacanthoides, which has smaller corollas (3 to 4 cm long) and shorter capsules (15 to 20 mm long). It also may be confused with R. paniculata, which has shorter corollas (3 to 4 cm long) and shorter, cylindric, glabrous to sparingly puberu- lous capsules. 76 FIELDIANA: BOTANY Ruellia metallica Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1253. 1938. Figure 16. Suffrutescent herb to 1 m tall; internodes between leaf- bearing nodes 1-9.5 cm long, 1-3 mm thick, subqua- drangular, sparingly puberulous mostly at the angles. Leaves on glabrous petioles 5-13 mm long, glabrous. Laminae elliptic to elliptic-ovate, varying from broadly to narrowly so, 4-17 cm long, 1-8 cm broad, apically acuminate, basally attenuate, margins undulate, glabrous above, sericeous on the costa and veins beneath, cys- toliths prominent above and beneath, upper surface often faintly lustrous or purplish, the lower surface often pur- plish or violet purple beneath, especially in younger leaves. Inflorescences axillary or terminal with sessile flowers in fascicles of 3-4; uppermost bracts linear to linear-ellip- tic, 2—4 cm long, lower bracts intergrading with leaves. Flowers with calyx segments equal, subulate, 5-7 mm long, glabrous except scant puberulence at the margins; corollas lavender, pink, or white, funnelform, 2.3-2.7 cm long, puberulous, cylindrical portion ca. 1.5 mm broad, 10 mm broad at the mouth, lobes suborbicular, 5-7 mm in diameter; stamens included. Fruits clavate, 1 1-13 mm long, 2.5-3.2 mm broad, 2.2-2.6 mm thick, minutely puberulous; seeds 2, suborbicular, 2.5-2.8 mm in diameter. This species occurs in and adjacent to lowland rain forest areas ranging from Guatemala to Pan- ama. In Costa Rica it has been collected in moist areas of both watersheds, excluding Guanacaste and San Jose provinces. Flowering collections have been made in January, February, March, May, June, September, and December. Ruellia metallica is recognized by its terminal spikelike inflorescences with axillary fascicles of three to four sessile flowers subtended by bracts which are linear to linear-elliptic (2 to 4 cm long) near the apex, but intergrading with leaves basally, its lavender, pink, or white corollas (2.3 to 2.7 cm long), its clavate, minutely puberulous capsules (11 to 13 mm long), and its leaves which are often purplish or violet purple beneath, lustrous or pur- plish above. It is most easily confused with R. geminiflora, which differs with its smaller (4 to 7 cm long) sessile to short petiolate leaves, its in- conspicuous bracts, its inflorescences with pedi- cellate, axillary flowers in fascicles of one to two, and its clavate capsules (6.5 to 8 mm long). Ruellia nudiflora (Engelm. & Gray) Urb., Symb. antill. 7: 382. 1912. Dipteracanthus nudiflorus Engelm. & Gray, Boston J. Nat. Hist. 5: 229. 1845. Ruellia nudiflora (Engelm. & Gray) Urb. var. puberula Leonard, J. Wash. Acad. Sci. 17: 517.1 927. Ruellia puberula (Leonard) Tharp & Barkley, Amer. Midi. Naturalist 42: 17. 1949. Figure 16. Herbs to 30 cm tall; internodes between leaf-bearing nodes 1-8 cm long, 1.4-2.7 mm thick, quadrangular, glandular-puberulous plus occasional larger pilose hairs. Leaves on glandular-puberulous petioles 0.4-3.2 cm long; laminae elliptic to obovate, 3.5-11.5 cm long, 1.3-3.7 cm broad, apically acute, obtuse or rounded, basally at- tenuate, margins undulate, puberulous above mixed with occasional pilose hairs, similar beneath, but densely pu- berulous, cystoliths dense, more visible above, obscured by hairs beneath. Inflorescences of axillary dichasia, pe- duncles 2-8 cm long, glandular-puberulous, pedicels 0.5- 2 cm long, glandular-puberulous, bracts oblong, 7-9 mm long, glandular-puberulous. Flowers with calyx lobes lin- ear-subulate, equal, 16-20 mm. long, glandular-puber- ulous; corolla light blue or lavender, funnelform, 3.5- 4.5 cm long, tube ca. 2 mm broad basally, expanding to 8-15 mm broad at the mouth, puberulous, the lobes suborbiculate, 9-1 1 mm broad; stamens included. Fruits elliptic, 16-18 mm long, 2.5-4 mm broad, puberulous; seeds ca. 12, ovate, basally asymmetrical, ca. 2.5 mm long, 2.0 mm broad. Found in dry savannas and pastures, this species survives burning and is then conspicuous for its showy flowers appearing close to the ground after growth is resumed. This variety ranges from Oa- xaca to central Panama. In Costa Rica it is found at elevations of 25 to 100 m in the lowlands of Guanacaste. Flowering collections have been made in January, February, May, and June. Ruellia nudiflora is recognized by its puberu- lous, elliptic to obovate laminae, its axillary di- chasia on glandular-puberulous peduncles to 8 cm long bearing pedicellate flowers with light blue or lavender corollas 3.5 to 4.5 cm long with stamens excluded, and its puberulous, elliptic capsules 1 6 to 1 8 mm long. It may be confused with R. pan- iculata, which differs in its elliptic to ovate leaves, its corollas with stamens slightly exserted, and its glabrous, cylindric capsule (10 to 13 mm long). Ruellia palustris Durkee, sp. nov. Figure 16. Herbae suffruticosae, usque ad 1 m altae, caulibus erectis, junioribus pilosis-adpressis. Folia petiolata, lam- inis ellipticis vel elliptico-obovatis, 5.5-15.5 cm longis, 2.2-8.5 cm latis, apice acuminatis, basi acutis vel parum attenuatis, maximam partem glabris, cystolithis abun- dantibus. Inflorescentiae fasciculatae, axillares et ter- minates, 1-5 floribus, bracteis 2 in quoque flore. Flores corollis albis infundibuliformibus, tubis 6-7 cm longis et lobis 1 .6-2 cm longis, staminibus paulo exsertis. Fruc- tus clavati, ca. 23 mm longi. Suffrutescent herb to 1 m tall; stems erect; internodes between leaf-bearing nodes 3-1 5 cm long, 2-4 mm thick, appressed-pilose. Leaves petiolate, petioles 6-18 cm long, BURGER: FLORA COSTARICENSIS 77 appressed-pilose; laminae elliptic to elliptic-obovate, 5.5- 15.5 cm long, 2.2-8.5 cm broad, apically abruptly acu- minate, basally acute to slightly attenuate, margins entire to slightly undulate, glabrous on both surfaces except for appressed hairs on costa and main veins, cystoliths abun- dant and visible on both surfaces. Inflorescences in ax- illary and terminal clusters of 1-5 flowers, flowers borne on short pedicels 2-4 mm long, strigose; bracts 2 per flower, shape variable from broadly elliptic to obovate or oblanceolate, 3-4 cm long, 0.6-2.3 cm broad, apically acute, glandular-puberulous. Flowers with a 5-merous calyx, calyx lobes equal, long-elliptic to lanceolate, 25- 30 mm long, 4-5 mm broad, glandular-puberulous; co- rolla white, funnelform, the tube 6-7 cm long, cylindrical portion 4.5-5 cm long, 3.5-4 mm broad, funnelform portion 1.5-2 cm long, 10-16 mm broad at the mouth, glandular-puberulous, the lobes equal, suborbicular, 1 .6- 2 cm long, 2-2.5 cm broad; stamens exserted to just beyond the mouth, anthers 4-5 mm long, ca. 0.8 mm broad, filaments attached at ca. the beginning of the funnelform portion of the corolla tube. Fruits clavate, ca. 23 mm long, 8 mm broad, 8 mm thick, puberulous; seeds 8. TYPE— Costa Rica: Heredia Prov. Collected at swamp's edge in mature rain forest at Finca La Selva on Rio Puerto Viejo, elev. ca. 100 m, 27 July 1980, Hammel 9341 (Holotype, DUKE). Ad- ditional Collections— All from the same vicinity and similar habitat, Grayum 1467, 2428 (DUKE); Hammel 10480 (DUKE), McDade 364 (DUKE); Da- mon Smith 614 (DUKE). Flowering collections have been made in July, August, and November. Ruellia palustris is recognized by its large white corollas (6 to 7 cm long) with two-thirds of the tube cylindrical, the two large, mostly elliptic bracts per flower and with axillary and terminal flowers in clusters of one to five. It most closely resembles R. praeclara of similar habitats in western Panama and southeastern Costa Rica, which differs in its shrubby habit, longer corollas (to 9 cm), and linear bracts and calyx lobes. Ruellia paniculata L., Sp. pi. 2: 635. 1753. Figure 16. Herbs to suffrutescent plants; internodes between leaf- bearing nodes, 2-6.5 cm long, 1.8-3.5 mm thick, qua- drangular, glandular-puberulous. Leaves on petioles 0.5- 4 cm long, glandular-puberulous; laminae ovate to el- liptic, 3-10(17) cm long, 1 .5-4.5 (6.5) cm broad, apically acute to obtuse, basally abruptly attenuate, margins un- dulate, glandular-puberulous plus occasional pilose hairs on both surfaces, but more dense and prominent above, less so beneath. Inflorescences in axillary, branching di- chasia forming dense clusters in mature plants, the branches glandular-puberulous; floral bracts narrowly el- liptic to obovate, 3-5 mm long, 0.5-0.7 mm broad, glan- dular-puberulous, lower bracts larger intcrgrading with the leaves. Flowers with calyx segments elongate-subu- late, unequal, 9-1 1 mm long, basally 0.7-0.8 mm broad, glandular-puberulous; corolla light blue, funnelform, puberulous, 2.5-4 cm long, unexpanded portion of tube ca. 1.3 cm long, 1-1.2 mm broad, lobes suborbiculate, 6-7 mm broad; stamens slightly exserted. Fruits cylin- dric, 12.5-15 mm long, 2-2.5 mm in diameter, glabrous to minutely puberulous apically; seeds 8, flat, ovate, ca. 2 mm long, 1.5 mm broad. This species is wide ranging from Mexico to Brazil where it grows in open disturbed habitats with a dry season at elevations of sea level to 200 m. It is not frequently collected, either because it is uncommon or, as I suspect, it is mistaken for the much more common R. inundata of similar habitats. Only two Costa Rican collections have been observed from the Bagaces and Canas areas of Guanacaste, both in flower— one in January, the other in April. Ruellia paniculata is recognized by its dense ax- illary dichasia, with most parts bearing a dense glandular-puberulence, blue corollas (2.5 to 4 cm long) with slightly exserted stamens, and cylindric capsules (12.5-15 mm long). It is most often con- fused with R. inundata of similar habitat, which differs in its pink or mauve corollas, included sta- mens, and clavate capsules (8 to 10 mm long). Ruellia pittieri Lindau, Bull. Herb. Boissier 5:655. 1 897. Ruellia longissima var. glabra D. Gibson, Fieldiana, Bot. 34: 80. 1972. Figure 16. Herbs to suffrutescent plants to 2 m tall; internodes between leaf-bearing nodes, 2.8-1 1 .5 cm long, 3-3.8 mm thick, quadrangular, puberulous. Leaves on petioles 2- 6.5 cm long, puberulous; laminae ovate to ovate-elliptic, 7-12 (19) cm long, 2.5-7 (8.5) cm broad, apically acu- minate, basally acute to obtuse, margins undulate, gla- brous above, puberulous on costa and veins beneath, cystoliths dense and prominent on both surfaces. Inflo- rescences in axillary, lax, spicate panicles that are often dichotomously branching; peduncles 1 2-2 1 cm long, pe- duncles and branches sparingly puberulous and slightly winged; nonfloral bracts oblong-spat ulate, 15-35 mm long, glabrous; floral bracts elliptic, 4-8 mm long, 1-2 mm broad, glabrous. Flowers sessile to subsessile, with calyx segments subulate, 2-3 mm long at anthesis, 3.5- 5 mm long at fructescence, ca. 0.5 mm broad basally, sparingly puberulous; corolla light blue or pale violet, funnelform, 2.3-2.8 cm long, puberulous, unexpanded portion of tube 8-10 mm long, 1-1.5 mm broad, 5-10 mm broad at the mouth, lobes equal, obtuse, 5.5-7 mm long, 5-7.5 mm broad; stamens included. Fruits clavate, 12-15 mm long, 3-4 mm broad, 1 .9-2.2 mm thick, spar- ingly puberulous to glabrous; seeds 4, 6, or 8, suborbicu- lar, 2.2-2.8 mm in diameter. 78 FIELDIANA: BOTANY This species is commonly found along streams and other openings in wet forests of Costa Rica and Panama. In Costa Rica it is found in southern Puntarenas Province at elevations of 30 to 270 m. Most collections have been made in the Golfo Dulce area and on the Burica Peninsula. Flowering collections have been made in December, March, and April. Ruellia pittieri is recognized by its lax, dichot- omous, spicate panicles, with slightly winged, te- trangular peduncles (and branches) which usually exceed 10 cm in length; its oblong-spatulate bracts, its funnelform corollas (2.3 to 2.8 cm long), and its clavate capsules (1 2 to 15 mm long). It is easily confused with R. stemonacanthoides, which differs in its longer corollas (2.8 to 3.2 cm long), its longer calyx lobes at anthesis (4 to 7 mm long vs. 2 to 3 mm), and its longer capsules (15 to 20 mm long). Ruellia pittieri is also easily confused with R. ton- duzii, which differs in its shorter peduncles, (most- ly less than 1 0 cm long), its oblong-elliptic bracts, and while the size ranges overlap, its generally shorter leaves (6 to 13.5 cm long vs. 7 to 19 cm). Ruellia praeclara Standley, Publ. Field Colum- bian Mus., Hot. Ser. 4: 263. 1929. Figure 16. Suffrutescent plants to 2 m tall; internodes between leaf-bearing nodes 2.4—4.5 cm long, 2.3-3.0 mm thick, quadrangular, appressed pilose. Leaves on petioles 1.5- 3 cm long, glabrous to appressed pilose; laminae oblan- ceolate, 5.5-30 cm long, 1.6-9 cm broad, apically acu- minate, basally acute to attenuate, margins entire to un- dulate, glabrous to sparingly strigose at the costa and veins, the cystoliths prominent on both surfaces. Flowers axillary, sessile, solitary to clusters of 3; bracts linear to linear-oblanceolate, 18-30 mm long, 2-3 mm broad, sericeous plus glandular hairs. Calyx segments linear to linear-lanceolate, 9-35 mm long, 1.5-2 mm broad, se- riceous with numerous glandular hairs; corolla white, 8- 10 cm long, funnelform, puberulous-glandular, unex- panded portion 4-5.5 cm long, 3 mm broad, expanding to 9-14 mm broad at the mouth, lobes orbicular 15-20 mm in diameter; stamens included, but extending to just below the mouth. Fruits not seen. Found in the shade of lowland rain forests of western Panama (Caribbean side) and one site in the Osa Peninsula of Costa Rica. Three flowering collections have been made, two in January by Burger & Liesner 7202 (F, NY), 7329 (F), and one in March by Burger & Gentry 8915 (us), all from near Rincon de Osa of Puntarenas Province. The Costa Rican population is at the lower range of measurements for leaves, bracts, and calyx lobes. Also, bracts and calyx lobes are linear-oblanceo- late and linear-lanceolate, respectively, in the Cos- ta Rican population as opposed to linear for both in the Panamanian population. Ruellia praeclara is recognized by its sessile, axillary, solitary flowers with white corollas (8 to 10 cm long), its large lanceolate leaves (to 30 cm long), and its linear to linear-lanceolate calyx lobes (9 to 35 mm long) with both sericeous and nu- merous smaller glandular hairs. It may be con- fused with R. tubifora var. tetrastichantha or var. hirsuta, both of which differ in their terminal and subterminal spicate inflorescences, the numerous glandular spots on the lower surfaces of leaves and bracts, and much shorter calyx lobes (3.5 to 4 mm long vs. 9 to 35 mm long). Ruellia stemonacanthoides (Oerst.) Hemsl., Biol. Centralbl. 2: 507. 1882. Arrhostoxylum stemo- nacanthoides Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854: 130. 1855. Figure 17. Herbs to suffrutescent plants 1 m tall; internodes be- tween leaf-bearing nodes, 3-10.5 cm long, 1.3-3.2 mm thick, quadrangular, puberulous. Leaves on petioles to 3.5 cm long, puberulous to glabrous; laminae ovate-el- liptic, 4-12 cm long, 2-5.7 cm broad, apically acuminate, basally acute and often slightly attenuate, margins un- dulate, glabrous above, frequently drying to a purplish color, puberulous beneath on costa and veins, cystoliths dense and visible on both surfaces although occasionally obscure. Inflorescences in axillary panicles, often di- chotomously branched, peduncles to 1 1 cm long, qua- drangular, often slightly winged, glabrous; bracts elliptic, 10-27 mm long, 2-6 mm broad, glabrous. Flowers ses- sile to pedicles 3 mm long, puberulous, with calyx seg- ments linear-subulate, 4-7 mm long at anthesis, 6-10.5 mm long at fructescence, sparingly puberulous; corolla pale violet, funnelform, 2.8-3.2 cm long, puberulous, unexpanded portion of tube 9-11 mm long, 1 . 1-1 .5 mm broad, 10-12 mm broad at the mouth, lobes equal, ob- tuse, 6-9 mm long, 5-8 mm broad; stamens included. Fruits clavate, 15-20 mm long, 3-5 mm broad, 2.25- 4.5 mm thick, sparingly puberulous; seeds 8-10, subor- bicular, 2.5-3.2 mm in diameter. This species is commonly found in or near re- cently cut forests or in new pastures in premontane forest areas of Costa Rica to Guatemala and west- ern Mexico. In Costa Rica it has been collected mainly from the Pacific watershed areas of San Jose, Alajuela, and Guanacaste provinces, with one collection from the Guapiles area of Limon Province, all at elevations of 300 to 1000 m. Flow- ering collections have been made in September through November, January, February, April, and July. BURGER: FLORA COSTARICENSIS 79 Ruellia stemonacanthoides is recognized by its paniculate inflorescences which are often dichot- omously branched on peduncles to 1 1 cm long, its pale violet corollas (2.8 to 3.2 cm long), its calyx lobes (4 to 7 mm long at anthesis), and its sparingly puberulous, clavate capsules ( 1 5 to 20 mm long). It is easily confused with R. pittieri and R. ton- duzii. See the description of R. pittieri to distin- guish these differences. Ruellia tonduzii Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 1 88. 1 898, and in Pitt, Prim, fl. costaric. 2: 203. 1900. Figure 17. Erect herbs to ca. 50 cm tall; internodes between leaf- bearing nodes 2.5-7 cm long, 1 .5-2.5 mm thick, quadran- gular, slightly winged, glabrous. Leaves on petioles 2-3 cm long, puberulous; laminae elliptic to ovate, 3.5-12.5 cm long, 1.1-5.2 (7) cm broad, apically acuminate, ba- sally acute and frequently attenuate or obtuse, margins undulate, glabrous above, puberulous on costa and veins beneath, cystoliths dense and prominent on both sur- faces. Inflorescences in axillary, dichotomously branch- ing panicles on glabrous, quadrangular, slightly winged peduncles 5-9.8 cm long; bracts elliptic-oblong, 8-13 mm long, 1.5-3 mm broad, glabrous. Flowers borne sin- gly on glabrous pedicels to 3 mm long; calyx segments subulate, 2.7-4 mm long, 0.5 mm broad basally, gla- brous to subglabrous; corolla lavender, funnelform, 2.2- 2.8 cm long, puberulous, unexpanded portion of tube ca. 9 mm long, 1.5 mm broad, 7 mm broad at the mouth, lobes equal, obtuse, 5-7 mm long, 3.5-5.5 mm broad, stamens included. Fruits clavate, 12-15 mm long, 3-4 mm broad, 2.5 mm thick, sparingly puberulous; seeds 8, suborbiculate, 2.4-2.6 mm in diameter. This species is found in the shade of moist forest areas of the Pacific coastal regions of Panama and Costa Rica. In Costa Rica it has been collected in Puntarenas and San Jose provinces at elevations of 5 to 340 m. L. D. Gomez 2208 (F, MO, NY) was not included in this description because of the corolla color described as yellowish red and its location at 2100 m elevation in Heredia, although in other respects this specimen very closely resem- bles R. tonduzii. Flowering collections have been made from February through August. Ruellia tonduzii is recognized by its axillary, dichotomously branching panicles on glabrous, quadrangular peduncles (ca. 5 to 9 cm long), flow- ers borne singly on short pedicels (to 3 mm long), elliptic-oblong bracts (8 to 13 mm long), calyx lobes (2.7 to 4 mm long), and capsules (12 to 15 mm long). It is easily confused with R. pittieri and R. stemonacanthoides. (See the description of R. pittieri to distinguish these differences.) Ruellia tubiflora Kunth in H.B.K., Nov. gen. sp. 2: 241. 1817. Figure 17. Ruellia tubiflora H.B.K. var. tetrastichantha (Lin- dau) Leonard, Contr. U.S. Natl. Herb. 31:91. 1951. R. tetrastichantha Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 2: 300. 1900. Fl. Costaricensis. 2: 300. 1900. SufTrutescent plants to 3 m tall; internodes between leaf-bearing nodes 4.5-1 1 cm long, 2.5-4.5 mm thick, quadrangular, puberulous. Leaves on petioles to 5 cm long, puberulous; laminae elliptic, 10-23 cm long, 3.5- 10 cm broad, apically acuminate, basally attenuate, mar- gins undulate, glabrous above, puberulous on costa and veins to glabrous beneath, gland-dotted beneath, occa- sionally gland-dotted above, cystoliths visible above, ob- scure beneath. Inflorescences in compressed, short, ter- minal, and subterminal spikes with a rachis to 1.2 mm long; bracts, pale green to greenish white, leafy, elliptic, 4-5.5 cm long, 12-18 mm broad, gland-dotted; brac- teoles small, subulate, ca. 2 mm long. Flowers sessile; calyx segments subulate, 3.5—4 mm long, 1 mm broad basally, puberulous; corolla white or yellow, funnelform, 7.5-9 cm long, puberulous, often gland-dotted, unex- panded portion of tube 3-3.5 cm long, 2.5-3.5 mm broad, expanded portion 1 5-23 mm broad at the mouth, lobes suborbicular, 9-15 mm long, 1 1-20 mm broad; stamens included. Fruits clavate, 2.2-3 cm long, 7.4-8 mm broad, 6 mm thick, glabrous to minutely puberulous; seeds 8- 12, suborbicular, 4.5-5.5 mm in diameter. Comparatively wide ranging, this species is found in shaded areas of wet forests or thickets at lower to middle elevations of Costa Rica, Panama, and Colombia. In Costa Rica it has been collected in Alajuela, Cartago, Limon, and Puntarenas prov- inces at elevations of 450 to 900 m. Flowering collections have been made in January through March. Ruellia tubiflora var. tetrastichantha is recog- nized by its inflorescences of very short, terminal spikes (1.2 cm or less long), its white, funnelform corollas (7.5 to 9 cm long) which are often gland- dotted, its leaves which are gland-dotted beneath, and its pale green to greenish white bracts which are also gland-dotted. It most closely resembles R. tubiflora var. tubiflora and var. hirsuta. Variety tubiflora is distinguished by its red corolla, and var. hirsuta, by its subtomentose internodes in younger stems and its leaves with relatively long pilose hairs on both surfaces. It can also be con- fused with R. praeclara, which differs in its axillary flowers, its calyx segments (1 to 3 cm long) with sericeous hairs combined with numerous shorter glandular hairs, and its puberulous glandular co- rolla. 80 FIELDIANA: BOTANY Ruellia tubiflora H.B.K. var. hirsuta Leonard, Contr. U.S. Natl. Herb. 31:93.1951. Figure 1 7. This variety resembles R. tubiflora var. tetra- stichantha most closely and differs mainly in its hairiness. Leafy internodes are subtomentose, and the leaves and bracts are pilose on both surfaces. This variety is also found in central Colombia. Only one collection, Allen 6316 (F, MO), has been made in Costa Rica in the Esquinas Forest be- tween Rio Esquinas and Palmar at 30 m. The collector described it as very common in the forest. This flowering collection was made in late No- vember. Sanchezia Ruiz & Pavon REFERENCES— E. C. Leonard, Notes on the genus Sanchezia, J. Wash. Acad. Sci. 16: 484-492. 1926. E. C. Leonard and L. B. Smith, Sanchezia and related American Acanthaceae. Rhodora 66: 3 1 3- 343. 1964. Erect or climbing herbs or shrubs, mostly glabrous. Leaves petiolate; laminae with numerous cystoliths on both surfaces, glabrous. Flowers solitary or more often fascicled, those that are fascicled subtended by bracts which are often large, partly connate and showy; calyx 5-merous; corolla usually large and showy, corolla tube cylindric. the lobes 5 and equal; stamens 2, usually exserted, the anthers 2-celled, mucronulate basally, pub- erulous, staminodes 2. Fruit an oblong capsule, 6-8 seeds; seeds orbicular. This genus contains about 59 species, all native to tropical America and found mainly in wet forest areas of the northern Andes with two exceptions: S. pennellii Leonard extends into the Darien Prov- ince of eastern Panama, and 5". parvibracteata is found in Mexico and Central America, but prob- ably escaped from cultivation. Sanchezia parvibracteata Sprague & Hutchinson, Bull. Misc. Inform. 253. 1908. Shrubs to 2.5 m tall, stems subquadrangular, glabrous; internodes between leaf-bearing nodes, 4-6 cm long, 4- 5.5 mm thick. Leaves with short, glabrous petioles to 3 cm long; laminae elliptic to obovate, 1 2-24 (35) cm long, 5-1 1 (1 5) cm broad, apically abruptly acuminate, basally attenuate, margins undulate-serrate, glabrous, costa and veins edged in yellow, cystoliths numerous and promi- nent on both surfaces, ca. 0.5 mm long. Inflorescences in sparingly branched terminal panicles to 20 cm long; bracts loosely imbricate, ovate, 15-20 mm long, 7.5-10 mm broad, apically obtuse, glabrous; bracteoles oblong ovate, ca. 1 5 mm long, 4 mm broad, apically rounded, glabrous; peduncles to 5 cm long, glabrous; rachis gla- brous. Flowers 3-5 per pair of bracts, sessile; calyx seg- ments oblong, 2-2.5 cm long, 3.5-5 mm broad, apically obtuse, minutely puberulous at apex outside, glandular- puberulous inside; corolla red or ochre, puberulous with yellowish appressed hairs, 4-5 cm long, 6-7 mm broad at the throat, 5 mm broad just above ovary, lobes oblong, reflexed, 5-7 mm long, 5 mm broad; stamens exserted to 8 mm beyond the mouth of corolla, filaments sparsely villous, anthers 6-7 mm long; staminodes 1 5 mm long, sparingly pilose. Fruit not seen. In Central America this species is usually found in gardens. It is probably native to the Northern Andes since collections have been made there (Leonard, 1951). Only one Costa Rican collection has been observed. This was a flowering collection from a thicket in the Monteverde area, altitude 1 500 m, Puntarenas Province, Palmer 36769 (CR). Sanchezia parvibracteata is recognized by its flowers in fascicles of three to five loosely enclosed by a bract, its large red or ochre, equally five-lobed corolla, and its exserted stamens with two-celled anthers which are basally spurred. It is not easily confused with other Acanthaceae from Costa Rica. Spathacanthus Baillon Shrubs or small trees. Leaves petiolate; laminae usu- ally large, oblong, or elliptic. Flowers large, few, borne on small terminal branches; bracts small; calyx spatha- ceous, valvate; corolla usually large, white, the tube sal- verform, the limb 5-lobed, bilabiate, the lobes subequal; stamens 4, didynamous; anther lobes parallel. Fruit a capsule, large, glabrous. There are five species in this genus of Mexico and Central America. Because of the spathaceous, two-lobed calyx, this genus is not easily confused with other genera of the family. Spathacanthus hoffmannii Lindau, Bull. Herb. Boissier 3: 370. 1895. Figure 17. Shrubs or small trees to 8 m tall; internodes between leaf-bearing nodes 1.5-4.5 cm long, 2-3 mm thick, gla- brous. Leaves subsessile to petioles 10 mm long, gla- brous; laminae elliptic to elliptic-obovate, 7.5-27 cm BURGER: FLORA COSTARICENSIS 81 long, 3-7.5 cm broad, apically acuminate, basally long attenuate, margins undulate, glabrous on both surfaces, cystoliths dense and prominent on both surfaces. Flow- ers axillary, solitary, in clusters of 2 or 3 or in small racemes, peduncles to ca. 2 cm long, glabrous; pedicels 6-10 mm long, glabrous; bracts subulate, ca. 2 mm long, 1 mm broad basally, bracteoles similar but smaller. Flowers with a spathaceous calyx 2.2-3.5 cm long, gla- brous, fused for ca. half its length and then separating into 2 mostly equal lobes that are apically acute and apiculate, lobes 8-12 mm broad; corolla white, 8-8.7 cm long, glabrous, ca. 2 mm broad basally, expanding to 13-15 mm broad at the mouth, the lobes subequal, obtuse, 12-15 mm long, 12-15 mm broad; stamens di- dynamous, included, anthers 2-celled, ca. 4.5 mm long, basally rounded. Fruits clavate, 7-8 cm long, 12-14 mm broad, ca. 1 1 mm thick, glabrous; seeds 4, flattened, suborbicular, 9-10 mm in diameter, 2-3 mm thick, sur- face dark brown, rugose. This species is found in moist habitats at middle elevations in forest clearings most frequently along streams and rivers. I have seen no collections of it outside Costa Rica where it has been collected at elevations of 750 to 1750 m, mostly in the Caribbean watershed areas of Alajuela, Cartago, and Heredia provinces. One collection, Standley & Valeria 44745 (F) is known from the Pacific watershed, Tilaran area, Guanacaste. Flowering collections have been made in July through Oc- tober and in January (the Guanacaste collection). Spathacanthus hoffmannii is recognized by its woody habit, its large elliptic to elliptic-obovate leaves, its large, showy, white flowers with a five equally lobed corolla, two large, lobed spathaceous calyx and strongly didynamous stamens with two- celled anthers which are rounded basally, and its large, clavate fruit. With this combination of char- acters, it is not easily confused with other Acan- thaceae. Streblacanthus Kuntze Herbs to suffrutescent plants. Leaves petiolate. Inflo- rescences of lax terminal and axillary spikes; bracts nar- rowly oblong to filiform, long and conspicuous. Flowers with a 4-merous calyx, segments narrow and equal; co- rolla tube expanded basally, narrowed to a slender throat, the limb bilabiate, spreading, the upper lip 2-lobed, the lower lip 3-lobed; stamens 2, exserted, the filaments short, attached to the throat of the corolla, anthers 2-celled, the cells unequal and widely separated, 1 usually fertile, calcarate basally, the other much smaller, rounded ba- sally and often abortive; staminodes absent. Fruit a slen- der clavate capsule; seeds 2-4. This is a genus of four species with one species each described from Mexico, Guatemala, Costa Rica, and Bolivia. Streblacanthus monospermus Kuntze, Revis. gen. pi. 2: 498. 1891. Streblacanthus macrophyllus Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and in Pitt., Prim. fl. costaric. 2: 306. 1900. Streblacanthus longiflorus Cufodon- tis, Arch. Bot. (Forli). 10: 48. 1934. Figure 17. Herbs to 1 m tall; internodes between leaf-bearing nodes 3-5.5 cm long, 3-5.5 mm thick, tetrangular, pu- berulous along 2 lines in younger portions to glabrous in older. Leaves on petioles to 2.2 cm long, puberulence of petioles that of the stem; laminae elliptic, 6-19 cm long, 2.5-8 cm broad, apically acuminate, basally acute, margins entire to sparingly crenulate, glabrous to spar- ingly strigose on both surfaces, cystoliths visible on both surfaces. Inflorescences in terminal and axillary spikes, to 25 cm long, peduncle to 2.5 cm long, brownish pilose; rachis brownish pilose-glandular; bracts narrowly ellip- tic to oblong, 13.5-20 mm long, 3-4 mm broad, densely pilose-glandular, bracteoles long-subulate, 6-10 mm long, 1-2 mm broad basally, pilose-glandular. Flowers with 4-merous calyx, lobes subulate, 4—4.5 mm long, 0.8-1 mm broad basally, pilose-glandular; corolla lavender or white, salverform, puberulous, the tube 2-3 cm long, 1.5 mm broad expanding to 3 mm broad around the ovary, lobes of the upper lip obovate, 9-10 mm long, 7-8 mm broad, apically acute, lower lip very similar, lobes ca. 5 mm broad; stamens exserted just beyond the mouth with a tuft of pilose hairs at the apex of each filament; anther cells unequal and widely separated, the upper cell 1 mm long, basally calcarate, the lower 0.5 mm long and pro- ducing pollen. Fruits clavate, long-stipitate, 2-2.5 cm long, 5.5-7 mm broad, ca. 2 mm thick, pilose-glandular; seeds 4, orbicular, 5-6 mm in diameter. This species is apparently endemic to Costa Rica where it is found in lowland rain forests of the Caribbean coastal region. Flowering collections have been made in March, May, and September. Since the type collection was made at the Rio Yor- kin at the Panamanian border, this plant should also be found in adjacent Panama. Streblacanthus monospermus is recognized by its white to lavender salverform corollas with two exserted stamens on very short filaments and widely separated anther cells, the upper cell cal- carate and the lower cell much smaller and lacking a spur. After examining the types of S. mono- spermus, Kuntze s.n. (us) and S. macrophyllus, Lindau 8547 (us), I have concluded that these are the same species. The only difference between them is in the length of the corolla lobes, and more 82 FIELDIANA: BOTANY recent collections of S. monospermus show that the original measurements were made from im- mature corollas. Teliostachya Nees REFERENCES— C. G. Nees von Esenbeck, Acan- thaceae. In DC, Prodr. 11: 262-264. 1847. C. E. B. Bremekamp, Notes on the Acanthaceae of Suri- nam. Recueil Trav. Bot. Neerl. 35: 149. 1938. Herbs, ascending or decumbent. Leaves petiolate; laminae ovate to lanceolate with numerous cylindrical cystoliths, the margins entire. Inflorescences in dense spikes either terminal or terminal and axillary, the spikes either cylindrical or ovoid; pedicels arranged in whorls of 3-7, the lateral flowers of each whorl subtended by a 1 - to 3-nerved bract; 2 bracteoles subtending the lateral flowers, 1 -nerved. Flowers with a 5-merous calyx, seg- ments unequal, divided nearly to the base, the posterior lobe much larger than the rest, 3-nerved, the lateral lobes 1 -nerved, the anterior lobes 1- or 2-nerved, longer than the lateral nerves; corolla about as long as the calyx, the tube cylindric, 2-lipped, the upper lip obtuse, emarginate or subentire, the lower lip 3-lobed, the lateral lobes ob- tuse, the middle lobe emarginate or obtuse; stamens 4, didynamous, the anthers 2-celled, unequal, the lower cell and sometimes both basally apiculate; stigma 2-lobed, the style curved toward the upper lip. This is a genus of about 1 0 species occurring throughout much of tropical America and in West Africa. Only one species occurs in Central Amer- ica. It closely resembles Lepidagathis, a genus en- demic to the Old World. According to Breme- kamp, Teliostachya differs in its definitely terminal spikes, calyx segments which are almost entirely free as opposed to at least partially united seg- ments, its unequal anther cells as opposed to equal cells, and its pollen grains which are entirely free from reticulations as opposed to the finely retic- ulate grains of Lepidagathis. 2.5-8 cm long, 1 .5-3.2 cm broad, apically acute to slight- ly acuminate, basally attenuate, margins entire, gla- brous above, strigose mostly on costa and veins beneath, cystoliths dense on both surfaces. Inflorescences in dense terminal or axillary cylindrical spikes to 5 cm long, 1.3 cm broad; flowers in sessile whorls of 3-5; 1 bract sub- tending each flower, obovate lanceolate, 5-6 mm long, 1.5-2.2 mm broad, apically caudate, basally acute, thin, translucent, veins prominent, 3-nerved, transparent when wet, the margins ciliate, bracteoles 2, lanceolate linear, ca. 5 mm long, 0.4 mm broad, translucent with prom- inent veins, 1 -nerved, ciliate. Flowers with a 5-merous calyx, segments unequal, the posterior pair oblanceolate, 4-5.5 mm long, the remaining segments lanceolate, 3-6 mm long, all venose, ciliate; corolla white to violet, tu- bular, 3.5-4 mm long, glabrous, 2-lipped, upper lip ca. 2.5 mm long, 1 mm broad, bidentate, lower lip 2.5 mm long, 3-lobed, lobes ca. 1 mm long; stamens exserted. Fruits oblong, 3.3-3.7 mm long, 1-1.1 mm broad, 0.8- 1 mm thick, apically acuminate, basally acute, seeds 4, near ovate, flattened ca. 1 mm long, 8 mm broad, basally oblique, pilose when dry, hairs densely mucilaginous when wet. This species occurs in moist areas along streams and trails in rain forests, in thickets, and in open fields. Its range includes most of South America, Central America, and the West Indies. Only one Costa Rican collection has been seen: Weston et al. 4292 (us) from near Repunta, 1 5 km south of San Isidro, San Jose Province, altitude 600 m. This flowering collection was made in January. Collec- tions from neighboring countries have been made from near sea level to 1400 m elevation. Teliostachya alopecuroidia is a short, small- leaved herb that is recognized by its dense, ter- minal spikes with flowers containing a fi ve-merous calyx with very dissimilar segments, the posterior and anterior segments longer and broader than the lateral segments, and its inconspicuous two-lipped corolla with four didynamous anthers which are two-celled and cells superposed. It is not easily confused with other species of Acanthaceae. Teliostachya alopecuroidea (Vahl) Nees in Mart., Fl. bras. 9: 72. 1 847. Ruellia alopecuroidea Vahl, Eclogae americanae 2: 49. 1798. Lepidagathis alopecuroides (Vahl) R. Br. ex Griseb., Fl. Brit. W.I. 453. 1864. Figure 18. Herbs to 25 cm tall; internodes between leaf-bearing nodes 2-5.5 cm long, 1-1.5 mm thick, quadrangular, puberulous mostly at the angles. Leaves on puberulous petioles to 1 cm long; laminae ovate to ovate-elliptic, Tetramerium Nees REFERENCE— G. B. Happ, Monograph of Tetra- merium and Henrya. Ann. Missouri Bot. Gard. 24: 501-582. 1937. Herbs to suffrutescent plants; stems erect or declining, much-branched, pubescence often arranged in two op- posite lines, the old bark often exfoliating. Leaves sessile or petiolate; laminae linear to ovate, glabrous or pubes- cent. Inflorescences in terminal and axillary 4-sided BURGER: FLORA COSTARICENSIS 83 spikes; bracts opposite, 4-ranked, conspicuous, often closely imbricate with mucronate apex; pedicels 1-3 per bract. Flowers with a 4-5-merous calyx of equal, ciliate segments; corollas funnelform to tubular, bilabiate, the upper lip entire, the lower lip 3-lobed, the lobes ca. equal; stamens 2, included in the upper lip, the anthers 2-celled, cells slightly unequal. Fruits clavate, apiculate, 4- or 2- seeded; seeds flattened, tuberculate, or muriculate. Twenty-three species have been recognized for this genus, and they range from the southern United States to Colombia and Ecuador. Tetramerium nervosum Nees in Benth., Bot. voy. Sulphur 148, pi. 48. 1844. T. hispidum Nees in DC., Prodr. 11: 468. 1847. Figure 18. Repent to upright herbs to ca. 30 cm tall; internodes between leaf-bearing nodes 2.5-6.5 cm long, 1.25-1.5 mm thick, terete, sparingly pilose. Leaves on pilose pet- ioles to 15 mm long; laminae ovate, 2-7 cm long, 0.8- 3 cm broad, apically acute to acuminate, basally rounded to acute, margins entire, sparingly strigose on both sur- faces, numerous cystoliths visible on both surfaces. In- florescences in terminal and axillary spikes to 7.5 cm long, 1.5 cm broad; bracts clearly 4-ranked, imbricate, ovate to elliptic, 7-1 1 mm long, 3.5-7 mm broad, api- cally mucronate, typically curved outward, basally ob- tuse, mostly 5-nerved, puberulous plus longer pilose hairs, ciliate; bracteoles narrowly lanceolate, 4-6.5 mm long, ca. 1 mm broad, ciliolate; peduncle 1-4 mm long, puber- ulous. Flowers with 4-merous calyx, calyx segments sub- ulate, 3-3.5 mm long, ca. 0.5 mm broad, ciliolate; corolla white or slightly yellowish, bilabiate, 16-17 mm long, the tube narrowly funnelform, 6.5-7.5 mm long, ca. 1.25 mm broad at the throat, 1 mm broad basally; the upper lip of one lobe, the lower lip 3-lobed, all lobes oblan- ceolate, 7-9 mm long, 2.5-3.5 mm broad, the lobes of lower lip at the narrower range; stamens exserted to ca. the length of the upper lip, attached to base of middle lobe of lower lip, cells of the anthers slightly unequal, ca. 1.5 mm long, gray or bluish. Fruit clavate, 5-6 mm long, 2.25-2.5 mm broad, 1.75-2.25 mm thick, puber- ulous; seeds 4, rhomboid, 1 .75-2 mm long, 1.5-1.75 mm broad, surface tuberculate. This species is found in secondary growth pas- tures, along roadsides and trails throughout trop- ical America, and extends into the southwestern United States. In Costa Rica it is found from near sea level to 500 m elevation. It has been collected from Guanacaste and the Pacific Coastal area of Alajuela. Because this is a weedy species and omit- ted in collecting, it is of much more common oc- currence in Costa Rica than these collections would suggest. Flowering collections have been made from December through March. Tetramerium nervosum is recognized by its dense, terminal and axillary spikes with common- ly outcurving, imbricate, mucronate bracts that are clearly four-ranked, its bilabiate, four-parted white or yellowish corollas with the two exserted stamens attached at the base of the middle lobe of the lower lip, and its two-celled anthers that are gray or bluish. It is sometimes confused with Ble- chum brownei, but can be distinguished from it by its four-merous calyx and corolla (vs. five-mer- ous) and its two stamens (vs. four that are didyn- amous). Thunbergia Retz. REFERENCE— C. E. B. Bremekamp, Delimitation and subdivision of the Acanthaceae, Bull. Bot. Surv. India 7: 21-30. 1965. Herbaceous to suffrutescent vines or shrubs. Leaves petiolate with hastate, rhomboid, or ovate laminae, cys- toliths absent. Flowers large, peduncled, solitary in the axils or in terminal or axillary racemes; bracts 2, folia- ceous, large; calyx short, cupuliform, truncate, or toothed; corolla 5-merous; stamens 4, didynamous, attached near the base of the corolla tube, anthers 2-celled with an apiculate connective; ovary fleshy, the style, apically di- lated; ovary subtended by a conspicuous, fleshy nectary. Fruit a 2-chambered capsule with a globose base ex- tending abruptly into a flattened beak; seeds 2 in each locule, borne on papilliform funicles, semiglobose to ovoid, ventral side conspicuously excavated. This is a genus of about 200 species mainly, if not exclusively, from tropical Africa and Asia. Key to Species of Thunbergia 1 a. Petioles winged; corollas orange or yellow, often with a dark purple throat (rarely white) T. alata Ib. Petioles not winged; corollas blue, scarlet, white, or yellow 2a 2a. Flowers solitary in the leaf axils; calyx reduced to 1 0-20 teeth 3a 2b. Flowers in terminal or axillary racemes; calyx reduced to a basal ring or annulus 4a 3a. Erect shrub; laminae basally acute T. erecta 84 FIELDIANA: BOTANY 3b. Vine; laminae basally hastate to cordate T. fragrans 4a. Corolla with blue lobes, yellowish or white throat; bracts irregularly oblong, 3-3.5 cm long T. grandiflora 4b. Corolla scarlet, tube yellow; bracts ovate, 2-2.5 cm long T. mysorensis Thunbergia alata Bojer ex Sims, Bot. Mag. 52: pi. 2591. 1825. Figure 18. Herbaceous trailing or climbing vine; internodes be- tween leaf-bearing nodes 4.5-13 cm long, 1-1.25 mm thick, strigose. Leaves with petioles to 6.5 cm long, pet- ioles winged ca. 1 mm broad on each side, hirsute; lam- inae deltoid to deltoid-ovate, 3.5-7.5 cm long, 2.5 cm broad, apically acute, basally hastate to cordate, margins undulate, sparingly to moderately hirsute on both sur- faces. Flowers axillary and solitary, peduncles to 8.5 cm long, hirsute; bracts 2 per flower, deltoid-ovate, 18-20 mm long, 9-10 mm broad, apically acute hirsute; calyx toothed with ca. 15-17 subulate lobes ca. 2 mm long, glandular-puberulous; corolla yellow or orange with dark purple throat; salverform, ca. 4 cm long, tube 2 cm long, lobes 5, ca. 2 cm long, apically emarginate. Fruits densely puberulous, 16-18 mm long, base ca. 7 mm in diameter, beak ca. 1 1 mm long, 5 mm broad, 2 mm thick; seeds 4, ca. 3.5 mm in diameter, honeycombed. This species is native to eastern Africa and is widely cultivated in tropical America. It frequent- ly escapes from cultivation and is found in fields and along trails and the edges of thickets. In Costa Rica it has been collected from near sea level to about 1000 m elevation. Flowering collections have been made from November through March and in July. Thunbergia alata is recognized by its herba- ceous, viny habit, its deltoid to deltoid-ovate lam- inae with winged petioles, and its yellow or orange salverform corolla with five equal emarginate lobes and a dark purple throat. It is most easily distin- guished from other species of the genus by its leaves with winged petioles. There is a variety with white corollas and another with a white throat and or- ange lobes, but I have seen no collections of these from Costa Rica. Thunbergia alata most closely resembles T. fragrans, which has similar though white corollas and lacks winged petioles. Thunbergia erecta (Benth.) T. Anderson, J. Linn. Soc., Bot. 7: 18. 1864. Meyenia erecta Benth., Niger. Fl. 476. 1849. Shrub to 3 m tall; internodes between leaf-bearing nodes 2.5-6.5 cm long, 1.25-2.5 mm thick, quadran- gular, glabrous. Leaves with glabrous petioles to ca. 5 mm long; laminae elliptic, 1.5-6.5 (8.5) cm long, 0.8- 2.6 (4.3) cm broad, apically acute, basally acute, margins undulate, glabrous to subglabrous on both surfaces. Flowers solitary in leaf axils, peduncles to 2.8 cm long, glabrous; bracts oblong-ovate, 10-20 mm long, 5-8 mm broad, apically acute, basally obtuse, glabrous to mi- nutely puberulous at the apex. Flowers with the calyx reduced to 10-16 subulate teeth ca. 3 mm long, glan- dular-puberulous; corolla all white or deep violet lobes with a yellowish throat, broadly salverform, glabrous, 6-8 cm long, 6-8 mm broad basally constricting to 2-3 mm broad just above the ovary, expanding to 1 5-20 mm broad at the throat, lobes suborbicular, 15-25 mm in diameter; cells of the anther unequal, the base and margins of the anther cells puberulous. Fruit not ob- served. This species is a native of tropical West Africa, but is widely cultivated. It has become naturalized from cultivation in many tropical areas, but all Costa Rican collections I have observed were from cultivation. These were from near sea level in Li- mon Province and from about 1000 m elevation from San Jose. Flowering collections were made in June and November. Thunbergia erecta is recognized by its shrubby habit and its all white or violet, lobed, yellow- throated corollas from 6 to 8 cm long. The all white corolla may be confused with that of T. fragrans, and the violet, yellow-throated corolla may be confused with that of T. grandiflora, but both of these species have a viny habit and shorter corol- las. Thunbergia fragrans Roxb., PI. Coromandel 1: 47. 1795. An herbaceous to somewhat woody vine; internodes between leaf-bearing nodes 3-11.5 cm long, 1-2 mm thick, quadrangular, strigose. Leaves with strigose peti- oles to 2.5 cm long; laminae ovate to ovate lanceolate, 5-1 1.5 cm long, 2-6.5 cm broad, apically acute, basally hastate to cordate, margins entire to slightly undulate, strigose on both surfaces. Flowers solitary in the leaf axils, pedicels to 8 cm long, strigose; bracts deltoid ovate, 1.3-2 cm long, 5-8 mm broad, apically acute, strigose. Flowers with a calyx of 12-20 teeth, the teeth ca. 3 mm long, puberulous; corolla white, salverform, pilose, the tube 2-2.5 cm long, 3-5 mm broad, the lobes cuneate, BURGER: FLORA COSTARICENSIS nearly as long as the tube, crenate; anther cells ca. 3 mm long, basally acute. Fruit ca. 2.5 cm long, the beak sub- ulate, flattened, 1-1.5 cm long. A native of India, this species is rather widely cultivated, as it is in Costa Rica. Collections from cultivation have been made from near sea level to about 1 000 m elevation. Flowering collections were made in October, February, April, and June. Thunbergiafragrans is recognized by its twining habit and solitary flowers with a toothed calyx and a white corolla. It most closely resembles T. alata, which differs in its winged petioles and yellow co- rollas with a dark purplish throat. Thunbergia grandiflora Roxb., Hort. bcngal. 45. 1814. Suffrutescent climbing or twining vine; inter nodes be- tween leaf-bearing nodes 6-19 cm long, 2.6-3.2 mm thick, subquadrangular, glabrous. Leaves on scabrous petioles to 6 cm long; laminae ovate to ovate-hastate, 8-16 cm long, 3-13.5 cm broad, apically acuminate, basally hastate or rounded, margins often coarsely toothed or lobed below the middle, scabrous mostly on costa and veins on both surfaces. Inflorescences of elongated ter- minal and axillary racemes with 2-4 flowers at each node; peduncles to 4.5 cm long, quadrangular, glabrous to sparingly scabrous; pedicels to 3.3 cm long, sparingly scabrous; bracts irregularly oblong, 3-3.5 cm long, 17- 20 mm broad, apically acute, sparingly puberulous plus numerous dark glandular appearing dots. Flowers with a very much reduced annular calyx at the base of the corolla, margin of annulus puberulent; corolla light or dark blue with a white or yellowish throat, campanulate, the tube 3.0—4.5 cm long, lobes 3-4 cm long; bases of the anther cells of the posterior stamens each with spurs to 3 mm long, only 1 anther cell spurred on each of the anterior stamens. Fruit base ca. 1 6 mm in diameter, beak flattened, ca. 2 cm long. This species is native to Bangladesh and is widely cultivated in tropical regions of the world. Only collections from cultivation have been made in Costa Rica in the Central Valley area from Ala- jucla. Heredia, and San Jose provinces. Flowering collections were made in September and March. Thunbergia grandiflora is recognized by its twining habit, its loose terminal and axillary ra- cemes, its annular calyx, and its large, showy, cam- panulate corolla with blue lobes and a white or yellowish throat. It is not easily confused with oth- er cultivated species of Thunbergia. Thunbergia mysorensis (Wight) T. Anderson ex Bedd. in Trans. Linn. Soc. London 25. 222: 1 865. Hexacentris mysorensis Wight, PI. asiat. rar. Ill: 78. t. 871. 1832. A woody vine; internodes between leaf-bearing nodes 2-8.5 cm long, 1-1.5 mm thick, subquadrangular, gla- brous. Leaves with glabrous petioles to 1.6 cm long; laminae lanceolate-oblong, 5-9.5 cm long, 1.2-2.3 cm broad, apically attenuate, basally obtuse to subcordate, margins undulate, mostly 3-nerved, glabrous on both surfaces. Inflorescences in terminal and axillary racemes; peduncles to 6 cm long, glabrous; pedicels to 2.5 cm long, glabrous; bracts red, ovate, 2-2.5 cm long, 8-10 mm broad, apically acute, glabrous. Flowers with a very much reduced annular calyx at the base of the corolla; corolla zygomorphic, lobes scarlet, tube yellow, 5-5.5 cm long, throat opening obliquely with lower lobes re- flexed and 2 upper lobes fused most of their length to partially enclose the stamens and style, stamens with anthers ca. 8 mm long, basally long-spurred, spurs ca. 4 mm long. Fruits not observed. Native to India, this species is occasionally cul- tivated in Costa Rica for its showy inflorescences. I have observed only two Costa Rican collections, Jimenez 447 (F) and Sanchez 4 (F). Both were in flower, one collected in January, the other in March. Thunbergia mysorensis is recognized by its lan- ceolate-oblong leaves with obtuse to subcordate bases, its showy flowers in terminal or axillary racemes with red bracts, its annular calyx, and its zygomorphic corollas with scarlet lobes and yellow tubes. It is not easily confused with other Thun- bergia species grown in Costa Rica. Trichanthera H.B.K. Shrubs or trees. Leaves petiolate; laminae ovate to oblong, entire to undulate, numerous cystoliths visible on the upper surface of dried specimens. Inflorescences in compact terminal panicles (or corymbs). Flowers sub- tended by 2 small, triangular bracteoles; calyx 5-merous with sepals separate, equal and mostly obtuse; corolla regular, 5-lobed, tomentose, campanulate; stamens 4, exserted, didynamous, the anthers ciliate, with 2 equal basally rounded cells. Fruit a 2-chambered capsule with 2 seeds per locule. Trichanthera is a genus of two species ranging from Central America to Brazil. It is most easily confused with the genus Bravaisia, but can be dis- tinguished from it by its basally rounded anthers (vs. spurred) and one- to four-seeded capsules (vs. eight-seeded). Trichanthera gigantea (Humb. & Bonpl.) Nees in DC, Prodr. 11: 218. 1809. Ruellia gigantea 86 FIELDIANA: BOTANY Humb. & Bonpl., PI. aequinoct. 2: 75, pi. 102. 1809. REFERENCE— T. B. Croat, Flora of Barro Colo- rado Island, p. 790. Stanford Univ. Press. 1978. Shrubs or small trees to 10 (17) m tall; internodes between leaf-bearing nodes 4-6 mm thick, tetrangular, brown tomentose. Leaves with petioles 1.5-6 cm long; laminae ovate to elliptic, 10-26 cm long, 6-10 cm broad, apically acuminate to acute, basally acute to obtuse or oblique, margins entire to slightly undulate, the surfaces glabrous except sparingly pilose on the costa of larger leaves. Inflorescences in terminal, compact panicles tending to be secund, to 1 5 cm long, 6 cm broad, branch- es light brown tomentose. Flowers subtended by 2 tri- angular bracteoles to 3 mm long; sepals 10-12 mm long, tomentose, apically rounded to acute; corollas shiny yel- low in the throat, brownish red in the expanded portion, 3-4 cm long, glabrous at throat, silky tomentose above, silky tomentose outside, the tube 1-1.5 cm long, the limb 2-3 cm across; lobes 4-6 mm broad, oblong to oblong- ovate; ovary brown tomentose, style 4-5 cm long. Fruits 1.5-2 cm long, 5 mm broad, apically obtuse, sericeous; seeds flattened, lenticular, 3-4 mm in diameter. This is a wide-ranging species of moist lowland forests, frequently along streams and swampy areas. While it is not abundant in areas where collected, it extends from Costa Rica through Colombia and Venezuela to the mouth of the Amazon in Brazil. Although T. gigantea occurs in Costa Rica (Leon- ard, 1938), this description is based upon Pana- manian collections. On Barro Colorado Island, Panama, Croat has observed flowering from Jan- uary to April, especially in February and March. Trichanthera gigantea is recognized by its ar- borescent habit; paniculate inflorescences with to- mentose sepals and corolla; more or less equally five-lobed corollas 3 to 4 cm long; didynamous stamens with two-celled anthers that are basally rounded; sericeous capsules with one to four seeds. It may be confused with Bravaisia integerrima, which differs in its basally spurred anthers and glabrous capsules that are eight-seeded. PLANTAGINACEAE By William Burger Herbs (rarely subshrubs with few-branched woody stems in island endemic species ofPlantago), annual or perennial, stems usually very short above the ground with closely congested internodes, xylem in discrete bun- dles or forming a cylinder, a rhizome often present; stip- ules absent. Leaves alternate (rarely opposite), simple and usually from basal rosettes, petioles often not strong- ly differentiated from the lamina, leaf-base clasping the stem; laminae linear to broadly ovate, entire or with small lobes or teeth, glabrous or puberulent, venation palmate or parallel. Inflorescences solitary from each leaf axil, flowers usually in dense spikes or capitula on a long, unbranched peduncle, each flower subtended by a single bract. Flowers small and sessile or subsessile, bisexual (in ours) or unisexual, radially symmetrical and usually 4-parted (rarely 3-parted); sepals 4 (3), free or partly united, imbricate in bud; petals united to form a short tube, the 4 (3) corolla-lobes spreading or reflexed at an- thesis, membranous to scarious, white to yellowish, often persisting; stamens as many as the corolla-lobes and al- ternate with them (1 or 2 in Bougueria), filaments borne on the corolla-tube and inflexed in bud, anthers exserted and versatile, 2-thecous, introrse and longitudinally de- hiscent, pollen 4-20-porate; pistil 1 , ovary superior, usu- ally 2-(3, 4)-locular in Plantago with 1 to many ovules in each locule, 1-locular with a single basal ovule in Bougueria and Littorella, style 1 , stigma 1 - or 2-lobed. Fruit a thin-walled capsule opening at or below the mid- dle circumscissilly (a pyxis) in Plantago, a small nut in Bougueria and Littorella; seeds often lustrous, endo- sperm present (absent in Littorella), embryo straight (curved in Bougueria). A family of three genera and about 250 species in temperate, arctic, and montane habitats. Bou- gueria nubicola is found in the high Andes of southern South America. Littorella is a genus of semiaquatic, freshwater habitats, with one species in Europe, one in North America, and a third species in southern South America. Plantago is cosmopolitan in cooler or montane habitats and the only genus found in Central America. While its close relationships are obscure, the tubular co- rolla and bilocular ovary with axile placentation (in Plantago) has suggested an affinity with the Scrophulariales. Plantago Linnaeus Herbs (in ours) or subshrubs, annual or perennials, usually scapose plants with very short stems, clustered basal leaves and erect, unbranched peduncles bearing small, congested flowers. Leaves all basal and rosulate in Central American species, alternate and tightly con- gested (in ours), petioles present or absent, poorly dif- ferentiated from the lamina, clasping the stem at their base; laminae linear to broadly ovate, glabrous or more often puberulent, entire or with short blunt lobes or teeth, venation palmate or parallel. Inflorescences spicate or capitate with long, erect, unbranched peduncles, flowers sessile in the axils of bracts. Flowers small and bisexual, sessile in the axils of bracts, sepals 4, free or partly united, often unequal with 2 longer and 2 shorter, imbricate in bud, margins scarius, petals united to form a short tube, BURGER: FLORA COSTARICENSIS the 4 lobes reflexed or spreading at anthesis, persistent and sometimes forming a cover over the fruit; stamens 4 or 2, exserted at anthesis; ovary 2-locular or 3- or 4- 1 ocular by intrusion of the placentae, ovules 1 to several in each locule. Fruit a thin-walled capsule with circum- scissile dehiscence around the middle; endosperm pres- ent, embryo straight. A genus of worldwide range but restricted to temperate, arctic, and montane environments; ab- sent in the lowland tropics except on islands. A number of island endemics have unusual woody growth forms, but most species have a very short aerial stem, leaves arising from near the ground and erect scapose inflorescences. These plants can be mistaken for sedges and other monocotyledons because of the often narrow leaves with parallel venation and the small, tightly clustered flowers subtended by brownish bracts. The small flowers with short corolla-tube, four scarious corolla-lobes, long, slender filaments, exserted versatile stamens, and fruit with circumscissile dehiscence easily dis- tinguish these plants. Dr. Knud Rahn has revised many New World species (see Nordic Journal of Botany 3: 331-342, 1983, and included refer- ences). Key to Species of Plantago la. Leaves becoming broadly ovate, lamina abruptly narrowed at the base (in larger leaves); corolla- lobes 0.5-1 mm long; ovary with many ovules, seeds usually more than 8 P. major 1 b. Leaves elliptic to oblanceolate, laminae tapering gradually at the base to join the poorly differentiated petiole; corolla-lobes 2-3 mm long; ovary with 6 or fewer ovules, seeds fewer than 7 2a 2a. Corolla-lobes persisting and erect in fruit, forming a pointed "cap" at the apex of the fruit; ovary with 3-6 ovules, seeds 1-6; native and common P . australis 2b. Corolla-lobes not persisting as a cap above the fruit; ovary with 2 ovules; seeds 1 or 2; introduced, rare . .P. lanceolata Plantago australis Lamarck, Encycl. Meth. Bot. Illust. Gen. 1: 339. 1793. P. hirtella H.B.K., Nov. gen. sp. 2: 187 or 229, t. 127. 1817. P. hartwegii Decne. in DC, Prodr. 13, pt. 1: 724. 1 852. P. schiedeana Decne. in DC., loc. cit. 723. P. sodiroana Pilger, Bot. Jahrb. Syst. 25: 733. 1898. P. ecuadorensis Pilger, loc. cit. 50: 232. 1913. P. australis ssp. hirtella (H.B.K.) Rahn, Bot. Tidsskr. 60: 50. 1964. Figure 19. Herbs, perennial, stems very short above the ground, internodes congested and obscured by the leaf-bases. Leaves usually in basal rosettes, quite variable in size on different plants, petioles to 15 cm long; laminae 4-25 (35) cm long 0.5-5 (7) cm broad, lanceolate to oblan- ceolate or narrowly elliptic-obovate, obtuse to acute at the apex, gradually narrrowed to the base and continuous with the petiole, margin entire or slightly undulate, very sparsely (rarely glabrous) to densely puberulent on both surfaces with slender septate and translucent hairs 0.5- 1 .5 mm long, drying thin-chartaceous and greenish, ve- nation parallel with (3) 5 or 7 primary veins. Inflores- cences 3-50 (100) cm long, peduncle sparsely to densely whitish villous with hairs to 2 mm long, flowering por- tion 3-50 cm long, flowers usually congested near the apex, more separate near the base, bracts 1.6-4.2 mm long, 0.6-1.4 mm wide, triangular to ovate, usually cil- iate along the margin. Flowers with sepals 1.7-2.7 mm long, ca. 1 mm broad, 1 often longer than the other 3, usually puberulent along the keel and ciliate along the margin distally, corolla-lobes 2-3.5 mm long, narrowly ovate, usually erect and spread apart only at anthesis, persistent and becoming stiff in fruit; anthers 1.2-1.6 mm long; ovary usually with 3 ovules. Fruit 2—4 mm long, to 6 mm together with the persisting beaklike im- bricate corolla-lobes, pale brown; seeds 1.2-2.4 mm long, ca. 1 mm broad, oblong, flat and with a small circular scar on the inner face, pale brown, smooth. Plants of open, sunny sites in regions of wet, evergreen, montane forest formations between 1500 and 3200 m elevation in Costa Rica; to as low as 400 m in northern Central America. Inflo- rescences are produced throughout the year, but there may be a flowering peak in August. The species ranges from Arizona and Mexico through the Central Highlands of Central America and western Panama through the highlands of northern and central South America to Bolivia, Brazil, and Argentina. Plantago australis is our only native species of Plantago and can be recognized by the stiff, erect corolla-lobes that are persistent and form a "beak" on the apex of the fruit. The corolla-lobes appear to be open for a very short period during anthesis and are otherwise erect and imbricate. As with other species, the name Llanten is commonly used for these plants. Rahn distinguishes a number of subspecies; most of our material would appear to belong to ssp. hirtella (H.B.K.) Rahn. 88 FIELDIANA: BOTANY FIG. 19. Plantaginaceae: Plantago australis. A, Flower viewed from above; B, a small plant; C, seeds (1-mm scale); D, fruit with bracts and perianth (5-mm scale); E, fruit (same scale as D); F, a large plant. BURGER: FLORA COSTARICENSIS Plantago lanceolata L., Sp. pi. 113. 1753. Herbs, annual or becoming perennial, aerial stem very short with congested internodes, secondary roots nu- merous. Leaves erect or spreading laterally, petiole weakly differentiated from the lamina, clasping the stem at the base; lamina 4-28 cm long, 0.5-4 cm broad, lanceolate to oblong-lanceolate, entire or with small teeth along the margin, glabrous or puberulent, the veins usually strong- ly impressed above. Inflorescences 1 5-60 (80) cm long, peduncle longitudinally ridged, spike 1.5-8 cm long, bracts 2.3-5 mm long, triangular. Flowers with sepals 2.4-3.5 mm long, the 2 anterior (abaxial) sepals almost completely united, posterior sepals ovate, corolla-lobes 1.8-2.8 mm long, thin, translucent, reflexed; anthers ca. 1.8 mm long; ovary with only 2 ovules. Fruit ca. 5 mm long, not covered by the persisting corolla-lobes; seeds 2.3-3 mm long, inner face concave. Plantago lanceolata has been reported on the slopes of Volcan Irazu and Volcan Turrialba (Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1264, 1938), but we have seen no specimens from Costa Rica or elsewhere in Central America. The species is now cosmopolitan and is natural- ized in many parts of the Andes; it can be a trou- blesome weed of lawns and grassland. Plantago major L., Sp. pi. 1 12. 1753. Herbs, perennial from a short, thick rootstock, inter- nodes tightly congested and hidden by the leaf-bases. Leaves in basal rosettes, flat on the ground or erect, petioles 2-1 5 cm long, expanded at the base and clasping the stem; 10-30 (50) cm long 3-12 (18) cm broad, ovate to elliptic-ovate or broadly elliptic, obtuse at the apex, the larger laminae abruptly narrowed to the base and obtuse to truncate, margins entire or irregularly short- dentate, drying chartaceous, glabrous or with slender septate translucent hairs 0.3-1 mm long, venation pal- mate with 3-7 (1 1) primary veins. Inflorescences spicate, 1 5-35 (50) cm long, flowering portion 4-28 cm long and 6-8 mm thick (dry), peduncle puberulent in early stages but becoming glabrous, flowers closely congested dis- tally, bracts 2-3 mm long. Flowers with sepals 1.2-2 mm long, broadly ovate to obovate, with thin margins, glabrous or minutely puberulent along the midvein abax- ially, corolla-lobes 0.5-1 mm long, narrowly triangular, obtuse or acute, rotate, not persisting as a covering on the fruit. Fruit 2-3.8 mm long, ovoid, usually with 5- many brown seeds; seeds ca. 1 mm long and 0.6 mm thick, variable in shape but mostly oblong. A cosmopolitan weed of open sunny sites such as recently cleared land, roadsides, and stream edges. It is naturalized in the Central Highlands between 500 and 1 500 m elevation and appears to flower throughout the year in Costa Rica. The plants have been used medicinally, fresh or ground as poultices; they are known by the name Llanten. The broad leaves and very short corolla-lobes are distinctive. 90 FIELDIANA: BOTANY Index The index includes all accepted names (in Roman type), synonyms (italics), common English names (Roman), and vernacular names (italics). New species are in boldface, and the page numbers of illus- trations are in boldface. Hyphenated words and multiple words are alphabetized by letter. Acanthaceae 1 Acanthoideae 2 Acanthus 23 mollis 2, 23 montanus 23 Adhatoda carthaginensis 52 umbrosa 50 Amasonia integerrima 33 Aphelandra 23 araoldii 30 aurantiaca 3, 24 aurantiaca var. aurantiaca 3, 24 aurantiaca var. stenophylla 3, 25 campanensis 3, 25 deppeana 28 dolichantha 3, 25 golfodulcensis 4, 26 haenkeana 28 leonardii 4, 26 lingua-bovis 4, 27 pectinata 28 pulcherrima 28 repanda 25 scabra 3, 27 seiberti 4, 28 sinclairiana 4, 26, 28 storkii 4, 29 tonduzii 5, 29 tridentata 5, 30 Aphragmia 75 inundata 72 Arrhostoxylum achimeniflorwn 76 stemonacanthoides 79 Asystasia 30 gangetica 5, 3 1 Barleria 3 1 discolor 3 1 m icans 5, 31 pyramidata 32 Beloperone 48 brenesii 5 1 guttata 5 1 urophylla 6 1 variegata 53 medium 32 brownei 5, 32 brownei f. puberulum 32 costaricense 5, 32 dariense 32 pyramidatum 32 Bravaisia 33 integerrima 6, 33 Buceragenia 33 glandulosa 6, 34 Carlowrightia 34 arizonica 34 costaricana 34 Chaetochlamys 35 panamensis 35 Chaetothylax 35 leucanthus 6, 35 Chamaeranthemum 36 durandii 6, 36 tonduzii 36 cultivated genera 2 Dianthera candelariae 52 comata 53 Diapedium trifurcatum 39 Dicliptera 37 imbricata 6, 37 iopus 6, 38 pallida 7, 38 podocephala 7, 39 skutchii 7, 39 trifurca 7, 39 trifurcata 39 unguiculata 7, 40 Dipteracanthus nudiflorus 77 Dyschoriste 40 valeriana 7, 40 Ecbolium refractifolium 58 trichotomum 60 umbrosum 50 Elytraria 4 1 imbricata 7, 4 1 Eranthemum atropurpureum 68 cuspidatum 68 praecox 69 Glockeria blepharorhachis 43 sessilifolia 44 stricta 44 ventricosa 45 Graptophyllum 4 1 hortense 42 pictum 8, 42 Habracanthus 42, 70 silvaticus 8, 42 Hansteinia 43 blepharorhachis 8, 43 gracilis 44 sessilifolia 8, 44 stricta 8, 44 ventricosa 8, 45 Hemisandra aurantiaca 24 Henrya 45 scorpioides 9, 45 Herpetacanthus 46 panamensis 9, 46, 47 Hexacentris mysorensis 86 Hygrophila 47 conferta 47 costata 9, 47 Hygrophila guianensis 47 Hypoestes 47 phyllostachya 9, 47 Jacobinia 48 aurea 50 crenata 54 spicigera 59 tinctoria 59 umbrosa 50 Justicia 48 angustibracteata 9, 50 asymmetrica 60 aurea 9, 50 brandegeana 11, 51 brenesii 10, 51 candelariae 10, 51 carthaginensis 10, 52 chamaephyton 10, 53 comata 10, 53 costaricana 10, 53 crenata 11, 54 ephemera 54 gangetica 3 1 graciliflora 59 guttata 51 imbricata 41 macrantha 11, 55 metallica 11, 55 microphylla 50 oerstedii 11, 55 orosiensis 11, 56 parvibracteata 12, 56 pectoralis 12, 57 picta 42 pittieri 12, 57 pseudopolystachia 58 refractifolia 12, 58 refulgens 58 sarapiquensis 12, 58 scabra 28 secunda 61 skutchii 12, 59 spicigera 13, 59 tinctoria 13, 59 tonduzii 13, 60 trichotoma 13, 60 tubiformis 66 umbrosa 50 urophylla 13, 61 valerii 13, 61 Kalbreyeriella 70 Kolobochilus blepharorhachis 43 leiorhachis 1 1 Lepidagathis 83 alopecuroides 83 Llanten 88, 90 BURGER: FLORA COSTARICENSIS 91 Louteridium 62 costaricensis 14, 62 tamaulipense 62 Megaskcpasma 62 erythrochlamys 15, 62 Mendoncia 63 brenesii 14, 63 costaricana 14, 64 gracilis 63, 64 lindavii 14, 63, 64 littoralis 63 retusa 14, 65 tonduzii 14, 65 Mendoncioideae 1, 2 Meyenia erect a 85 Nelsonia 65,66 canescens 15, 66 Nelsoniodiae 1, 2 Odontonema 66 flagellum 66 strictum 66 tubiforme 15, 66 Onchyanthus speciosus 33 Pachystachys 2 Plantaginaceae 87 Plantago 87 australis 88, 89 australis ssp. hirtella 88 ecuadorensis 88 hartwegii 88 hirtella 88 lanceolata 90 major 90 schiedeana 88 sodiroana 88 Poikilacanthus 67 macranthus 15, 67 Pseuderanthemum 67 atropurpureum 15, 68 cordatum 34 cuspidatum 15, 68 pittieri 16, 69 praecox 16, 69 standleyi 16, 69 Razisea 70 breviflora 43 citrina 16, 70 spicata 16, 71 wilburii 16, 71 Rhytiglossa candelariae 51, 52 microphylla 50 Ruellia 72 achimeniflora 76 alopecuroidea 83 ha rhi liana 73 biolleyi 17, 73 blechum 32 caroliniensis 75 cooperi 17, 74 geminiflora 17, 74 golfodulcensis 17, 75 inundata 17, 75 jussieuoides 17, 76 longissima 73 longissima var. glabra 78 malacosperma 76 metallica 18, 77 nudiflora 18, 77 nudiflora var. puberula 77 palustris 18, 77 paniculata 18, 78 pittieri 18, 78 praeclara 18, 79 puberula 77 stemonacanthoides 19, 79 tonduzii 19, 80 tuberosa 75 Ruellia tubiflora 8 1 tubiflora var. hirsuta 19, 8 1 tubiflora var. tetrastichantha 19, 80 Sanchezia 8 1 parvibracteata 8 1 pennellii 81 Scrophulariaceae 1 Sericographis tinctoria 59 Sornia 32, 52 Spathacanthus 8 1 hoffmannii 19, 81 Standleyacanthus costaricanus 46 Streblacanthus 82 longiflorus 82 macrophyllus 82 monospermus 19, 82 Teliostachya 83 alopecuroidea 20, 83 Tetramerium 83 hispidum 84 nervosum 20, 84 scorpio ides 45 Thunbergia 84 alata 20, 85 erecta 65 fragrans 85 grandiflora 86 mysorensis 86 Thunbergioideae 1, 2 Thyrsacanthus flagellum 66 si rictus 66 Trichanthera 86 gigantea 20, 86 Tubiflora 4 1 squamosa 41 Zornia 32, 52 92 FIELDIANA: BOTANY 11 UNIVERSITY OF ILLINOIS-URBANA 30112028175997