HARVARD UNIVERSITY -^ Library of the Museum of Comparative Zoology L't- VOLUME 24 1983 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. Contrib- utors need not be members of the Tulane faculty. Manuscripts of 20 or more pages, double-spaced, are preferred. We recommend conformance with the principles stated in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double spaced. Two additional copies should accompany the original to expedite editing and publication. Legends for figures should appear on a separate page and in sequence. Illustrations should be proportioned for one or two column width corresponding to our printed page size, and should allow for insertion of the legend if occupying a whole page. Guidelines for letter and other extraneous markings should be done with a non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Lettering in any illustrative or tabular material should be of such a size that no letter will be less than 1 V2 mm high when reduced for publication. An abstract not exceeding three percent of the length of the article must accompany the manuscript. Separates of published articles are available to authors at a nominal cost. Page charges, calculated at $45/page, are solicited from authors who have funds for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs but excessive illustrations and tabular matter may be charged to the author. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- changes are invited from institutions publishing comparable series. Subscriptions are billed in advance. A price list of back issues is available on request. Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University." Subscription rates: Volume 24, $8.50 domestic, $9.50 foreign Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor CONTENTS OF VOLUME 24 Number Page 1. A REVISION OF THE MIDDLE AMERICAN THECOPHYLLOID VRIESEAS (BROMELIACEAE) JOHN F. UTLFY 1 2. FISHES OF THE BAYOU SARA DRAINAGE, LOUISIANA AND MISSISSIPPI, WITH A DISCRIMINANT FUNCTIONS ANALYSIS OF FACTORS INFLUENCING SPECIES DISTRIBUTION lAMFS M. (.RADY , ROBIRT C. C ASHNER AND JAMES S. ROGERS 83 PLANTS AND PLANT COMMUNITIES OF JEAN LAFITTE NA- TIONAL HISTORICAL PARK, LOUISIANA DAVID A. WHITE, STEVEN P. DARWIN, AND LEONARD B. THIEN 101 ISSN 0082-6782 f I^IL^SfH i^I0®I!IS§ urn Volume 24, Number 1 $5.50 May 9, 1983 A REVISION OF THE MIDDLE AMERICAN THECOPHYLLOID VRIESEAS (BROMELIACEAE) JOHN F. UTLEY tViuS. COMP. ZOOL LIBRARY UNlV£RSiTY TULANE UNIVERSrry NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. Contrib- utors need not be members of the Tulane faculty. Manuscripts of 20 or more pages, double-spaced, are preferred. We recommend conformance with the principles stated in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double spaced. Two additional copies should accompany the original to expedite editing and publication. Legends for figures should apf)ear on a separate page and in sequence. Illustrations should be proportioned for one or two column width corresponding to our printed page size, and should allow for insertion of the legend if occupying a whole page. Guidelines for letter and other extraneous markings should be done with a non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Lettering in any illustrative or tabular material should be of such a size that no letter will be less than 1 Vi mm high when reduced for publication. An abstract not exceeding three percent of the length of the article must accompany the manuscript. Separates of published articles are available to authors at a nominal cost. Page charges, calculated at $45/page, are solicited from authors who have funds for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs but excessive illustrations and tabular matter may be charged to the author. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- changes are invited from institutions publishing comparable series. Subscriptions are billed in advance. A price list of back issues is available on request. Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University." Subscription rates: Volume 24, $8.50 domestic, $9.50 foreign Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 24, Number 1 $5.50 May 9, 1983 A Revision Of The Middle American Thecophylloid Vrieseas (Bromeliaceae) John F. Utley Department of Biological Sciences. University of New Orleans, New Orleans. Louisiana 70148 CONTENTS Introduction 2 Habitat and Ecology 2 Morphology 5 Reproductive Biology 12 Intrageneric Relationships '4 Systematic Treatment '8 Artificial Key to the Species of Middle American Thecophylloid Vrieseas 18 1 . V. leptopoda 22 2. V. pedicellata 24 3. V. diffusa 27 4. V. triflora 28 5. V. greenbergii sp. nov 30 6. V. umbrosa 32 7. V. latissima 33 8. V. lyman-smithii sp. nov 35 9. V. nephrolepis 36 10. V. ororiensis ^^ 11. V. balanophora 46 1 2. V. luis-gomezii sp. nov 48 13. V. viridis 50 14. V. hainesiorum 52 15. V. leucophylla 53 1 6. V. vietoris 55 17. V. uxoris 57 18. V. picta 58 19. V. stenophylla 61 20. V. williamsii 62 21 . V. attenuata 64 22. V. comata 66 23. V. singuliflora 68 24. V. hygrometrica 69 25. V. kathyae sp. nov 71 26. V. notata 73 27. V. bracteosa 75 28. V. capitata 77 Doubtful and Excluded Taxa 78 Literature Cited 80 1 Tulane Studies in Zoology and Botany Vol. 24 Abstract The thecophylloid vrieseas were long maintained in a distinct genus, Thecophyllum, because of their enlarged primary bracts and reduced secondary inflorescence branches. These taxa currently form a geographically and morphologically cohesive alliance within section Xiphion of Vriesea. In addition to the nocturnal flowers generally encountered in section Xiphion several thecophylloid vrieseas display crepuscular and diurnal anthesis and floral syndromes suggestive of adaptation to a wide range of pollen vectors. Keys, descriptions and synonymies are provided for the Middle American taxa and their relationships are discussed. Vriesea green- bergii, V.kathyae, V.luis-gomezii and V.lyman-smithii are illustrated and described as new species. Introduction The genus Thecophyllum was described by Andre, on the basis of enlarged primary bracts and aborted lateral inflorescence branches. His treatment included two spe- cies, Thecophyllum poortmanii Andre and T. wittmackii Andre. In 1896 Mez reduced Thecophyllum to a section of Guzmania and included 14 species in the new section. Mez (1903a) reinstated Thecophyllum at the generic level and transferred the 14 species from his section to the renovated genus; he concomitantly emended Andre's original description to include species with enlarged primary bracts, aborted or highly reduced secondary inflorescence branches and free, appendaged petals. Mez's action resulted from the opportunity to study additional material of Guzmania ororiense Mez from Costa Rican collections of Pittier. During the following 13 years Mez (1903-1916), added a number of species to Thecophyllum, so that when he treated the Bromeliaceae in Das Pflanzenreich (1934-1935), Thecophyl- lum contained 45 species. Smith (1932, 1938) described four new species in Thecophyllum, and Suesenguth and Goeppinger (in Sues- senguth, 1942) added two species and a var- iety, bringing the total number of species in the genus to 51. Smith and Pittendrigh (1953) transferred all Thecophyllum species to Guzmania, Tillandsia or Vriesea. The two species considered by Andr^ when he origi- nally established the genus and 1 1 other spe- cies were relegated to Guzmania, two species were transferred to Tillandsia and the vast majority (38) of the taxa were transferred to Vriesea, where they were placed in section Xiphion. Those species transferred to Vriesea, as well as several species named as vrieseas since 1953, form a homogeneous group within sec- tion Xiphion. These thecophylloid vrieseas are more than 80% endemic to Costa Rica and Western Panama. Within this area they are largely confined to the moister parts of the lower montane, premontane and mon- tane regions. They are separated from the remainder of sections Xiphion by the same characteristics which distinguished Theco- phyllum sensu Mez reduced lateral inflores- cence branches and enlarged primary bracts. Moreover, the thecophylloids are distingui- shed from the majority of vrieseas by a uni- que stigmatic morphology that they share with a group of low to middle elevation members of the section Xiphion. The vrie- seas, like the thecophylloids, are highly diverse in Middle America. Habitat And Ecology From my extensive field work in Costa Rica. I interpret that the ecological dis- Editorial Committee for this Paper: Dr. Robert W. Read, Curator, Department of Botany, Smithsonian Institution, United States National Museum, Washington, D.C. 20560 Dr. Lyman B. Smith, Botanist Emeritus, Department of Botany, United States National Museum, Washington, D.C. 20560 No. I Middle American Vrieseas tribution of most thecophylloids exhibits a positive correlation with the life zone class- ification developed by Holdridge et al. (1971). The elevational distribution of the thecophylloids is confined to the area between 300 and 3200 m, while the vast majority of species are found between 900 and 2100 m (Fig. 1). Within this elevational range the taxa are confined to five life zones; premontane rain forest and wet forest, lower montane rain forest and wet forest, and montane rain forest. Most thecophylloid vrieseas are restricted to one or two life zones. When a species occupies more than two life zones, the exten- sion occurs into life zones that border on the principal one (s). Alternatively, species may occur in the center of an additional life zone, but this is invariably in conjunction with an isolated peak or ridge system that is regularly exposed to cloud and mist conditions. For terrestrial vegetation the general life zone of such exposed elevations may be a "wet zone" in the Holdridge system, but the additional atmospheric moisture present in mist and clouds effectively makes the epiphytic habi- tat equivalent to a "rain zone." A prime example of this situation is the crest of the Cedral or Cerros de Escazu, located south of San Jose, Costa Rica. This chain is located in the lower montane wet formation of Holdridge, but the crest and peaks of this chain receive almost daily clouds and mist. The area is an extremely rich collecting location for Vriesea neph- rolepis, a species common in the lower mon- tane wet forest life zone. However, near the crest of this range, and in particular crests of the individual peaks, V. ororiensis and V. williamsii, species typical of the montane and lower montane rain forest life zones, are common. These taxa are further stratified; V. williamsii is found only within a few yards of the peaks, whereas V. ororiensis is com- mon in a narrow band below the crest. This distribution apparently parallels subtle changes in the availability of atmospheric moisture. This assumption is reinforced by my observations from field and horticulture that indicate that V. williamsii exhibits less tolerance for desiccation than V. ororiensis. The extension of these two species into a drier life zone is undoubtedly facilitated by the regular availability of atmospheric moisture. Anomalies in the life zone system due to cloud and mist drift were predicted by Hol- dridge (1967), and Walter (1971) mentioned that epiphytes are good indicators of microc- limatic factors such as ambient humidity. Moerover, Pittendrigh (1948) related mois- ture requirements of bromeliads to their dis- tribution in Trinidad. The phenomenon was discussed for bromeliads in particular by Gilmartin (1973) who concludes that knowl- edge of bromeliad distribution in conjunc- tion with a physiognomic-ecological classifi- cation (e.g. the Holdridge Life Zone System) might be effectively used to indicate vegeta- tional types. I agree with Gilmartin's find- ings, but, I feel that use of epiphyte distribu- tion might be better described as a 'Tme tuning" of the life zone system. Knowledge of epiphyte distribution in conjunction with the life zone system allows one to infer condi- tions of atmospheric moisture or seasonal variation in precipitation that are not re- flected in general vegetational structure. As previously mentioned the thecophyl- loids are highly endemic to Costa Rica. All species found in Central America are also found in Costa Rica. Three species native to Costa Rica extend northward as far as Hon- duras, and one of these extends into south- ern Mexico. Vriesea pedicellata is common in the Cordillera Central of Nicaragua and has been collected as far north as Montana Uyuca in Honduras. Vriesea nephrolepis extends into Guatemala and V. hygrome- trica has been collected at Vista Hermosa in the state of Oaxaca, Mexico. The southeast- ern extension of thecophylloids into Panama Tulane Studies in Zoology and Botany Vol. 24 SPECIES attenuata balanophora bracteosa capitata comata diffusa greenbergii hainesiorum fiygrometrica kathyae latissima leptopoda leucophylla luis-gomezii lyman-smithii nepfirolepis notata ororiensis pedicellata picta singuliflora stenophiylla triflora umbrosa uxoris vietoris viridis williamsii 600 1200 ELEVATION IN METERS 1800 2400 3000 3600 Fig. 1. Elevational ranges of thecophylloid vriesea species in Middle America. No.l Middle American Vrieseas is to be expected considering the phytogeo- graphic continuum provided by the Cordil- lera de Talamanca which extends into cen- tral Panama. Several species, Vriesea lep- topoda, V. leucophylla, V.ororiensis and V. picta, are now known from northwestern Panama and V. capitata has been collected as far southeast as Cerro Jefe. Judged by the available data those collections were made at elevations and in life zones similar to those in which the species occur in Costa Rica. Morphology Habit. The thecophylloid vrieseas are primarily epiphytic, although many s'pecies will grow terrestrially. The majority of spe- cies are acaulescent with leaves densely rosu- late but, with a few species becoming extemely long caulescent. There apparently are taxonomic as well as ecological cor- relations associated with caulescence. Vrie- sea williamsii and a few related species, the only thecophylloids that regularly exhibit extreme caulescence, have stems ranging from ca 6 cm to over 1 m long. Extreme caulescence is frequently encountered in ter- restrial individuals. Vestiture. The single most characteristic feature of the Bromeliaceae is the almost universal presence of absorbing trichomes on leaf surfaces. These structures are want- ing in only Navia lopezii L.B. Smith, where their absence apparently is a derived condi- tion resulting from the failure of initial cells to subdivide (Robinson, 1969). The general structure of the tillandsioid tri- chome has been described by Tomlinson (1969) and Benzing (1976) and illustrated with great clarity by Ehler (1977). The lower portion of the trichome consists of three cell types, 1) two or more small basal foot cells, 2) two to three stalk cells and 3) a large distal dome cell. This stem-like structure supports a shield one cell layer thick that consists of four central cells, immediately distal to the dome cell and peripheral concentric series of ring cells. Each series contains twice as many cells as the adjacent inner ring cell series. The of the outermost series, or wing, frequently are elongate and distinct in appearance from the subtending ring cells. The wing cell series usually contains more than twice the number of cells in the preceding ring cell series and the cells often are allomorphic, producing asymmetric, or in extreme cases, "tailed" shields. Unlike shield cells, which are dead at maturity, cells of the stalk are living. The distal dome cell has a dense protoplast, an enlarged nucleus, an elaborated plasmalemma and numerous mitochondria, modifications that Dolzmann (1964) believed to be in- volved with the absorptive function of these trichomes. In addition the trichomes have a specialized pattern of cutinization which serves to direct water and solutes to the dome cell and ultimately to the leaf interior (Ehler, 1977). Moreover, this cuticular pattern coupled with the hygroscopic changes in orientation of the shield cells enables the trichomes to function as "one-way valves" in water and solute uptake. This mechanism was reviewed by Benzing (1976). Within the family, the structure of these absorbing trichomes is variable. An increase in complexity is apparently linked to adap- tive radiation into the epiphytic habitat. Moreover, this increase in complexity has apparently occurred along two separate lines. Such a view is supported by the fact that scale structure of epiphytic taxa in the Bromelioideae is strikingly different from the structure of scales found in the Till- andsioideae (Tomlinson, 1969; Benzing, 1976). The scales of the former subfamily, like those of the Tillandsioideae, have a well developed shield, but they lack the highly organized arrangement of central disc and peripheral ring cells found in the latter sub- family (Fig. 2). Variability in wing structure is frequently encountered within the blade of a single leaf Tulane Studies in Zoology and Botany Vol. 24 or bract. The shields in the central portion of the leaf are more or less symmetrical but near the leaf margin the shield becomes pro- gressively more asymmetrical. Moreover, scales of the adaxial and abaxial surfaces are frequently differentiated from each other by size and/ or density, and in the extreme case of Vriesea capitata, the trichomes from opposing leaf surface display radically dif- ferent wing morphologies (Figs. 3 & 4). The most striking examples of variation in tri- chome morphology in the thecophylloid Fig. 2. Arrangement of cells in the shield of absorptive trichomes; a) Pitcairioideae and Bromelioideae, and b) Tillandsioideae. vrieseas are found between trichomes from sheath and blade areas of the same leaf (cf. Figs. 3, 4, 5, 7, 8, 9, 10). The sheath area trichomes from different species tend to be similar morphologically (Figs. 5, 7, 8). This similarity becomes even more striking when one considers that the blade trichomes of the same species show distinct differences (Figs. 3, 4, 9, & 10). Interspecific similarity in sheath trichomes and frequent dissimilarity between those from blade regions may be due to convergence in function and struc- ture. Since these taxa are tank epiphytes, the area of greatest water and nutrient availabil- ity is within the tightly overlapping leaf sheaths thus trichomes of the sheath area will play a potentially important role in absorp- tion. Involvement in this essential process may have limited the toleration for variabil- ity in structure and resulted in the mainte- nance of similar trichomes in divergent taxa. The taxa surveyed in this study have a common shield morphology of four central cells and one series of eight ring cells; how- ever, the number of wing cells can be either 16 or 32. As already mentioned, great varia- tion in trichome morphology is encountered between the sheath and blade area of the same leaf, as well as between the abaxial and adaxial blade surfaces. This emphasizes the need to utilize trichomes from homologous areas in any comparative study. In addition to the ubiquitous foliar tri- chomes, the species under consideration also display other epidermal ornamentation, which is diagnostic of some taxa. The axis of the scape and inflorescence of Vriesea tri- flora is covered with densely packed papillae of characteristic appearance (Fig. 6). The rachis and scape of V. diffusa, a closely related species, also have these papillae, although they are not as well developed. While no other thecophylloids have these structures, the floral bracts of many com- monly have small wart-like excrescences, as well as pellucid, castaneous margined, punc- No.l Middle American Vrieseas Figs. 3-6, 3. Foliar trichome from the abaxial leaf-blade surface of Vriesea capitata. X ca. 375 (Utley & Utley 4477). 4. Foliar trichome from the adaxial leaf-blade surface of Vriesea capitata, X ca. 190 (Utley & Utley 4477). 5. Foliar trichome from the leaf-sheath area of Vriesea triflora, X ca. 300 (Utlev & Utley 4477). 6. Papillae on the inflorescence of Vriesea triflora. X c&. 85 (Utley & Utley 5193). Tulane Studies in Zoology and Botany Vol. 24 tulae and minute pits; the punctulae appar- ently represent poorly developed or abortive stomata and the pits are associated with absorptive trichomes. Leaves. The leaves of thecophylloid vrie- seas are typically densely rosulate on a much shortened stem or occasionally spirally arranged on an elongated caudex. The cau- lescent taxa rarely retain leaves over the entire length of the -stem; the older regions are usually denuded or covered with the remains of leaf bases, while functional leaves are found at the distal portion of the stem. The leaves consist of two morphological regions, the sheath and blade areas, which can usually be delimited fairly accurately, but occasionally the zones merge so gradu- ally that they can only be approximately defined. The sheath comprises that portion of the leaf delimited by the bounds of an open "S" curve; in addition there are frequently char- acteristic trichome morphologies and color or marking patterns, which change at or near the sheath-blade interface. As previously mentioned, the trichomes in the sheath area tend to be more highly developed and are very densely packed, frequently forming a mosaic pattern of shields. The abaxial sur- face or occasionally both surfaces may be densely castaneous, and in the distal portion of the sheaths, the foliar markings character- istic of many thecophylloids become evident. The leaf blade is that portion of the leaf distal to the sheath. Shapes encountered range from ligulate to elongate-triangular, the entire range of shapes occasionally being found within the rosette of a single in- dividual. The blades are frequently suffused with various shades of red, or maroon, a condition most evident when a plant is about to produce an inflorescence. However, plants in exposed locations are frequently colored throughout the year. In addition to general suffusion, many thecophylloids have wavy transverse lines or straight longitudinal stripes of pink, red ,purple or maroon. The wavy transverse bands are frequently dis- cernible in totally green leaves as wavy darker green lines. Leaves of taxa under consideration were sectioned and mounted using the techniques of Robinson (1969). The following gener- alized description of leaf structure is based on these slides. The cells of the adaxial epidermis, which frequently contain spherical, spinulose silica bodies, are comparatively flat in crosssection and the anticlinal walls are thickened and sinuous. The hypodermis is differentiated into an outer sclerotic layer with thickened walls and an inner layer of thinner walled "water storage" cells with anticlinally ex- tended, frequently plicate, walls. Immediately beneath the hypodermis is the chlorenchyma, which consists of two cell forms: I) a com- pact area of isodiametric or slightly elongate cells which are most abundant over the inter- costal aerating canals, extending down the sides of the vascular bundles or up the adax- ial bundle buttresses and at times completely overtopping the vascular bundles, and 2) stellately branched cells which occur as a ramifying network within the aerating can- als. The vascular bundles are located within strongly sclerified bundle sheaths, which form 'Tigure-8's" around the vascular ele- ments of the principal veins. While the cen- tral bar of the "8" is variable in degree of development it often is prominent. The adaxial surfaces of the bundle sheaths are typically strongly buttressed, with the distal portion of the longer buttresses coming within one cell layer of the sclerotic hypo- dermis; the abaxial surface is rarely but- tressed to the same degree. Below the vascu- lar bundles is the abaxial "water storage" hypodermis which frequently extends to the vascular bundles and also beneath the aerat- ing canals where it is interrupted by the sub- stomatal chambers. Beneath the abaxial water storage hypodermis is an outer layer of No.l Middle American Vrieseas sclerotic cells with very much thickened anticlinal and internal periclinal walls. There are taxa that are sufficiently unique in one or more anatomical features to be recognized on the basis of their leaf anatomy alone (e.g. V. bracteosa and V. capitata), however the sample size involved in this sur- vey is not adequate to support any broad generalizations regarding the use of anat- omical characters in the classification of the Figs. 7-10. Foliar trichomes of vrieseas. 7. From leaf-sheath area of Vriesea tnflora, \c&.215(Utley & Utley5193). 8. From the leaf-sheath area of Vriesea diffusa. X ca. 275 (Utley & Uiley 1857 c). 9. From the abaxial leaf-blade surface of Vriesea triflora. Xca. 275 (Utley & Utley 5193) 10. From the abaxial leaf-blade surface of Vriesea diffusa. X ca. 275 (Utley & Utley 1857 c). 10 Tulane Studies in Zoology and Botany Vol. 24 thecophylloid vrieseas. There is apparent variation in the amount of hypodermis, the amount and arrangement of sclerenchyma and the arrangement of chlorenchyma, indi- cating that further study of leaf anatomy is warranted. Inflorescences. The basic thecophylloid vriesea inflorescence consists of a scape sup- porting a once-compound inflorescence var- ying in length from one or a few centimeters (subcapitate) to several decimeters. The scape is usually erect but is occasionally curved laterally or even pendent. It is pro- vided with a series of spirally arranged scape bracts which are transitional in form be- tween leaves and the primary bracts of the inflorescence. The bracts consist of a sheath and blade area, which, as in the case of leaves, are occasionally difficult to delimit precisely. The length of these bracts relative to the internodes, and their color and mark- ing, frequently are of use in identifying spe- cies. The primary bracts, which subtend the lateral branches, are similar in shape and morphology to the upper scape bracts, but they are often more intensely colored and are usually spreading or even recurved in con- trast to the erect posture of the scape bracts. A third series of bracts, the floral bracts, subtends the flowers. The morphology of these bracts has been used to separate species or species groups. The condensation of lateral branches and concomitant enlargment of primary bracts influenced Andre (1889) to establish The- cophyllum and, although the genus is no longer recognized, this basic inflorescence structure is fairly distinctive within Vriesea. The inflorescence consists of a panicle in which the secondary branch system has become reduced to the point that the lateral branches are sessile or short pedunculate in the axils of enlarged primary bracts. The flowers are alternately or suboppositely spi- cate, racemose on a short axis, collateral or subfasciculate. The simplest of the inflor- escences encountered is the single flowered spike of Vriesea singuliflora, where the com- pound nature of the inflorescence is revealed only by the presence of a floral bract and a primary bract subtending each flower. In contrast the well-developed lateral spikes or racemes of V. greenbergii contain three to five flowers (Fig. 19). In several species, including V. ororiensis, the lateral fascicles regularily contain two flowers, with varia- tion largely confined to differences in pedun- cle and pedicel length (Fig. 1 1). The general morphological trends exhi- bited by the thecophylloid vrieseas are con- densation or abortion of the peduncles, pedicels or floral rachises and a reduction in the number of flowers per lateral branch. The diversity of inflorescence forms dis- played by this group can be derived through a reduction series from the laxly flowered panicle commonly encountered in Vriesea. Calyx. The calyx is imbricate in bud and closes around the developing capsule after anthesis. Because the sepals are often thickly coriaceous, the calyx-envelope potentially serves as a barrier against predation in early stages of fruit development. The durability of the sepals has resulted in these structures being utilized extensively in traditional spe- cies delimitation. The size, shape and texture Fig. 1 1. Lateral inflorescence branch and flowers of Vriesea ororiensis showing the two colater- al flowers and the subtending primary bract. No.l Middle American Vrieseas 11 of the sepals are useful but the general range in the former two characters and the qualita- tive nature of the latter make an "a poste- riori " application imperative. Corolla. The three free petals of the corolla are imbricate in bud, inserted alter- nately to the sepals and bear two scale-like appendages near the base. At anthesis the corolla may assume a wide variety of shapes ranging from campanulate to more or less tubular to strongly zygomorphic (cf. Figs. 11-14). The petals vary in size from about one cm to over five cm in length and from translucent white or greenish-white to char- treuse or yellow-green. The corollas of this group of vrieseas are highly variable; many species that are very similar or almost identi- cal vegetatively or in fruit, have vastly differ- ing floral structures and phenologies. Androecium. The androecium is charac- teristically asymmetric. The filaments are displaced so that the anthers, which are usu- ally connivent by their lateral margins, form a hood over the gynoecium (Fig. 12). Occa- sionally the androecial hood is represented by two pairs of three anthers, or rarely the anthers are free but the filaments are dis- placed so that the anthers are dorsal to the stigma. Gynoecium. The ovary is inserted on the receptacle in such a way that the petal appendages contact the gynoecium at the base of the style, loosely partitioning the intrafloral chamber into two compartments. The style is usually bent in a dorsal direction at the style -ovary junction and then arches distally until, near the stigma, it curves ven- trally, orienting the stigma surface away form the overarching androecial hood. The degree of proximal torsion varies from almost a right angle to practically none and the ventral torsion of the stigmatic surface varies but is usually present to some degree (Figs. 12& 13). As previously mentioned the thecophyl- loids share a unique stigmatic structure with a few other species in section Xiphion. The stigma presents the appearance of three cir- cular cups deposited symmetrically around a common axis (Fig. 15c). The remainder of the vrieseas have stigmas that are somewhat flattened and expanded distally. Deriving the thecophylloid stigma from the latter form through progressive flattening, expan sion and inrolling of the lateral margins, fol- lowed by, or concomitant with, an axial tor- sion of the tubular stigma branches is not conceptually difficuh. The distribution of this stigmatic type and its consequent signifi- cance in classification is not entirely known; the results of a preliminary survey and some tentative conclusions are presented in the section dealing with intergeneric relation- ships. Fruit. The fruit is a tough, septicidal cap- sule, which varies in shape from fusiform to strongly obovate, castaneous to olive-green in color and from about 1 .5-5.5 cm long. At maturity the fruit splits basipetally and the chartaceous septa break down or tear free from the outer wall releasing the plumose appendaged seeds, which are wind dispersed. Seed. The seeds, which are fairly uniform within and among the species, have a plu- mose appendage derived from the integu- ments. The outer integument splits longitud- 12 Fig. 12. Flower of Vriesea nephrolepis. 12 Tulane Studies in Zoology and Botany Vol. 24 13 Fig. 13. Flower of Vriesea williamsii. inally into hair-like segments that are at- tached only at the proximal end of the seed, thus forming a parachute-Uke coma. The inner integument also becomes longitudi- nally divided, but remains attached along the length of the seed coat. Seed characteristics have not yet proven of value within the the- cophylloids. The development and taxo- nomic significance of seeds in Bromeliaceae is reviewed by Downs (1974) and Szidat (1922). REPRODUCTIVE BIOLOGY The reproductive biology of the Bromelia- ceae has not been studied to any great extent. Mc Williams (1974) presented a survey of recent work and a list of putative breeding systems for 14 taxa. Brewbaker and Gorrez (1967) elucidated the genetics of self -incomp- atibility in the cultivated pineapple. The state of our knowledge concerning pollen vectors is only slightly better. There have been reports of bat, hummingbird, bee, fly and wind pollination (McWilliams, 1974). With respect to the thecophylloid vriesea species there are only two reports. Porsch (1932) considered Thecophyllum to be bat polli- nated and Salas (1973) inferred bat visita- tions to a species that he identified as Theco- phyllum irazuense. The thecophylloid vrieseas present a variety of phenologies: 1) day-blooming taxa, 2) crepuscular taxa, and 3) night-blooming taxa. The day-blooming taxa have greenish or greenish-yellow flowers and the primary bracts are usually red to pink, occasionally maroon or rarely brown at anthesis. Mem- bers of this group can be further subdivided based on the orientation of the inflorescence at anthesis. In V. ororiensis the inflorescence is erect and in V. leucophylla, V. hainesio- rum and V. uxoris inflorescence posture is downwardly slanting to strongly pendent. This entire group may be hummingbird pol- linated and I have made field observations for most of the species. In the case of V. ororiensis extensive observations and photo- graphs are available. Vriesea ororiensis usu- ally occurs in large local populafions and flowers during the dry season. The exact time of flowering varies among populations, but within a population flowering is syn- chronous. Within an inflorescence flowering is sequential, beginning basally and con- tinuing acropetally, with a few flowers at a time (one to four) opening over a period of several days to more than a week. The petals begin to expand prior to sunrise, sometimes as early as the preceding evening, and about dawn the corolla has assumed the form of a Fig. 14. Flower of Vriesea hygrometrica. No.l Middle American Vrieseas 13 narrow tube, with a distal portion of each petal flaring slightly. The androecial hood, which has been in tubular configuration around the style and stigma, separates ven- trally and assumes a dorsal hood config- uration over the ventrally facing stigma (Fig. 11). Pollen is shed in clingy masses about dawn or shortly thereafter. Stigma receptiv- ity, as measured by the production of viscid drops, occurs concomitantly with pollen release or as much as an hour later. The flowers last a single day and have usually become flaccid by the second morning, or on rare occasions they persist through the second day but become senescent by the third morning. Hummingbirds begin to work the opening flowers before sunrise and continue through late afternoon; they hover or occasionally rest lightly on a lower prim- ary bract while visiting the open flowers and then depart to visit other inflorescences in the vicinity (Fig. 16). Because of the paucity of open flowers on an inflorescence, self pol- hnation is kept to a minimum and the gener- ally high density of individual plants in- creases pollinator fidelity and concomitantly outcrossing (Baker, 1961). Vriesea leuco- phylla is also visited by hummingbirds, and ahhough extensive observations are not available the phenology and population den- sity are similar to V. ororiensis and the same generalizations may hold true. Vriesea hainesiorum is similar to V. leucophylla but the former species is consistently smaller and has fewer flowers per inflorescence. Although no potential pollinators have been observed visiting V. hainesiorum, the general form of the inflorescence, with red scape bracts and primary bracts and diurnal flowers, argues for ornithophilous pollination. Vriesea uxo- ris also possesses a syndrome of characters that suggest hummingbird pollination. Hum- mingbirds have been observed hovering near the species but no actual visitations have been recorded. Only two taxa, V. balanophora and V. vir- idis, are known to be crepuscular. Both spe- cies have white flowers and maroon sepals, whereas the primary bracts are green with prominent wavy maroon transverse bands or occasionally entirely maroon in V. viridis. This syndrome agrees with that proposed by Baker ( 196 1 ) for plants adapted to crepuscu- lar, ornithohilous pollination. Phenological observations are available only for V. viridis. Anthesis begins in the early afternoon and is completed by about 5:00 p.m. As in V. oro- riensis the direction of flower maturation is acropetal and only a few flowers are open on any given day; moreover, this taxon also tends to occur in extensive local populations. On the two occasions observations were made, hummingbirds visited the flowers from about 3:00 p.m. until light was insuffi- cient for further observations; even at this point, flight noise was audible in the vicinity of the plants. Other crepscular vectors possi- bly visit this taxon, however the flowers are odorless and hummingbirds work the in- florescences diligently, indicating that the Fig. 15. Diagrammatic representation of stigma forms. 14 Tulane Studies in Zoology and Botany Vol. 24 flowers are probably primarily omithophilous. In contrast to the diurnal and crepuscular taxa which seem to be groups of more or less similar species, the noctui-nal taxa are sub- divisible into two distinct groups based on inflorescence structure. The first of these groups consists of species with capitate to elongate inflorescences, with collaterally bi- flowered lateral branches and flowers sessile or subsessile in the axils of the primary bracts. Corolla shape varies from open cyl- indric in V. stenophylla to strongly zygo- morphic in V. williamsii and V. nephrolepis. In the latter case two of the petals are fre- quenUy associated into a dorsal hood and the third petal is positioned as a ventral, labellum-like structure (Fig. 12 & 14). Based on observations of living flowers, Porsch (1932) suggested the possibility of bat pollination in an unidentified species of night-blooming Costa Rican Thecophyllum; he likely observed a member of this group. Salas (1973) studied a species of the pan- iculate subgrouping with strongly zygomor- phic flowers. The determination of the spe- cies as Thecophyllum irazuense was in error however, because this name is a synonym of the day-flowering, omithophilous Vriesea ororiensis. Unfortunately, voucher specimens were not deposited, but the extensive floral description and the site location provided by Salas leave little doubt that the taxon he studied was Vriesea nephrolepis (Thecophyl- lum werckleanum). Salas observed bats fly- ing among the flowering plants from 6:30 p.m. until 1 1:00 p.m. and inferred bat visita- tions to the bromeliad flowers by trapping the animals and examining fur for presence and type of pollen. He reported finding cop- ious amounts of "Thecophyllum" poWen on the tops of the bats' heads and the sides of their faces. He also noted that the flowers began to open about 5:00 p.m. and had a musky odor. My observations of this species generally bear out Salas' reports on the time of anthesisand smell of the flowers, however many flowers in the population I observed began opening as early as 3:30 p.m. and several were fully open by 5:00 p.m. In addi- tion, hummingbirds were photographed vis- iting open flowers in a manner which was as conducive to pollination as visitations by bats. The second major group of nocturnal taxa consists of species with well developed lateral branches with obvious peduncles, pedicels or rachises and frequently more than two flow- ers per branch. Corolla shape varies from campanulate in V. pedicellata to strongly zygomorphic in V. umbrosa. Phillip DeVries (pers. comm.) has observed bats visiting what is probably V. latissima in the Monte Verde forest perserve in Costa Rica. I have observed that V. umbrosa has a decidedly sweet, spicy odor during the early evening. This odor, in conjunction with noc- turnal anthesis, argues strongly for pollin- ation by a night flying insect but confirm- atory observations are lacking. In summary, the thecophylloid vrieseas present a wide range of floral and inflor- escence types as well as divergent phenolo- gies. A variety of potential pollinators has been observed or inferred on the basis of presumptive evidence. INTRAGENERIC RELATIONSHIPS The genus Vriesea is distinguished from the remainder of the Tillandsioideae on the basis of distichous inflorescences and free ap- pendaged petals. It is subdivided into two subgenera and two sections on the basis of seed and floral characters. The subgenus Alcantarea is characterized by seeds that have the apical appendage enlarged and divided to form a short coma in addition to the usual basal coma and ephemeral petals that become flaccid soon after anthesis. In contrast, subgenus Vriesea has a single basal plumose appendage and comparatively dur- able petals. The subgenus Alcantarea contains only seven species and has not been further subdi- No.l Middle American Vrieseas 15 vided. Subgenus Vriesea, however, has over 200 species and contains two sections which are separated from each other on the basis of androecial structure. In section Vriesea the stamens are exserted from the corolla, whe- reas in Xiphion they are included. This div- ision is correlated with several other charac- ters. Section Vriesea contains species that frequently have highly colored bracts which are red, yellow or orange, but section Xiphion contains species that generally have dull, green or brown colored bracts. Furth- ermore, section Vriesea has a center of diversity in southeastern Brazil whereas Xiphion has a center of diversity in northern South America and southern Central Amer- ica, with a secondary center of diversity in southeastern Brazil. Within section Xiphion the thecophylloids are most closely allied to a complex of low to middle elevation, secund-flowered species which are concentrated in Costa Rica, Panama and Brazil. However, a few species are distributed through northern Central 4 Fig. 16. Hummingbird (Panterpe insignis) visiting Vriesea ororiensis. America, the Antilles and northern South America (Smith, 1941). Most species of this group exhibit the three-cupped stigmatic structure typical of the thecophylloids (Fig. 15c). The only taxa examined that lacked this stigmatic structure were V. gigan- tea Gaud, and V. unilateralis (Baker) Mez from southern Brazil. Both of these species have a stigma form consisting of three flat- tened, suborbicular branches (Fig. 15b). Moreover, the secund-flowered vrieseas have nocturnal flowers which are essentially identical to the larger, campanulate, noc- turnal flowers of the thecophylloids. More data are needed but the similarity in floral morphology, including the unique stigmatic structure, tempt one to hypothesize that the thecophylloid vrieseas and (at least the majority of) the secund-flowered species of section Xiphion share a common origin. If the foregoing assumption is true then the ancestral group of the thecophylloids likely had night-blooming flowers. This is sug- gested by the predominance of night- flowering in the thecophylloids and their secund-flowered allies. If this is so, and if one accepts the likelihood that the montane spe- cies endemic to Costa Rica envolved in situ, then it follows that the ornithophilous taxa have undergone adaptation to avian pollina- tion independently of other ornithophilous vrieseas. Even more interesting is the corol- lary that these diurnal, ornithophilous spe- cies are derived from a nocturnal, potentially chiropterophilous, ancestor, a situation that is the converse of the usual evolutionary sequence (Grant and Grant, 1968; Skog, 1976). The thecophylloid vriesea species can be divided into three basic groups on the basis of inflorescence morphology. The first of these groups, the pedicellata complex, is characterized by well-developed secondary inflorescence branches with obvious pedun- cles, pedicels, or rachises and usually more than two flowers per lateral branch. The other groups, ororiensis and hygrometrica 16 Tulane Studies in Zoology and Botany Vol. 24 complexes, are characterized by collaterally biflowered lateral inflorescence branches with little, if any, peduncle, rachis or pedicel development. Four species groups can be recognized within the pedicellata complex on the basis of pedicle length, the size and texture of floral bracts and floral morphology. The first of these, the pedicellata species group, is characterized by membranous floral bracts which are shorter than or equal to the pedic- els, and campanulate flowers. The pedi- cellata species group is similar to the triflora group that is distinguished by subcoriaceous to coriaceous floral bracts, which exceed the pedicels, and by the papillose surfaces of the inflorescences. As in the pedicellata group, the flowers are campanulate. The third spe- cies group, the umbrosa group, is distin- guished by coriaceous floral bracts which exceed the pedicels, by the smooth surfaces of the inflorescences and by the bilaterally symmetrical corollas. The fourth species group in the pedicellata complex, the latis- sima species group, contains a single species, V. latissima. This species is most similar to the species in the umbrosa group, but is dis- tinguished by its larger floral bracts and flowers and much stouter inflorescence branches. In the second of the complexes, the ororien- sis complex, the floral bracts are shorter than or about equal to the sepals and the inflores- cence usually is elongate. This complex con- tains five species groups that are differen- tiated on the basis of floral bract length, inflorescence orientation, phenology and growth habit. The first of these groups, the ororiensis species group, is characterized by diurnal or nocturnal anthesis, floral bracts that are no more than one half as long as the sepals, and erect inflorescences. This group consists of three taxa which are very similar in the vegetative and fruiting states. How- ever, they are readily separable in flower on the basis of phenology, primary bract color and floral morphology. Members of this group display floral syndromes typical of adaptation to bat or hummingbird pollen vectors. The second group, the balanophora group, is characterized by floral bracts that are about as long as the sepals, crepuscular anthesis, maroon or purple sepals and ciner- ous pubescence on the abaxial leaf surfaces. This group is most similar to the leucophylla species group which is characterized by pendent inflorescences, diurnal anthesis and floral bracts that are about as long as the sepals. The species in this latter group are apparently adapted for diurnal humming- bird pollination. The fourth group consists of a single species, V. uxoris, which has a pendent inflorescence and diurnal flowers. This species differs in floral morphology, ecology and trichome type from the pendent- inflorescenced species in the leucophylla group and is best considered as distinct from these species. The final group in this com- plex, the stenophylla species group, consists of species with nocturnal anthesis, strongly zygomorphic corollas, a caulescent habit and tank-roots. Diurnal anthesis in conjunc- tion with other characteristics typical of ornithophily (Faegri and van de Pijl, 197 1) is found only in the ororiensis complex. Signif- icantly this complex contains those species that have been the most successful in adapt- ing to the montane environment. Moreover, the presence of crepuscular anthesis and noc- turnal anthesis in conjunction with a chirop- terophilous syndrome (Faegri and van de Pijl, 1971) indicates that this complex is potentially adapted to a diversity of pollina- tors. Neither the pedicellata complex nor the hygrometrica complex exhibits a similar pol- linator diversity and these groups are poorly represented at higher elevations (i.e. above 2,000 m). Perhaps the pollinator diversity of the ororiensis complex has contributed to their success in the higher montane regions. The third complex, the hygrometrica com- plex, contains taxa with floral bracts that are usually much longer than the sepals and inflorescences, which are frequently subcap- No.: Middle American Vrieseas 17 itate or short, dense cylindric. This complex consists of three species groups. Two of these, the comata group and the hygrome- trica group, are closely atllied and are distin- guished from each other on the basis of leaf size and markings, density of the inflores- cence and primary bract color. The comata group contains V. comata and V. attenuata. The latter species has floral bracts that are regularly shorter than or equal to the sepals. The two taxa are virtually indistinguishable in the vegetative state but fertile material is readily indentifiable. In light of its overall similarity to V. comata, V. attenuata is best considered part of this complex despite the aberrant floral bract length. I also place V. singuliflora in the comata group. The floral bracts of this species are also shorter than the sepals. However, its castaneous leaf sheaths, unmarked green leaves, apparently white, nocturnal flowers and occurrence at rela- tively low elevations (300 m), suggest a rela- tionship with the comata group. In contrast to the usually green leaves of the comata species group, the leaves of the hygrometrica group are frequently suffused with purple to maroon and often have longitudinal or transverse lines of red, purple or maroon. Moreover, the inflorescences of this group elongate post anthesis, a condition which is not encountered in the comata species group. The third group in the complex, the bracteosa species group, is characterized by its dense, capitate inflorescences, green unmarked leaves in an ascending rosette and by the presence of a well developed layer of extrafascicular sclerenchyma in the adaxial hypodermis. The arrangement of the Middle American thecophylloid vrieseas into species groups is summarized below (Table 1). In summary, the thecophylloid vrieseas may be closely allied to a group of secund- flowered species in section Xiphion. This alliance is supported by floral morphology, phenology and a common stigma structure. The thecophylloids are divisible into group- ings on the basis of the structure of the lateral Table i. Species groups of Middle American thecophylloid vrieseas. I. pedicellata complex A. Pedicellata group B. Triflora group C. Umbrosa group D. Latissima group II. ORORIENSIS COMPLEX E. Ororiensis group F. Balanophora group G. Leucophylla group H. Uxoris group I. Stenophylla group III. HYGROMETRICA COMPLEX J. Comata group K. Hygrometrica group L. Bracteosa group V. leptopoda & V. pedicellata V. diffusa & V. triflora V. greenbergii & V. umbrosa V. latissima V. lyman-smithii, V. nephrolepis & V. ororiensis V. balanophora, V. luis-gomezii & V. viridis V. hainesiorum, V. leucophylla & V. vietoris V. uxoris V. picta, V. stenophylla & V. williamsii V. attenuata, V. comata & V. singuliflora V. hygrometrica, V. kathyae & V. notata V. bracteosa & V. capitate Tulane Studies in Zoology and Botany Vol. 24 inflorescence branches, inflorescence form and the relative length of floral bracts and sepals. In contrast to the general pattern of nocturnal anthesis found in Xiphion, several thecophylloids have developed diurnal anthesis coupled with an ornithophilous floral syndrome in addition to creuscular anthesis in conjunction with a syndrome adapted to pollination by a crepuscular vec- tor. Moreover, this variety of pollination syndromes is confined to a single group of species that have been highly successful in the middle and upper montane environ- ments. The success of these species at higher elevations likely is related to their adaptation to a variety of potential pollinators. TAXONOMIC TREATMENT Although the thecophylloid vrieseas have not been given nomenclaturally valid status as a group within section Xiphion, they and their allies form a distinct, probably phy- letically related alliance. However, further knowledge of the group and the remainder of Xiphion is an essential prerequisite to the erection of a formal intrasectional frame- work. The thecophylloids and their allies can be easily separated from the remainder of the Costa Rican vrieseas by their unique three- cupped stigmatic surfaces. Those species that were or would have been considered in the genus Thecophyllum by Mez (1934-1935) can generally be separated from their secund- flowered allies on the basis of general inflo- rescence structure combined with develop- ment and form of the lateral inflorescence branches and primary bracts. The following synoptic key summarizes the salient differ- ences between the thecophylloid vrieseas and their allies. 1. Inflorescences simple or compound; the lateral branches much exceeding the primary bracts, the floral bracts exceeding the sepals and the pedun- cles and rachises smooth, definitely not papillose thecophylloid allies 1. Inflorescences compound, or if ap- pearing simple than a series of two bracts subtending each flower indi- cates a compound inflorescence; primary bracts exceeding or about equaling the lateral branches; if the lateral branches exceed the primary bracts then the floral bracts are shorter than the sepals or the rachis and peduncles are densely papillose thecophylloid vrieseas The characters that are used to separate the species of thecophylloids have already been detailed, but a brief discussion of the mea- surement of floral parts is necessary before the following key to species can be used effectively. Floral parts especially the floral bracts, sepals and petals, shrink on drying, in some cases resulting in as much as 25% change in dimensions. Therefore floral material used in identifying species must be fresh, liquid preserved or rehyd rated by boil- ing the flowers. With the exception of floral parts the following key is based on dried material. ARTIFICIAL KEY TO THE SPECIES OF MIDDLE AMERICAN THECOPHYLLOID VRIESEAS 1. Proximal lateral branches 1 -flowered 2. Floral bracts longer than wide or the two dimensions about equal; sepals to 1 .2 cm long. 3. Floral bracts as long as or shorter than the pedicels V. pedicellata 3. Floral bracts longer than the pedicel V. diffusa No.l Middle American Vrieseas 19 2. Floral bracts wider than long, subrotund; sepals 1.5 cm long .... V. singuliflora 1. Proximal lateral branches bearing 2 or more flowers. 4. Lateral branches of the inflorescence highly developed, frequently more than 2- flowered; peduncles, rachises or pedicels elongated, usually slender. 5. Floral bracts membranaceous, shorter than or about equalling the pedicels. 6. Lateral branches 5- to 8-flowered; floral bracts 1.4-1.8 cm long; sepals 1.4-1.6 cm long; petals 2.5-2.7 cm long V. leptopoda 6. Lateral branches 2- to 5(6)-flowered; floral bracts 0.4-0.8 cm long; sepals 0.8-1 .2 cm long; petals 1 .8-2. 1 (2.4) cm long V. pedicellata 5. Floral bracts chartaceous to subcoriaceous, distinctly exceeding the pedicels. 7. Rachis of the inflorescence, peduncles or pedicels densely papillose. 8. Floral bracts 0.9- 1 .2 cm long; petals 1 .9-2.0 cm long; leaf blades with wavy transverse lines proximally V. triflora 8. Floral bracts 0.5-0.7 cm long; petals 1.5-1.6 cm long and leaf blades with longitudinal purphsh lines proximally V. diffusa 1 . Rachis of the inflorescence, peduncles and pedicels glabrous or with punctu- lae, definitely not papillose. 9. Leaf blades deeply and evenly suffused with maroon abaxially; floral bracts 2.0-3.0 cm long V. latissima 9. Leaf blades green, occasionally suffused with maroon distally, abaxial surface never evenly suffused with maroon; floral bracts 1.0-1.8 (2.2) cm long. 10. Floral bracts 1.0-1.3 cm long; petals 2.3-3.1 cm long V.greenbergii 10. Floral bracts 1.4-1.8 (2.2) cm long; petals 3.6-4.2 cm long V. umbrosa 4. Lateral branches of the inflorescence not well developed, typically collaterally 2-flowered or, if proximal branches 3-flowered, then the inflorescence capitate or pendent; peduncles, rachises and pedicels highly reduced, or when developed then short and stout. 1 1 . Floral bracts distinctly longer than the sepals. 12. Floral bracts at most 1.8 cm long; sepals 0.7-1.0 cm long; petals 1.7-1.9 cm long V. kathyae 12. Floral bracts greater than 2.0 cm in length; sepals greater than 1.3 cm in length; petals longer than 2.0 cm. 20 Tulane Studies in Zoology and Botany Vol. 24 13. Sepals 2.0 cm long or longer; leaves brittle, thick-coriaceous, in cross- section the bundle sheaths not buttressed and not extending into the adaxial hypodermis; highly developed layer of extrafascicular scleren- chyma present in the adaxial hypodermis. 14. Floral bracts 2.0-2.2 cm long, oblong to obovate, unkeeled or weakly keeled distally; petals 3.1-3.4 cm long V. capitata 14. Floral bracts 3.8-4.6 cm long, broadly elliptic or ovate to orbicular, strongly keeled; petals 3.8-4.3 cm long V. bracteosa 13. Sepals at most 2.0 cm long and usually much shorter; leaves quite flexible; bundle sheaths buttressed, these extending well into the adaxial hypodermis, extrafascicular sclerenchyma wanting. 15. Scape 16.0-26.0 cm long; leaves green, blades with fine longitudinal maroon lines proximally V. comata 15. Scape 29.0 cm long or longer, if shorter than the leaves then strongly suffused with maroon; leaves green or maroon frequently with wavy transverse lines. 16. Leaves and bracts green, frequently suffused with reddish-brown wavy transverse lines V. hygrometrica 16. Leaves and bracts frequently suffused with maroon or red, or if green then with longitudinal maroon or red lines V. notata 1 1 . Floral bracts shorter than or about equaling the sepals or if slightly longer than the sepals, the inflorescence pendent. 1 7. Inflorescence pendent; primary bracts brightly colored at anthesis; flowers diurnal. 18. Floral bracts at most Vi as long as the sepals; sepals 2.2-3 cm long; leaf blades ligulate or rarely broadly triangular; at least 3.5 cm wide; known from the montane rainforest life zone V. uxoris 18. Floral bracts distinctly more than Vi as long as thesepals and usually about equal to the sepals; sepals less than 2. 1 cm in length; leaf blades triangular or rarely subligulate, less than 3 cm wide; known from the premontane rainforest, wet forest and moist forest life zones. 19. Inflorescence with 7 or more lateral branches; sepals 1.5-1.9 cm long; found in the premontane rainforest and wet forest life zones. 20. Leaf blade and upper sheath area usually marked with con- spicuous green or dark maroon wavy transverse lines; floral bracts green or rarely weakly suffused with pink; sepals green to yellow-green; primary bracts frequently geniculate-reflexed; sheaths of lower primary bracts 2.6-3.8 cm wide or, if rarely No.l Middle American Vrieseas 21 narrower, then the leaves with obvious maroon transverse lines; most common in the premontane rainforest life zone V. leucophylla 20. Leaf blade and upper sheath area unmarked, never with conspic- uous green or maroon transverse lines; floral bracts deep pink to crimson; sepals deep red at anthesis; primary bracts subscalari- form not geniculate-reflexed; sheaths of lower primary bracts 2-2.4 cm wide; most common in premontane wet forest life zone V. vietoris 19. Inflorescence with 5 or fewer lateral branches; sepals 1.9-2.1 cm long; most common in the premontane moist forest life zone V. hainesiorum 1 7. Inflorescence erect; flowers diurnal, crepuscular or nocturnal; primary bracts green, brightly colored or dried brown at anthesis. 2 1 . Primary bracts pink to scarlet at anthesis; flowers diurnal, green to yellow %^^^^ V. ororiensis 2 1 . Primary bracts green, maroon or dried brown at anthesis; flowers crepus- cular or nocturnal or if rarely diurnal, then the primary bracts dried brown at anthesis. 22. Plants caulescent and producing tank-roots in the axils of the older leaves; leaves usually with longitudinal maroon lines at least proximally. 23. Inflorescence greater than 6.0 cm in length. 24. Floral bracts 1.6 cm long, lanceolate, occasionally wanting; longitudinal lines wanting V. picta 24. Floral bracts 1 .4 cm long or shorter ovate, oblong or cuniform, always present; longitudinal lines usually present V. williamsii 23. Inflorescence 6.0 cm long or less in length. 25. Floral bracts 0.7-0.9 cm long; maroon lines broad, color not confined to the veins; inflorescences lax V. stenophvlla 25. Floral bracts 1.1-1.4 cm long; maroon lines very narrow, color apparently confined to bundle sheaths of veins; infiorescence subcapitate to dense cylindric V. attenuata 22. Plants acaulescent or if caulescent, then tank-roots not produced; longitudinal colored lines infrequently produced. 22 Tulane Studies in Zoology and Botany Vol. 24 26. Sepals (at least some) entirely maroon; leaf blades cinereously pubescent abaxially. 27. Floral bracts 1 .5- 1 .7 cm long; petals 2.9 cm long; corolla tubular or campan- ulate at anthesis V. luis-gomezii 27. Floral bracts 1.2 cm long or less; petals to 2.5 cm long; corolla tubular or campanulate at anthesis 28. Inflorescence 3.0-6.0 (9.0) cm long, with 8 or fewer lateral branches; floral bracts usually exceeding one-half the length of the sepals at anthesis; corolla campanulate V. balanophora 28. Inflorescence 1 1.0-31.0 cm long, with considerably more than 10 lateral branches; usually less than the length of the sepals at anthesis floral bracts corolla tubular geniculate V. viridis 26. Sepals green, olive-green or suffused with maroon marginally but not entirely maroon; abaxial leaf surfaces not obviously pubescent. 29. Flowers green, diurnal, petals 2.6 cm long or less, corolla tubular at anthesis V. ororiensis 29. Flower white or greenish white, nocturnal, petals 3.1-5.4 cm long. 30. Corollas strongly zygomorphic at anthesis; style 2.4-3.7 cm long; sepals elliptic to obovate; mature capsules to 4.3 cm long V. nephrolepis 30. Corollas subtubular at anthesis; style 1 .7-2. 1 cm long; sepals frequently suborbicu- lar; mature capsules 4.9-5.3 cm long V. lyman-smithii 1. Vriesea leptopoda L. B. Smith & Pitt. J. rounded and acuminate; sheaths 19.0-27.0 Wash. Acad. Sci. 43: 403. 1953. (310) cm long and 7.0-12.0 cm wide, pale ^ , „ , »# OL green drying to light brown with a coating of Thecophyllum rubrum Mez &^. ■^ r .^ »A, 1 1 ■ »# n 11 Tf u n ■ cmereous tnchomes, frequently with a cas- Werckle m Mez, Bull. Herb. Boiss. , ^ , ^ .,• • II A- 878 1904 taneous band near the base, elhptic to ^, . ' ' ' ' ,oc-T weakly obovate. SCAPE erect or curved- Vriesea rubra Beer, 1857. ^^ « ^^ « , . « ^ , -, TYPE: COSTA RICA: Werckle, ^'^''^ ^'^f'^ ^"^ '«"f ""A , '"" '1 diameter; bracts erect, 3.5-4.5 cm long and (1.9) 2.2-3.2 (3.7) cm wide, green the middle and upper soon dryingbrown, exceeding or Epiphytic or terrestrial, acaulescent. about equaling the internodes, ovate to ellip- Bromel. Costaric. 100 (HOLOTYPE: B!; ISOTYPE: GH). LEAVES in a spreading rosette, 38.0-55.0 tic, apex acute to acuminate. INFLORES- (62.0) cm long; 6/fl^e5 14.0-3 1.0 cm long and CENCE erect, glabrous, (23.5) 35.0-55.0 5.0-8.0 cm wide, green or green suffused with (67.0) cm long and 8.0-12.0 cm in diameter; purple, especially on the lower surface or pr/mflrv^rac/5 3.5^.2cmlongand(1.6)2.0- near the apices, ligulate, apex acute to 2.8 (3.7) cm wide, dried brown, ovate to No.l Middle American Vrieseas 23 elliptic, apex acute to acuminate, LATERAL BRANCHES 3.5-6.7 cm long, 5-to 8-flow- ered, peduncle 0.4-0.7 (1.1) cm long, rachis 3. 1-4.7 (5.5) cm long. FLOWERS 1.7-2.0 cm pedicellate;y7orfl/ bracts 1 .4-1 .8 cm long and 0.5-0.9 (1.2) cm wide, dried brown, surface nerved, frequently short tuberculate, keel usually present as a narrow longitudinal thickening, occasionally confined to the dis- tal portion or absent, elliptic to ovate, navic- ular, apex acute or torn and appearing bifid; sepals 1.4-1.6 cm long and 0.8-1.1 cm wide, coriaceous, green or green suffused with maroon drying to brown or yellow-brown, elliptic to obovate, apex obtuse to rounded; petals 2.5-2 J cm long and 1.2-1.3 cm wide, white, elliptic; appendages 0.5-0.6 cm long and 0.2-0.3 cm wide; stamens in a hood con- figuration over style, anthers not connivent, filaments 1.4-1.7 cm long, anthers 0.5-0.6 cm long; ovary 0.5-0.6 cm long and 0.3-0.4 cm in diameter, style about 1.9 cm long, exserted beyond the staminal hood; capsules 2.2 cm long and about 0.6 cm in diameter, fusiform, castaneous. PHENOLOGY: Flowering specimens have been collected in March and May. DISTRIBUTION AND HABITAT: Costa Rica and northwestern Panam^ between 1600-2300 m in the premontane and lower montane rain forest life zones. DISCUSSION: Prior to my collections Vriesea leptopoda was only known from the type material. Since the inflorescence of the type specimen is either immature or aborted and only the inner rosette leaves are present, exact interpretation of this specimen was dif- ficult. However, its broad leaves and the nature of the pedicels and floral bracts, when taken together, are a unique combination of characters which eliminates other interpre- tations. Leaves of this taxon, which are green suffused with maroon abaxially, form a broad, spreading rosette. The inflorescence is erect, with well developed lateral branches. Floral bracts of this taxon are membranace- ous and about equal to or shorter than the slender, elongate pedicels. The white, cam- panulate, nocturnal flowers of V. leptopoda are characteristically downwardly secund at anthesis. In vegetative condition, V. leptopoda can be easily confused with V. latissima. The features that distinguish these taxa are pres- ented under V. latissima. Phenetically, V. leptopoda is most similar to V. pedicellata. In both taxa the flowers are slenderly pedi- cellate, the floral bracts are membranaceous and shorter than, or equal to, the pedicels and the flowers are white, campanulate and nocturnal. These taxa are readily separated by pedicel length, floral bract size, and sepal and petal size. SPECIMENS EXAMINED: COSTA RICA: CARTAGO PROVINCE: Aseri, on tree in cloud forest, 19 March 1966, L. B. Smith, etal. 15324. Region between Cascajal and ca 8 km northeast on CR Highway 216 or about 6 km slightly south of due east from Alto de La Palma, 1600-1800 m, 23 August 1975, Utley & Utley 2925 (DUKE). BOR- DER OF CARTAGO AND SAN JOSE PROVINCES: ca 17 km south of El Empalme on the Interamerican Highway, ca 2600m, 17 March 1975 (grown in hort., pressed 22 May 1975), Utley & Utley 2003 (DUKE). HEREDIA: Between the junction of Costa Rican highways 9 and 1 20 and 3 km south on Hwy. 9, 1900-2100 m, 1 1 May 1975, Utley and Utley 2486 (CR), 27 March 1976, Utley & Utley 4374 (NOLS), 3 May 1976, Utley & Utley 4695 (DUKE), secondary road north of CR Hwy. 1 13 connecting with Calle Gallito, saddle area between Cerro Chompipe and the southeast flank of Volcan Barba, 2000-2100 m, 8 December 1975, Utley & Utley 3581 (NOLS) & 3582 (DUKE). SAN JOSE: Cerros de Escazu (Cedral) approach to and summit of Cerro Daser, 2220-2318 m, 25 May 1975, Utley & Utley 2517 (DUKE). PANAMA: CHIRI- QUI PROVINCE: Bajo Chorro, 17 Febru- ary 1938, Davidson 307 (F). 24 Tulane Studies in Zoology and Botany Vol. 24 2. Vriesea pedicellata (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953 Fig. 17. Thecophyllum pedicellatum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 3: 136. 1903. TYPE; COSTA RICA: province not given: near Cartago, elevation 1200- 1600 m, Werckle in Inst. Phys.-Geogr. Costar. /<5/97 (HOLOTYPE B!, pho- tograph US!) Thecophyllum violascens Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 877. 1904. TYPE: COSTA RICA: province not given, near La Palma, Werckle Bromel. Cos tar ic. 43, 91. (LECTOTYPE: Werckle Bromel. Costaric. 43, B!, photograph US! SYNTYPE: Werckle Bromel. Cosraric. 9/,B!, photograph US!). Thecophyllum turbinatum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 1122. 1904. TYPE: COSTA RICA: location not given, Werckle Bromel. Costaric. 116 (HOLOTYPE: B!, photograph US!, ISOTYPE: GH). Vriesea violascens (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43:403 1953. Vriesea turbinata (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43:403.1953. Epiphytic or terrestrial, acaulescent. LEAVES in a spreading to subascending rosette, 19.0-47.0 (57.0) cm long; blades (7.0) 17.0-34.0 (41.0) cm long and (2.0) 2.4-5.3 cm wide, green usually suffused with purple or reddish-maroon abaxially or infrequently both surfaces, frequently with longitudinal purple or maroon lines, ligulate to subligu- late or broadly triangular, apex acute to acuminate or broadly acute to rounded and short acuminate; sheaths (7.0) 9.0-20.0 (22.0) cm long and 3.4-7.0 (8.4) cm wide, drying pale green with a coating of cinereous tri- chomes, usually suffused with longitudinal purple lines or purple to red distally, elliptic to ovate in outline. SCAPE erect to curved erect, (18.0) 24.0-50.0 (60.0) cm long and 0.2-0.5 (0.8) cm in diameter; bracts 3.3-9.0 (12.8) cm long and 1.3-3.9 cm wide, blade and sheath areas not clearly delimited, pale green, frequently suffused with longitudinal purple or maroon stripes, erect and clasping in the sheath area, divergent in distal portion of blade, sheath elliptic to ovate, blades tri- angular, apex acute to acuminate. INFLO- RESCENCE erect (6.0) 10.0-38.0 (54.0) cm long and (4.0) 6.0-1 1 .0 cm in diameter, cylin- dric, rachis green or suffused vwith purple; primary bracts (3.1) 3.6-8.0 (10.4) cm long and 1.5-3.5 cm wide, sheath and blade areas not clearly delimited, green frequently suf- fused with purple, drying brown at anthesis, sheaths ovate to elliptic or rarely obovate, blades triangular to elongatetriangular, apex acute to acuminate. LATERAL BRAN- CHES spreading to ascending (0.3) 0.9-2.3 (3.3) cm long, 2-5 (6) flowered, flowers race- mose, peduncles wanting to 0.3 (0.6) cm long, rachis 0.3-2.5 cm long. FLOWERS (0.6)0.8-1.9 (2.2) cm pedicellate;y7ora/^ram 0.4-0.8 cm long and 0.3-0.6 cm wide, drying pale brown at anthesis, membranous, gla- brous, finely nerved, even or slightly keeled, ovate to elliptic or rarely oblong, apex acute to obtuse or rounded; sepals O.S- 1 .2 cm long and 0.5-0.8 (1.0) cm wide, green or purple drying olive-green to castaneous, coriaceous, glabrous, elliptic to obovate, apex obtuse to rounded, frequently torn incised; pe/a/^ 1.8- 2. 1 (2.4) cm long and 0.9-1.1 ( 1 .4) cm wide, white occasionally lightly suffused with maroon marginally and apically, obovate; appendages 0.3 cm long and 0. 1 - 1 .2 cm wide; stamens loosely arranged over the gynoe- No.l Middle American Vrieseas 25 cium, anthers not connivent by their lateral margins, filaments 1.1-1.7 cm long, anthers 0.3-0.4 cm long; ovary 0.5 cm long and 0.25- 0.35 cm in diameter, style 0.7-0.9 (1.6) cm long; capsules 2.0-2.4 cm long and 0.4-0.7 cm in diameter, castaneous, fusiform. PH ENOLOG Y: Flowering specimens have been collected in December, and January but flowering Hkely occurs during other months as well. DISTRIBUTION AND HABITAT: Known from Costa Rica, Nicaragua and Honduras between 900-2100 m in the lower montane and premontane rain forest life zones and the lower montane and premontane wet forest life zones. DISCUSSION: V. pedicellata is a com- monly encountered, but not often collected taxon. Of the thecophylloid vrieseas, it is the most variable species, exhibiting much greater variability than either V.nephrolepis Figs. 17. Vriesea pedicellata inflorescence or V. ororiensis. Variation has been ob- served in size and general robustness, leaf coloration and markings, the degree of de- velopment of the lateral branches of the inflorescence and the number of flowers. Within a single population, the inflorescence can vary from 6-10 cm in length to over 50 cm. This is correlated with both the size of the flowering individuals and the degree of development of the lateral branches. Smaller plants tend to have shorter inflorescences with fewer flowered lateral branches, while larger individuals have extremely robust inflorescences with as many as five or six flowers per lateral branch. Variation in robustness is accompanied by, but appar- ently not correlated with, a great variation in leaf color and markings. Leaves of V. pedi- cellata range from green to maroon or deep red -purple and may be unlined, weakly lined or strongly lined throughout the leaf blade. Although V. pedicellata is a highly variable taxon, individuals are united by a series of common characters, including ligulate leaves and membranaceous floral bracts that are generally shorter than the pedicels. The white, campanulate flowers of this taxon are nocturnal. The variation encompassed by V. pedicel- lata was not recognized by Mez (1933-1934) because the fragmentary collections availa- ble at that time permitted little insight into this difficult species. As a result, Mez (1904) described Thecophyllum violescens, a spe- cies he distinguished from T. pedicellatum on the basis of a lack of development of the lateral branches of the inflorescences and only two flowers per axil. Because the devel- opment of lateral branches of V. pedicellata is a variable character displaying many intermediate forms, it is impossible to main- tain V. violescens using this criterion. Be- cause flower number varies and is dependent on the development of the lateral branches, it cannot be used to characterize V. violescens. I consider V. violescens to be well within the variation I observed in V. pedicellata popu- 26 Tulane Studies in Zoology and Botany Vol. 24 lations and cannot maintain it as a distinct species. Vriesea turbinata, which I also con- sider to be conspecific with V. pediceliata, is represented by very fragmentary material. The scape is entirely wanting from the type and it is even doubtful that the entire post- fruiting inflorescence has been preserved. Moreover the leaf material is difficult to interpret, being either badly decomposed, mutilated or from the central portion of the rosette and consequently not representative. In light of the poor condition of this speci- men, I could not reach an unequivocal deci- sion regarding the disposition of the binom- ial, V. turbinata. However, the texture and size of the floral bracts and pedicel length in conjunction with the general form of lateral branches of the inflorescence conforms with that of V. pediceliata. In light of the available information regarding these taxa, I feel that V. turbinata should be considered a syn- onym of V. pediceliata. Several anomalous collections have been made of V. pediceliata at higher elevations in the Talamanca Mountains between El Empalme and Villa Mills (Utley & Utley 1518, 2763. 2754 and 5266). In all these col- lections the scape bracts are about equal to the internodes and flowers are frequently subtended by enlarged floral bracts. When further collections are available, this element may prove to be a distinct species. V. pediceliata is phenetically most similar to V. leptopoda. The characters distinguish- ing these two taxa are discussed under V. leptopoda. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE:Viento Fresco, 13 February 1926, Standley & Ruben Torres R. 48020 (US); north of San Ramdn, region from Los Angeles Norte to about 12 km north of La Balsa de San Ramdn, 15 December 1974, Utley & Utley 1653 (NOLS) & 1670 (DUKE); 8 August 1975, Utley & Utley 2768 (DUKE) & 2770 (CR); 27 January 1976; Utley & Utley 3794 (K) & 3795 (DUKE). CARTAGO PRO- VINCE: 10 miles south of Cartago on the Interamerican Highway, Talamanca Moun- tains, 10 July 1962, Haines & Haines 687 (US); 21 miles south of Cartago on Intera- merican Highway, 10 July 1962, Haines & Haines 701 (US) & 702(US); 16 miles south of Cartago on Interamerican Highway, 17 July 1962, Haines & Haines 719 (US); 22 miles south of Cartago on Interamerican Highway, 17 July 1962, Haines & Haines 723 (US),729 (US) & 752 (US); 29 miles south of Cartago on Interamerican Highway, 19 July 1962, Haines & Haines 734 (US); Cerro de La Carpintera, February 1924, Standley 34304 (US); north of El Muneco, 5 April 1974, Utley & Utley 785 (DUKE); Tapanti Hydroelectric Project about 7-14 km sou- theast of Orosi on slopes above the Rio Grande de Orosf, 13 December 1974, Utley & Utley 1605 (DUKE) & 1610 (F); 14 April 1975, Utley & Utley 2109 (DUKE), 2110, (MO) & 2111 (CR); 5-6 miles south of San Isidro de Cartago, 8 June 1975, Utley & Utley 2536 (DUKE), 2540 (US), 2541 (MICH) & 2542 (F); Tapanti Hydroelectric Project about 7-14 km southeast of Orosi, 5 December 1975, Utley & Utley 3415 (B) &3420 (MO), 3422 (CR), 3424 (F) & 3435 (DUKE); 22 March 1976, Utley & Utley 4357 (DUKE), 4358 (US), 4359 (CR), 4361 (DUKE), 4362 (CAS), 4365 (MO), 4366 (NOLS) «fe 4367 (DUKE). BORDER OF- C ARTAGO AND S AN JOSE PROVINCES: along the Interamerican Highway about 20 km south of Cartago, in the vicinity of Casa Mata, 17 March 1975, Utley & Utley 7995 (DUKE); 3 August 1975, Utley & Utley 2760 (DUKE); HEREDIA PROVINCE: 2-3 km southeast of the junc- tion of highways 9 and 120 on Hwy. 9, 27 March 1976, Utley & Utley 4375 (DVKE) & 4385 (F). SAN JOSE PROVINCE: San Jeronimo, March 1909, Biolley fil. 17365 (US); La Hondura, on tree, 2-4 March 1924, Standley 36220 (US); Alto de U Palma, north of San Jeronimo, 12 January 1974, Utley & Utley 617 (DUKE); 15 February No.l Middle American Vrieseas 27 1974, Utley & Ut ley 625 (CR); 1 June 1974 Utley & Utley 891 (US); 17 October 1974, Utley & Utley 1407 (NOLS); 15 April 1975, Utley & Utley 2136 (CAS), 2147 (NY), 2149 (CR), 2151 (DUKE) & 2152 (GH); 9 June 1975, Utley & Utley 2559 (DUKE) & 2560 (NOLS); 19 June 1975, Utley & Utley 2564 (DUKE) & 2569 (CR); vicinity of Bajo La Hondura, north of Alto de La Palma, 23 October 1975, Utley & Utley 3186 (DVKE); about 5 km northeast of San Isidrode Here- dia on Calle Yerbabuena, 3 April 1976, Utley & Utley 4465 (DUKE) 4467 (US) & 4468 (CR); between Aho de La Palma and Bajo La Hondura or 6-8km northeast of San Jeronimo, 6 April 1976, Utley & Utley 4514 (DUKE); at Cascajal about 3 km north of Las Nubes on Hwy. 216, 3 July 1976, Utley & Utley 5251 (DUKE). PUNTARENAS- PROVINCE: Monteverde, 17 km west of Las Juntas de Abangares, in the cloud forest above the Quaker settlement, 27 January 1975, Utley & Utley 1883 (DVKE, CR, US). HONDURAS: FRANCISCO MORAZAN: Montana Uyuca, off km 25, Tegucigalpa- Zamorano highway, 19 July 1964, Gilmartin 995 (US). NICARAGUA: DEPARTMENT OF JINOTEGA: Cordillera Central de Nicaragua above and east of Jinotega, 20 February 1963, Williams, et al. 24761 (F). DEPARTMENT OF MATAGALPA: cloud forest area at Disparate de Potter, near Sta. Maria de Ostuma, 20 & 24 February 1963, Williams, etal. 25052(F), 15 February 1965, Williams, et al. 27681 (F). 3. Vriesea diffusa L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Thecophyllum laxum Mez «fe Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 1123. 1904, not Vriesea laxa Mez, 1896. TYPE :COSTA RICA: without further location, Werckle Bromel. Costaric. 90 (HOLOTYPE: B!, photograph US!). Epiphytic or terrestrial, acaulescent. LEAVES in a spreading to subascending rosette, ( 1 3.5) 1 7.0-25.0 cm long; blades 8.0- 13.5 (15.0) cm long and 1.7-3.5 cm wide, green with a dense coat of appressed, cinere- ous trichomes on the abaxial surface, fre- quently with longitudinal maroon lines proximally, ligulate or subligulate to broadly triangular, apex acute to acuminate; sheaths (4.9) 5.5-1 1.5 (13.0) cm long and 2.6^.6 cm wide, pale green with longitudinal maroon lines, frequently suffused with maroon cen- trally. SCAPE erect or curved-erect, 20.0- 32.0 (42.5) cm long and 0.2-0.3 cm in diame- ter; bracts 1.8-2.8 (3.4) cm long and 0.7-1.6 cm wide, erect-clasping, occasionally spread- ing apically, green drying to brown, fre- quently with longitudinal maroon markings in the lower blade, much shorter than the internodes. INFLORESCENCE erect, (3.5) 5.5-16.0 cm long and 2.0-6.0 cm in diameter, more or less cylindrical; primary bracts 1 .2- 2.4 cm long and 0.6-1 .3 (2.0) cm wide, green drying brown, frequently with longitudinal maroon lines, apex acuminate. LATERAL BRANCHES spreading or ascending to rarely suberect, (1.8) 2.2-4.2 cm long, (1)2 (3)- flowered, these collateral to spicate, peduncle (1.6) 2.2-3.6 cm long, rachis want- ing or 0.5-0.9 (1.3) cm long. FLOWERS 0.3-0.4 cm pedicellate; floral bracts 0.5-0.7 cm long and 0.5-0.7 cm wide, brown, char- taceous, nerved, midvein frequently thick- ened and the bract slightly keeled, ovate to subtriangular, apex acute to obtuse or rounded and minutely apiculate; sepals 0.8- 1.0 cm long and 0.5-0.8 cm wide, light cas- taneous to olive-brown, coriaceous, elliptic, apex rounded; petalsl. 5-1. 6 cm long and 0.8- 0.9 cm wide, white, obovate; appendages 0.3 cm long and 0.1-0.15 cm wide; stamens in hood configuration over the gynoecium, fil- aments 1.0-1.1 cm long, anthers 0.2-0.3 cm long; capsules ( 1 .6) 2.0-2.7 cm long and 0.4- 0.6 cm in diameter, castaneous, fusiform. 28 Tulane Studies in Zoology and Botany Vol. 24 PHENOLOGY: Flowering specimens col- lected during June and August. DISTRIBUTION AND HABITAT: Known from Costa Rica and Panama between 900-1500 m in the premontane rain forest life zone. DISCUSSION: The characters that distin- guish Vriesea diffusa are pale green leaves with green blades that are frequently long- itudinally lined with purple proximally and pale green sheaths with longitudinal purple striations. Its slender scape (0.2-0.3 cm in diameter) bears scape bracts that are usually much shorter than the internodes. The peduncles of the lateral branches are un- usually long (2.2-3.6 cm) while the floral bracts exceed the pedicels. The flowers of V. diffusa are white, campanulate and noctur- nal, resembling in general morphology those of both V. triflora and V. pedicellata. Based on a number of characters, V. diffusa appears to be most closely related to V. tri- flora. Both taxa characteristically display papillose induments on the inflorescence rachis, peduncles and pedicels, a feature not encountered among other thecophylloid vrieseas. Inflorescence form in both taxa is very similar although inflorescences of V. triflora are much larger and more fully deve- loped than those of V. diffusa. The two taxa are readily separable on the basis of a number of characters. Both floral bracts and petals are shorter in V. diffusa than in V. triflora. In the vegetative condition, the taxa may be distinguished from each other on basis of leaf markings. Leaves of V. triflora characteristically have proximal wavy trans- verse lines, while those of V. diffusa are longitudinal. Both V. triflora dind V. diffusa diVQ^xohahXy most closely related to V. pedicellata and V. leptopoda, from which they are easily dis- tinguished by their densely papillose bran- ches as well as a suite of other characters. From the available collections, V. diffusa appears to have a more widespread dis- tribution than V. triflora. This may be a reflection of its occurrence at lower eleva- tions (900 m) than V. triflora (1400 m). How- ever, much of the Atlantic watershed of both Costa Rica and Panama is poorly collected and as these areas become better known flo- ristically, the distribution of V. triflora will possibly be expanded. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA: north of San Ramon, the region from La Balsa to about 4.5 km north, Utley & Utley 7,^57 (DUKE, US, F, CR, MO, K, CAS, GH, MICH, NOLS, CR); between La Balsa de San Ramon and the Rio Cataratas or about 12 km north of La Balsa, Utley & Utley 3740 (DUKE); about 4 km north of La Balsa de San Ramon on the road leading to the Atlantic water- shed, Utley & Utley 4121 (DUKE). BOR- DER OF ALAJUELA AND HEREDIA PROVINCES: vicinity of Colonia Virgen del Socorro, or about 3-6 km east of Carib- lanco, Utley & Utley 2880 (NOLS) & 2881 (DUKE). CARTAGO: Tapanti Hydroelect- ric Project, between 7 and 12 km south of the bridge over the Rio Grande de Orosi at Tapanti, Utley & Utley 3410 (DVKE). SAN JOSE: La Hondura, Standley & Valerio 51912 (US); La Palma, Werckle 9 (US). REPUBLIC OF PANAMA: PANAMA: Cerro Campana, trail to summit. Dressier 5141 (VS). 4. Vriesea triflora L. B. Smith & Pitt. J. Wash. Acad. Sci 43: 403. 1953. Thecophyllum panniculatum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. 11. 4: 1128. 1904, not Vriesea panicu- lata Mez. 1896. TYPE: COSTA RICA: SAN JOSE: la Palma, Werckle 55 (HOLOTYPE: B!, photograph US!; ISOTYPE: B!; photograph US!). Epiphytic or terrestrial, acaulescent. No.l Middle American Vrieseas 29 LEAVES in a spreading rosette, (23.0) 26.0- 34.0 cm long; Z)/a^e5( 10.0) 13.0-18.0 cm long and 2.8-4.2 (4.8) cm wide, green strongly suffused with wavy maroon cross-bands proximally, these more pronounced on the abaxial surface, ligulate to subtriangular with an acute to acuminate apex; sheaths (12.0) 13.0-15.5 (16.5) cm long and 4.0-6.0 cm wide, green densely suffused with maroon wavy cross-bands and longitudinal maroon lines, occasionally with a castaneous band at the base, covered with a coating of pale brown trichomes, elliptic. SCAPE erect, (3 1 .5) 36.0-46.0 cm long and about 0.3 cm in diameter; bracts frequently spreading distally, 2.0-3.0 cm long and 1 .0-2.0 cm wide, green frequently suffused with maroon and maroon longitudinal lines proximally and with maroon spots or transverse lines dis- tally, erect, middle and upper bracts much shorter than the internodes. INFLORES- CENCE erect, (29.0) 38.0^9.0 cm long and (11.0) 15.0-20.0 cm in diameter, cylindrical to conical in shape, rachis frequently sinuous distally, the surface densely papillose, purple- green in color with a metallic sheen; pr/marv bracts 1.8-2.5 cm long and 0.9-1.6 cm wide, green with longitudinal maroon lines and occasionally with maroon spots or trans- verse lines distally, apex acuminate to acute. LATERAL BRANCHES spreading, the distal portion becoming ascending and the fertile portion suberect, 8.0-14.0 cm long, racemosely (2) 3 (5)-flowered with a non- functional terminal bud, flowers frequently secund, peduncles (5.6) 7.0-10.0 cm long, rachis 1.8-4.4 cm long. FLOWERS 0.1-0.5 cm pedicellate; yZora/ bracts 0.9-1.2 cm long and 0.7-1 .2 cm wide, brown at anthesis, char- taceous, finely nerved, the central vein fre- quently thickened distally, ovate or orbicu- lar, apex rounded frequently apiculate, often torn and appearing incised; sepals 0.1 -1. 2 cm long and 0.6-0.9 cm wide, castaneous, cori- aceous, elliptic, apex acute to rounded; petals 1.9-2.0 cm long and 1.0-1.1 cm wide, white, obovate; appendages 0.3-0.4 cm long and 0. 1 -0.2 cm wide; stamens in a hood con- figuration over the gynoecium, anthers lat- erally connivent, filaments 1.4 cm long, anthers 0.4 cm long; ovary 0.4 cm long and 0.3 cm in diameter, style 1.2 cm long; cap- sules 1.5-2.5 cm long and about 0.5 cm in diameter, fusiform. PHENOLOGY: Flowering specimens col- lected in June. DISTRIBUTION AND HABITAT: Known from Costa Rica between 1400-1700 m in the premontane rain forest life zone and possibly extending slightly into the lower montane rain forest. DISCUSSION: Until my recent collections from Costa Rica, Vriesea triflora was poorly represented in herbaria. In both the vegeta- tive and flowering condition this taxon is difficult to distinguish from surrounding vegetation. In contrast to other thecop- hylloid vrieseas, the inflorescences of this taxon, although large, are slender and in- conspicuous and lack large scape and prim- ary bracts. Vriesea triflora is characterized by ligulate leaves with proximal, wavy transverse lines and an erect scape with bracts much shorter than the internodes. Its inflorescence is lax, with a characteristically papillose rachis. The lateral branches of the inflorescence are elongate and also densely papillose. The flowers of this taxon are white, nocturnal and campanulate, and very similar to those encountered in V. pedicellata and V. diffusa. Although V. triflora shares this common floral syndrome with V. pedicellata , its clos- est affinities are clearly with V. diffusa. This relationship is discussed in greater detail under V. diffusa. SPECIMENS EXAMINED: COSTA RICA: CARTAGO PROVINCE: Tapanti Hydroelectric Project, 7-12 km south of the bridge over the Rio Grande de Orosi at Tapanti, 5 December 1975, Utley & Utley 3417 (DUKE) & 3434{CR)\ 25 June 1976, Utlev & Utley 5160 (DUKE), 5162 (US), 30 Tulane Studies in Zoology and Botany Vol. 24 5163 (F), 5193 (NOLS), 5194 (MO), 5195 (DUKE), 5196 (CR), 5797 (NOLS),579« (CR). 5199 (F), 5200 (CAS), & 5201 (NY). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Heredia along the Rio Para Blanco, 28 May, 1976, Utley & Utley 5097 (DUKE). SAN JOSE PROVINCE: 4 km north of Cascajal on Hwy. 216, southwestern slopes of Volcan Irazu, 3 July 1976, Utley & Utley 5257 (DUKE); La Palma, Werckle 11 (US). COSTA RICA: without further loca- tion: September 1955, Lankester. C. s.n. (US). 5. Vriesea greenbergii Utley, sp. nov. Fig. 18 A V. umbrosa L. B. Smith, cui valde affmis, sepalis et petalis minoribus, follis rotundatis abrupte acutis differt. Epiphytic or terrestrial, acaulescent. LEAVES in a spreading to subspreading rosette, 32.0-40.0 (45.0) cm long; ^/a^e5 15.0- 22.0 (28.0) cm long and 3.8-5.8 (6.6) cm wide, green occasionally suffused with maroon distally, ligulate, apex acute or acuminate to rounded and abruptly acute; sheaths 12.0- 20.0 (26.8) cm long and (3. 1) 4.6-6.8 (8.0) cm wide, pale green drying light brown with a coating of brownish trichomes, usually with a dark castaneous band near base and occa- sionally with a trace of longitudinal lines, elliptic. SCAPE erect or curved-erect, (33.0) 43.0-49.0 cm long and 0.2-0.4 (0.7) cm in diameter; bracts clasping-erect, 3.0-4.8 cm long and 1 .8-3.4 (4.0) cm wide, green to green suffused with maroon distally, usually ex- ceeding the internodes but occasionally equaling or slightly shorter than the inter- nodes. INFLORESCENCE erect, (7.0) 13.0- 20.0 (32.5) cm long and 5.0-1 1 .0 ( 14.0) cm in diameter,laxly cylindric; primary bracts 2.4- 4.2 cm long and 2.0-3.6 cm wide, green fre- quently suffused with maroon distally, ovate to elliptic, apex acute or acuminate. LATERAL BRANCHES spreading to sub- ascending, (I.O) 1.8-3.2 (5.4) cm long, race- mosely 2-to 3 (5)-flowered, rachis (0.2) 1.0- 2.5 (4.2) cm long. FLOWERS 0.5-1.2 cm pedicellate; Jloral bracts 1 .0-1.3 cm long and 0.9-1 .2 ( 1 .5) cm wide, green occasionally suf- fused with maroon drying to brown, subco- riaceous, ovate to broadly elliptic or oblong, navicular, apex acute to obtuse or rounded and short acuminate; sepals (1.0) 1.3-1.7 cm long and 0.9-1 . 1(1.3) cm wide, green drying to brown, coriaceous, elliptic to obovate; petals 2.3-3.1 cm long and 1.1-1.5 cm wide, white, obovate; appendages 0.3-1.4 cm long and 0.15-0.2 cm wide; stamens in a hood configuration over the gynoecium, filaments 1.7-2.3 cm long, anthers 0.4-0.5 cm long; ovary 0.4-0.5 cm long and 0.3 cm in diame- ter, style 1 .2-2.0 cm long; capsules about 3.5 cm long and 0.8 cm wide, fusiform. TYPE: COSTA RICA: ALAJUELA PROVINCE: north of San Ramon, between La Balsa de San Ramon and the Rio Catara- tas or about 12 km north of La Balsa, elev. 900-1000 m, 27 January 1976, Utley & Utley 3741 (HOLOTYPE: DUKE! ISOTYPES: CR!, DUKE!, F!, US!). PHENOLOGY: flowering specimens have been collected in August, December, Janu- ary and February, with the peak occurring in January and February. DISTRIBUTION AND HABITAT: Endemic to Costa Rica, occurring between 800-1500 m in the premontane rain forest life zone. DISCUSSION: Vriesea greenbergii is an easily recognized taxon that is distinguished by its green, usually unmarked leaves borne in a spreading rosette, by well developed lateral branches (1.8-3.2 cm long) and by white, nocturnal flowers with weakly zygo- morphic corollas. Although V. greenbergii xs very abundant in the type locality, it was never encountered in similar habitats in other parts of Costa Rica. Future collecting may, however, reveal a more widespread No.l Middle American Vrieseas 31 Fig. 18. Vriesea greenbergii. A. Inflorescence and upper portion of the scape; B. Leaf; C. Flower; D, Flower with sepals and floral bract; E. Floral bract, torn apically; F. Fruit with sepals and floral bract. A, B, F, from (Uilev & Utiey 3741 J C, D, E, from (Utley & Utley 1688). 32 Tulane Studies in Zoology and Botany Vol. 24 distribution Vriesea greenbergii is most closely related to V. umbrosa. Although these taxa share a common floral morphology, no pleasant fragrance was observed in the flowers of V. greenbergii. V. greenbergii differs from V. umbrosa in floral bract length, petal length and leaf shape. Leaf blades of V. greenbergii are rounded and abruptly acute apically, in contrast to the acute to acuminate leaf apices of V. umbrosa. These species also have dif- ferent elevational distributions, with V. umbrosa occurring between 1200 and 1900 m and V. greenbergii between 800 and 1200 m. Both species share characters in common with V. latissima, which are discussed under this latter taxon. ETYMOLOGY: This species is named in honor of the noted aquatic plant horticultu- rist, Mr. Albert Greenberg, of Tampa, Florida. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: north of San Ramdn, region between Los Angeles Norte and about 7 km north, 15 December 1974, Utley & Utley 1688 (DUKE) & 1689 (CR); between La Balsa de San Ramdn and 12 to 13 km north, 8 February 1976, Utley & Utley 4059 (NOLS), 4064 (US), 4122 (CR), 4123 (GH), 4124 (DUKE), 4125 (MO) «& 4126 (NY). 6. Vriesea umbrosa L. B. Smith, Phytologia 23: 328-329. 1974. TYPE: COSTA RICA: SAN JOSE: La Palma area, northeast of San Jero- nimo, above the La Hondura valley, elev. 1500 m. Burger & Stolze 5333 (HOLOTYPE: US!; ISOTYPES: F!, CR!). Epiphytic or terrestrial, acaulescent. LEAVES in a speading to subascending rosette, (45.0) 51.0-80.0 (93.0) cm long; blades (25.0) 33.0-53.0 (63.5) cm long and (3.6) 4.8-6.4 cm wide, green ligulate to sub- ligulate or long triangular, apex acute to acuminate; sheaths 16.0-26.0 (29.0) cm long and 5.0-8.5 (10.0) cm wide, pale green drying to light brown with a coating of brownish grey to cinereous trichomes, elliptic, blend- ing gradually into the blade. SCAPE erect or curved-erect (47.0) 55.0-68.0 (77.0) cm long and 0.4-0.8 cm in diameter; bracts (4.0) 5.0-8.0 (9.0) cm long and 3.0-5.2 cm wide, green the middle and upper soon drying brown, clasping erect, divergent to spreading distally, exceeding the internodes, ovate to broadly elliptic or suborbicular, apex acute to acuminate. INFLORESCENCE erect, (17.0) 21.0-39.0 (51.0) cm long and 6.0-10.0 cm in diameter, cylindric, the surface gla- brous with minute black linear punctulae; primary bracts (3.5) 4.0-6.0 (7.6) cm long and 3.0-5.2 cm wide, green soon drying brown, ovate to elliptic, broad-elliptic or suborbicu- lar, apex acute to acuminate. LATERAL- BRANCHES spreading, to subspreading, (1.0) 2.0-3.4 cm long, racemosely (2) 4-to 5-flowered peduncles 0.1-0.6 (0.9) cm long, and the rachis (0.6) 1.2-2.2 (2.8) cm long. FLOWERS 0.3-0.8 (1.0) cm pedicellate; floral bractsl .A-\ .% (2.2) cm long and 1.1-2.2 cm wide, dried brown, subcoriaceous, sur- face even to indistinctly nerved, frequently with minute raised, clear to light brown, puncutulae, usually with a more or less thickened central area along which the bracts tend to fold, ovate to elliptic or subor- bicular, apex acute to obtuse or rounded, at times minutely incised due to tearing; sepals 1 .4-2.0 cm long and 0.9-1 .3 cm wide, green to yellow-green drying to light brown or cas- taneous, coriaceous, elliptic to obovate, the apex acute to obtuse rounded, at times min- utely incised due to itdiung; petals 3.6-4.2 cm long and 1 .6-2. 1 cm wide, white, tipped with maroon; appendages 0.4-0.6 cm long and 0.2-0.25 cm wide, stamens coalescent by the lateral margins of the anthers into a hood over the gynoecium, filaments 2.4-3.4 cm long, anthers 0.7-0.9 cm long; ovary 0.5-0.7 cm long and 0.3-0.4 cm in diameter, style No.l Middle American Vrieseas 33 3.0-3.4 cm long; capsules about 3.5 cm long and 0.7 cm in diameter, fusiform. PHENOLOGY: Flowering specimens have been collected during April and May. DISTRIBUTION AND HABITAT: Known from Costa Rica and Panama between 1200-1900 m in the lower montane and premontane wet forest life zones and lower montane and premontane rain forest life zone. DISCUSSION: Vriesea umbrosa was only recently described by Smith (1974) from fragmentary material. It is a distinctive spe- cies with green, unmarked leaves in a gener- ally spreading rosette. The inflorescence of V. umbrosa is erect, with well developed lateral branches bearing nocturnal, white zygomorphic flowers. Floral bracts in this taxon exceed the pedicels in length. On the occasions when I have collected V. umbrosa in flower, the flowers have had a sweet, plea- sant fragrance in the early evening and have produced copious nectar. Although confir- matory observations are lacking, this odor, combined with nocturnal anthesis, suggest a night flying insect pollinator. Vriesea umbrosa is most closely related to V. greenbergii and their relationship and the features which distinguish them are discussed under the latter species. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: west on the road from Zapote to Santa Elena or about 7-10 km north-northwest of Zarcero, 16 April 1976 Utley & Utley 4656 (CR), 4657 (US), 4658 (DUKE), 4659{NOLS), 4660 (GH), 4671 (MO), 4672 (F), 4673 (NY). CARTAGO PROVINCE: about 8 km north of El Empalme near the road to Palmi- tal,6 April 1974, Utley & Utley 814 (DUKE); on the road from Tapanti to Taus and Tau- sito about 3 km northeast of the Rio Grande de Orosi at Tapanti, 27 May 1976, Utley & Utley 5019 (DUKE). BORDER OF CAR- TAGO AND SAN JOSE PROVINCES: about 1 km north of the road to San Cristo- bal Norte on the Interamerican Highway, 9 May 1976, Utley & Utley 4753 (DUKE). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Heredia on Calle Yerbabu- ena, 3 April 1976, Utley & Utley 4478 (DUKE) & 4480 (F). SAN JOSE PRO- VINCE: between San Jeronimo and Alto de La Palma on Hwy. 220, 19 June 1975, Utley & Utley 2570 (DUKE) & 2571 (CAS); between Bajo La Hondura and Alto de La Palma, 24 May 1976, Utley & Utley 5004 (NOLS) & 5005 (DUKE). PANAMA: PROVINCE OF PANAMA: road to Altos de Pacora, near schoolhouse, 8 October 1974, Mori & Kallunki 2344 (MO). 7. Vriesea latissima (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Thecophyllum latissimum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 1122. 1904. TYPE: COSTA RICA: without further location, Werckle 82 (HOLOTYPE: B!, photograph US!). Vriesea rainfera L. B. Smith, Phyto- logia 9: 242. 1963. TYPE: COSTA RICA: SAN JOSE PROVINCE: in cloud forests of El Alto de la Palma, C.K. Horich s.n.; University of Cali- fornia Botanical Gardens accession number 58.482-1 (HOLOTYPE: US!). Epiphytic acaulescent. LEAVES in a sp- reading rosette, 45.0-67.0 cm long; blades 23.0-35.0 (40.0) cm long and 9.0-11.0 cm wide, green adaxially, strongly suffused with maroon abaxially, ligulate, apex rounded and acuminate; sheaths 24.0-28.0 cm long and 1 1.0-12.0 cm wide, scarcely wider than the blades, pale brown becoming green suf- fused with maroon distally, broadly elliptic. SCAPE erect 60.0-67.0 cm long and about 1.0 cm in diameter; bracts 6.0-7.0 cm long and 4.0-5.0 cm wide, exceeding the inter- 34 Tulane Studies in Zoology and Botany Vol. 24 nodes, green to green strongly suffused with maroon, erect. INFLORESCENCE erect, 32.0-52.0 cm long and 9.0-10.0 cm in diame- ter, axis green or green deeply suffused with maroon; primary bracts 5.0-6.0 cm long and 3.0-5.0 cm wide, green frequently mottled with maroon to entirely purple or brown- maroon, divergent to subspreading, apex acute to rounded and acuminate. LATER- AL BRANCHES divergent to spreading, 3.5-6.0 cm long, distichously 4-to 5-flowered, occasionally subsecund at anthesis, peduncle 1.0-2.0 cm long, rachis 2.0-3.7 cm long. FLOWERS 0.1-0.5 cm pedicellate; /7ora/ bracts 2.0-3.0 cm long and 1.0-3.0 cm wide, slightly shorter than to about equalling the sepals, green to green spotted or suffused with purple, subcoriaceous, ovate to elliptic, apex acute; sepals 2.1-2.8 cm long and 1.1- 1.3 cm wide, green, frequently suffused with maroon proximally, coriaceous, elliptic, apex broadly acute or obtuse to rounded; petals about 3.5 cm long and 1.5 cm wide, white, obovate; appendages 0.4 cm long and 0. 1-0.2 cm wide; stamens in a hood configu- ration over gynoecium, anthers laterally connivent, filaments 2.2-2.3 cm long, anthers 0.7 cm long; ovarv 0.5 cm long and 0.4 cm in diameter, style 2.5 cm long. PHENOLOGY: Flowering specimens have been collected in April. DISTRIBUTION AND HABITAT: Known only from Costa Rica in the region between 900-1500 m in the premontane rain forest life zone. DISCUSSION: The conspicuous features which set Vriesea latissima apart from most other thecophylloid vrieseas include a spr- eading rosette, broad (9.0-1 1.O cm), ligulate leaves that are green adaxially but maroon abaxially, a stout scape and coriaceous floral bracts which are 2.0-3.0 cm long. Unlike other members of the pedicellata complex, lateral branches of the inflorescence of V. latissima are coated with a gelatinous exu- date. This condition had been noted in sev- eral groups in the hygrometrica complex, the comata, capitata and hygrometrica species groups. The white nocturnal flowers of V. latissima become subsecund at anthesis. The observed floral syndrome of these large white, noc- turnal flowers, borne on a stout inflorescence well above the surrounding foliage, is com- patible with that presented by Faegri and van der Pijl (1971) for chiropterophilous plants. In fact, bats have been observed visit- ing the flowers of V. latissima (Phillip De Vries, pers. comm.). Vriesea latissima shares a number of char- acters with V. greenbergii and V. umbrosa. In all three species the lateral branches of the inflorescence are short, stout pedunculate, with well-developed rachises. Their short pedicellate flowers are subtended by cori- aceous floral bracts which exceed the pedic- els in length. V. latissima is easily separated from V. greenbergii and V. umbrosa on the basis of leaf width, floral bract length and floral morphology. In the vegetative condition, V. latissima may be mistaken for V. leptopoda since they share a common leaf and rosette morphol- ogy. However, these taxa are not sympatric and have different elevational distributions. V. latissima is consistently found between 900 m and 1500 m in elevation; in contrast, V. leptopoda grows at higher elevations (1600-2300 m). In flower or fruit V. latissima may be distinguished from V. leptopoda by pedicel, sepal and floral bract length. Vriesea ranifera was characterized by Smith (1963) and Smith and Downs (1977) as having scape bracts which equaled or exceeded the internodes; in contrast, scape bracts of V. latissima appeared to be shorter than the internodes. While the scape bracts of the type specimen of V. latissima are, in fact, shorter than the internodes, this is a result of the age of the inflorescence, which is well past fruiting, and is also a consequence of the drying and weathering of the scape No.l Middle American Vrieseas 35 bracts, which are largely wanting. This con- dition, though not readily observable in a photograph of the type of V. latissima, is obvious upon critical examination of the type material. The type of V. ranifera agrees well with that of V. latissima in numerous characters, including rosette shape, leaf shape, width and color pattern, lateral in- florescence development, peduncle length, and sepal length and width. In the absence of demonstrable differences in critical charac- ters and in view of a compelling series of common characters, V. ranifera must be considered a synonym of V. latissima. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: fields, road banks and forest remnants between Los Angeles Norte and ca 7 km north of La Balsa de San Ramon or about 10-17 km north of San Ramon, 8 August 1975, Utley & Utley 28 14 {DVKE)& 2884 (CR). BORDER OF HEREDIA AND SAN JOSE PROVINCES: 5 km northeast of San Isidro de Heredia on Calle Yerbabuena, 3 April 1916. Utley & Utley 4470 (DUKE) & 4479 (NOLS). SAN JOSE PROVINCE: La Palma, Werckle 12 (US); between La Hondura and Alto de La Palma, 24 May 1976, Utley & Utley 5006 (DUKE). 8. Vriesea lyman-smithii Utley, sp. nov. Fig. 19 A V. nephrolepis L.B. Smith & Pitt, cui valde affmis corollis subtubularibus, sepalis rotundis, stylis brevioribus, capsulis longio- ribus differt. Epiphytic, acaulescent. LEAVES in a spreading to subspreading rosette, 45.0-61.0 cm long; blades 25.0-35.0 cm long and 4.2- 7.0 cm wide, green to green suffused with purple, ligulate to broadly triangular, acute; sheaths 20.0-26.0 cm long and (7.5) 9.5-1 1 .5 cm wide, pale brown at times with an abax- ial, basal castaneous band, usually w'*h maroon wavy crossbands in the distal region, ovate to elliptic. SCAPE erect (45.0) 55.0-68.0 cm long and 0.7-1.0 cm in diame- ter; bracts 7.0-15.0 cm long, blades 3.5-9.0 cm long and 1 .8-4.0 cm wide, green or green with a fine maroon margin and apex, drying to brown, erect to spreading or divergent, occasionally spirally involute, triangular, acu\.e;sheaths (3.2) 3.6-5.2 (5.8) cm long and 4.0-6.6 cm wide, drying ochre abaxially and light brown adaxially, sheathing-erect ovate. INFLORESCENCE erect, 16.0-33.0 cm long and 9.0-14.0 cm in diameter, cylindric to conical in outline, surface glabrous; pr/m- ary bracts 7.0-13.2 cm long, blades (3.0) 3.6- 5.2 (7.8) cm long and 2.0-3.0 cm wide, green, divergent to spreading frequently reflexed at or near the junction with the sheath, triangu- lar with an acute apex, sheath 3.5-5.7 cm long and 4.2-6.7 cm wide, drying ochre abax- ially, pale brown adaxially, ovate. LATER- AL BRANCHES spreading, 0.4-1.0 cm long, bearing 2 collateral or subcollateral flowers, occasionally with a vestigial third flower, peduncles 0.4- 1 .0 cm long, rachis 0.0- 0.3 cm long. FLOWERS sessile or to 0.3 cm pedicellate;y7ora/ bracts 0.9-1 .4 cm long and 1.6-2.3 cm wide, membranaceous to subco- riaceous, finely nerved, usually brown to pel- lucid punctate abaxially and obscurely lepi- dote adaxially, broadly ovate to reniform, apex obtuse to broadly rounded, frequently minutely acuminate or torn-incised; sepals 2.0-2.6 cm long and 1.6-2.1 cm wide, green with maroon margins, drying brown, cori- aceous, occasionally keeled, glabrous aba- cially, lepidote adaxially, elliptic to obovate, apex obtuse to rounded or broadly rounded; frequently suffused with maroon proxim- ally, coriaceous, elliptic, apex broadly acute or obtuse to rounded; petals 3.6-4.0 cm long and 1 .7-1 .9 cm wide, white to greenish white, ovate to elliptic, incised on either side where the petals are exserted from the sepals; appendages 0.7-0.8 cm long and 0.3-0.5 cm wide, stamens in a hood configuration over the stigma, filaments 2.2-2.9 cm long, anth- 36 Tulane Studies in Zoology and Botany Vol. 24 ers 0.9-1.1 cm long; ovary 0.8-1.0 cm long and 0.5-0.6 cm in diameter, style 1.7-2.1 cm long; capsule 4.9-5.3 cm long and 1 .3-1.5 cm in diameter, castaneous, obovate. TYPE: COSTA RICA: HEREDIA PROVINCE: 1-2 km south of the junction of highways 9 and 120 on Hwy. 9, slopes above the Rio Desengano, 22 May 1976, Utle\&Vtle\ 4960 {WOhOJ^?^,: DUKE!; ISOTYPES: CAS!, CR!, DUKE!, F!, MO!, US!). PHENOLOGY: Flowering specimens have been collected in May and June. DISTRIBUTION AND HABITAT: Known from the Cordillera Central de Costa Rica and the northern slopes of the Cordil- lera de Talamanca between 1500-2100 m in the lower montane rain forest life zone. DISCUSSION: Vriesea lyman-smithii is recognizable at once by its relatively broad leaves which form a spreading rosette, its stout scape and dense inflorescence. The sheaths of the primary bracts are broad and tend to cover the relatively large sepals. Flowers of V. lyman-smithii are white to greenish white and nocturnal. The corollas are subtubular with the two adaxial petals arranged in a position to form a normal tubular corolla; however,the abaxial petal is flexed slightly outward. Vriesea lyman-smithii is most closely allied with V. nephrolepis from which it differs by a combination of characters. The scape is gen- erally longer than that of V. nephrolepis, while the sheaths of the primary bracts are broader than those of V. nephrolepis. The broad primary bract sheaths in conjunction with a more dense nature of the inflorescence give the inflorescence of Vriesea lyman- smithii a characteristic appearance in both living plants and dried herbarium material. These features, when taken together with the subtubular flowers, generally rotund sepals and shorter styles ( 1 .7-2. 1 cm vs. 2.4-3.7 cm) and larger capsules (4.9-5.3 cm vs. 2.4-3.7 cm), consistently separate Vriesea lyman- smithii from V. nephrolepis. Despite the rather recondite nature of some of the characters, the two species are readily separable in the field as well as in the herba- rium. Although both taxa occur in the lower montane rain forests, they are generally allo- patric. However, they have been encoun- tered growing sympatrically at Cerro Zurqui in the Cordillera Central. ETYMOLOGY: This species is named in honor of Dr. Lyman B. Smith of the Smith- sonian Institution whose monumental labors in the taxonomy of the Bromeliaceae have stimulated and formed the basis for studies of the biology, ecology and sys- tematics of this unique epipthytic family. Dr. Smith has willingly shared his vast know- ledge and experience with all who have sought his advice. SPECIMENS EXAMINED: COSTA RICA: HEREDIA PROVINCE: region north of Cerro Chompipe, about 1 5 km nor- theast of Heredia and 3 km north of Cerro Redondo de La Cruz, 15 October 1974, Utley & Utley 1313 (DUKE); 8 December 1975, Utley & Utley 3583 (DUKE). BOR- DER OF HEREDIA AND SAN JOSE PROVINCES: along the Rio Para Blanco in the vicinity of Cerro Zurqui, 18 June 1974, Utley & Utley 963 (DUKE). SAN JOSE PROVINCE: 4 km north of Cascajal or 7 km north of Las Nubes on Hwy. 216, 3 July 1976, Utley & Utley 5271 (DUKE). 9. Vriesea nephrolepis L.B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953 Figs. 14, 20. Thecophyllum werckleanum Mez, Bull. Herb. Boiss. ser. II. 3: 139. 1903 not Vriesea werckleana Mez, 1903. TYPE: COSTA RICA: near Cartago, elev. 1200-1400 m, Werckle in Herb. Inst. Phys.-Geogr. Cos tar. 16201 (HOLOTYPE: B!; ISOTYPES: B! GH). No.l Middle American Vrieseas 37 Thecophyllum montanum L. B. Smith in Yuncker, Field Mus. Bot. 17: 319. 1982. TYPE: HONDURAS: COMAYAGUA: El Achote summit , above Siguate- peque, Yuncker, Dawson & Youse 6423. (HOLOTYPE: GH!). Fig. 19 Vriesea lyman-smithii. A. Inflorescence; B. Leaf; C & D. Flower with floral bract; E. Mature capsule with sepals and floral bract. A, B, from (Utley & Utlev 4960); C & D from (Utley & Utley 4981 X E, from (Utlev & Utley 1313). 38 Tulane Studies in Zoology and Botany Vol. 24 Vriesea montana (L.B. Smith) L.B. Smith & Pitt. J. Wash. Acad. Sci. 43: 69. 1953. Epiphytic or rarely terrestrial, acaulescent to 15 cm caulescent. LEAVES in a spreading rosette, (20.0) 27-46 (61.0) cm long; blades (9.0) 13-35 (42.0) cm long and 3.4-6.2 cm wide, green or at times suffused with maroon, usually with dark green wavy crossbands, these frequently suffused with maroon near the blade-sheath junction or occasionally suffused with maroon through- out the blade, ligulate, apex acute to acumi- nate or rounded and acuminate or rarely attenuate; sheaths 10.0-22.0 (26.0) cm long and (5.2) 6.2-9.6 ( 10.4) cm wide, pale brown, frequently with a basal castaneous band or medial castaneous spot on the abaxial sur- face or rarely the entire abaxial surface cas- Fig. 20. Vriesea nephrolepis. habit. taneous, elliptic to ovate or rarely obovate, usually with fine maroon wavy crossbands in the upper portion. SCAPE erect or curved- erect, 27.0-57.0 (67.0) cm long and (0.3) 0.4- 0.7 ( 1 .0) cm in diameter; bracts erect to erect and spreading distally, reflexed or rarely spi- rally involute distally, (3.6) 5.0-12.0 (16.2) cm long, blades ( 1 .6) 2.0-8.6 ( 1 1 . 1) cm long and 0.9-3.3 (4.5) cm wide, green with green, maroon or pinkish transverse lines, the upper usually dried brown at anthesis, tri- angular to subligulate with an acute apex, sheaths clasping erect, (1.6) 2.0-4.4 (5.8) cm long and (2.2) 2.8-5.0 (5.6) cm wide, ochre to green in living material, drying to ochre or brown to castaneous abaxially and pale brown adaxially, ovate to elliptic. INFLO RESCENCE erect, 9.0-28.0 (32.0) cm long and 4.0-10.0 (12.0) cm in diameter, cylindric, rachis glabrous; pr/marv bracts spreading to subascending, usually reflexed or occasion- ally spirally involute at the blade sheath junc- tion (3.2) 4.0-8.6 (10.8) cm long, blades 1.6- 5.6 (7. 1 ) cm long and 0.8-3.2 cm wide, green, at times tinged with maroon near margins, frequently with dark green or pink to red wavy crossbands, usually drying to brown at anthesis, triangular in shape with an acute or rarely acuminate apex; sheath 1.6-4.2 cm long and (2.2) 2.8-5.4 cm wide, green or green with a trace of ochre to strongly ochre in living material, drying brown to ochre abaxially and pale brown to white adaxially, ovate to rarely elliptic. LATERAL BRANCHES spreading, 2flowered, occa- sionally with a third vestigial bud, sessile to 0.6 (1.0) cm pedunculate, rachis usually wanting and the flowers collateral, or to 0.3 cm long and the flowers subcollateral. FLOWERS sessile to 0.6 (0.8) cm pedicel- late; /7ora/ bracts 0.8-1.4 cm long and (1.1) 1.3-2.0 (2.2) cm wide, subcoriaceous or rarely coriaceous, pale green or green, at times suffused with maroon, drying brown, glabrous or minutely brown to pellucid punctulate abaxially and scattered lepidote adaxially, suborbicular, broadly ovate to ovate or reniform, apex truncate to obtuse or No.l Middle American Vrieseas 39 broadly rounded, at times minutely acumi- nate, frequently tearing medially and appear- ing incised, keel usually wanting, occasion- ally present and inflated; sepals 1.5-2.2 (2.6) cm long and 1.0-2.0 cm wide, coriaceous, green frequently suffused with maroon espe- cially marginally or apically drying brown to castaneous, elliptic to obovate, apex rounded to broadly rounded or ohiu^t, petals (3.1) 3.4-4.5 (5.4) cm long and 1.2-1.9 cm wide, white to greenish white, ovate-lanceolate to lanceolate or slightly obovate, constricted at the point where the petals emerge from the sepals; appendages 0.3-0.8 cm long and 0.2- 0.4 cm wide, stamens in a hood over the gynoecium, usually connivent by the lateral margins of the anthers into a single grouping of six or two groupings of three, rarely free, filaments (2.5) 2.9-3.9 (4.7) cm long, anthers 0.6-1.0 cm long; ovary 0.6-0.8 cm long and 0.4-0.5 cm in diameter, style 2.4-3.7 (5.0) cm long; capsule 3.3-4.3 cm long and 1.1-1.7 cm in diameter, castaneous, fusiform to obovate. PHENOLOGY: Flowering specimens have been collected in March through July with a distinct peak occurring March-May. DISTRIBUTION AND HABITAT: Known from Costa Rica and Panama north- ward to Guatemala between 1500-2800 m in the lower montane wet forest and rain forest life zones and the premontane and montane rain forest life zones. DISCUSSION: Vriesea nephrolepis is the most commonly encountered night flower- ing taxon in the Cordillera de Talamanca and the volcanos of the Cordillera Central in Costa Rica. It is a distinctive species charac- terized by a spreading rosette form and green leaves which are frequently suffused with maroon and green to maroon wavy trans- verse lines. The inflorescence of V. nephrole- pis is erect and basally lax, bearing large white to green-white nocturnal flowers with zygomorphic corollas. The primary bracts are generally green, or rarely green with maroon wavy transverse lines at anthesis. Specimens have also been collected in which the primary bracts are either dried and brown at anthesis or very infrequently green suffused with red-maroon at anthesis. The primary bract sheaths of V. nephrolepis are frequently ochre colored and become rugose upon drying. Although the type of V. nephrolepis lacks corollas, the rugose sheaths of the primary bracts are prominent. The dense aggreated wavy crossbands which are present in the leaves of many collections are also character- istic of the type material of V. nephrolepis. The type collection of this taxon was made in the vicinity of Cartago between 1200 and 1400 m. This is more than 500 meters below the lowest elevation at which I have collected V. ororiensis, but within 100 m of the lowest collections of V. nephrolepis . Collections from the lower slopes of the Cordillera de Talamanca near Cartago agree in general size, leaf markings, vigor and form of the inflorescence and the form of the primary bracts and sepals with the type collection. This correlation of elevation, general locality and critical morphological features support the interpretation of V. nephrolepis that I employ. Although Smith (1977) stated that V. neph- rolepis was only known from the type mate- rial, there were a number of collections of this taxon available but they were identified as V. irazuensis, a taxon which is conspecific with V. ororiensis. This is presented in greater detail in the discussion of V. ororien- sis. Both Mez (1934-1935) and Smith (1977) emphasized the number of flowers in the lowermost branches of the inflorescences. In Mez's treatment V. nephrolepis ( Thecophyl- lum werckleanum) keys out only under the lead that requires three flowers per lateral branch. Smith's monograph is similar in this respect. While the type of V. nephrolepis does have three flowers in the lowermost lateral branches, this is an unusual condi- tion. The most frequent number of flowers per lateral branch that I observed in my 40 Tulane Studies in Zoology and Botany Vol. 24 collections was two. Smith (1938) estab- lished Thecophyllum montanum (Vriesea montana) and distinguished the new taxon from Thecophyllum werckleanum on the basis of sheath color, size and density of primary bracts and shape of the floral bracts. The type and other specimens of V. montana I have seen fall well within the range of varia- tion observed in Vriesea nephrolepis and I consider the two taxa to be conspecific. The solution to the problems associated with V. ororiensis, V. nephrolepis and V. montana was dependent on extensive field work. The collections extant at the time of Mez's and Smith's treatments were insufficient to add- ress these problems. Vriesea nephrolepis is similar to V. lyman- smithiiand their relationship is elaborated in the discussion of the latter species. In the vegetative or fruiting condition, V. neph- rolepis is easily confused with V. ororiensis. The best character for separating these mor- phologically very similar taxa in fruit is the marked tendency for the primary bracts of V. nephrolepis to appear rugose in sheath area upon drying. In flower the floral syn- dromes of these taxa are distinctly different and permit unequivocal identification of both V. ororiensis and V. nephrolepis. SPECIMENS EXAMINED: COSTA R1CA:ALAJUELA PROVINCE: between the junction of highways 9 and 120 and the summit of Volc^n Poas, 11 March 1975, Utley & Utley 1927 (DUKE) & 1928 (CR); on the finca of Mike Canon near the junction of highways 9 and 120,20 July 1975, Utley & Utley 2664 (DUKE); between Poasito and Volc^n Poas, along Hwy. 120. CARTAGO PROVINCE: 6 miles south of Cartago on Interamerican Highway, Talamanca Moun- tains, 10 July 1962, Haines & Haines 676 (US); 10 miles south of Cartago on Inter- american Highway, Talamanca Mountains, 10 July 1962, Haines & Haines 685 (US); 19 miles south of Cartago on Interamerican Highway, Talamanca mountains, 10 July 1962, Haines & Haines 700 (US); 16 miles south of Cartago on Interamerican High- way, north slope of Talamanca Mountains, 17 July 1962, Haines & Haines 722 (US); 21 miles south of Cartago on the Interamerican Highway, north slope of Talamanca Moun- tains, 19 July 1962, Haines & Haines 730 (US); Estate of C.H. Lankester, 27 February 1966, L.B. Smith 15301 (US); on tree, cloud forest, Aserri, 19 March 1966, L.B. Smith 15325 (US) & 15326 (US); Cerro de La Car- pintera, February 1924, Standley 34463 (US); south of Cartago, about 3 km south of the Interamerican Highway on a secondary road, vicinity of Palo Verde, 17 April 1975, Utley & Utley 2190 (NOLS), 2191 (DUKE) & 2192 (UC); 3-6 miles south of San Isidro de Cartago on the Interamerican Highway; 8 June 1975, Utley & Utley 2518 (DUKE) & 2537 (US); about 1/2 km south of Pacayas on Hwy. 230, 25 July 1975 Utley & Utley 270,S(DUKE), 2709(F) &2770(DUKE); on Hwy. 230 between Trinidad and Coliblanco, 25 July 1975, Utley & Utley 2711 (DUKE), 2720 (MO) & 2721 (NOLS); on Hwy. 216 beyond Cascajal, northwestern slopes of Volc^n Irazu, 10 December 1975, Utley & Utley 3617 (DUKE); BORDER OF CAR- TAGO AND SAN JOSE PROVINCES: in the vicinity of a montane bog 17 km sou- theast of El Empalme on the Interamerican Highway, 6 April 1974, Utley & Utley 809 (DUKE); within 2-3 km of the Interamerican Highway on the road from El Empalme to Santa Maria de Dota, 6 April 1974, Utley & Utley 816 (DUKE); junction of highways 2 and 223 about 5 km north of El Empalme, 16 April 1975, Utley & Utley 2169 (DUKE); Interamerican Highway about 12 km sou- theast of El Empalme in the vicinity of Paso Macho, 3 August 1975, Utley & Utley 2745 (DUKE), 2749 (CR) & 2750 (DUKE). HEREDIA PROVINCE: roadside and tree falls between Vara Blanca and Chinchona on Highway 9, 16 October 1974, Utley & Utley /J20(DUKE), 1321 (US)& /i22(F); upper slopes of Volcan Barba about 8 km beyond Porosati, 16 March 1975, Utley & No. Middle American Vrieseas 41 Utley 1964 (DUKE); 12 April 1975 Utley & Utley 2W0 (DUKE), 2046 (GH), 2047 (NY), 2048 (CAS) & 2049 (NOLS); about 1 km south of Monte de la Cruz on Hwy. 1 13, 1 April 1976, Utley & Utley 4448 (DUKE); 2 km south of the entrance to Parque Nacional Volcan Poas, 3 May 1976, Utley & Utley 47IliDVKE),4713i^OLSl47i4(F),4715 (GH), 4716 (B), 4717 (MO), 4718 (K), 4719 (NY), 4720 (BM), 4721 (DUKE), 4722 (MICH), 4723 (CR), 4724 (NOLS), & 4725 (DUKE); near the junction of highways 9 and 120, 21 June 1976, Utley & Utley 5153 (DUKE) & 5156 (NOLS). BORDER OF- HEREDIA AND SAN JOSE PROVIN- CE: along the Rio Para Blanco near Cerro Zurqui, 10 July 1974, Utley & Utley 935 (DUKE); 18 June 1974, Utley & Utley 973 (DUKE); 22 December 1974, Utley & Utley 1748 (DUKE). SAN JOSE PROVINCE: between San Jerbnimo and Alto de La Palma on Hwy. 220, 1 June 1974, Utley & Utley 892 (DUKE); between San Jeronimo and Alto de La Palma on Hwy. 220, 29 June 1974, Utley & Utley 991 (DUKE); 1-2 km south of the Interamerican Highway on the road from El Empalme to Santa Maria de Dota, 16 April 1975, Utley & Utley 2159 (MO), 2161 (F), 2164 (DUKE) &. 2168 (MO); between San Jeronimo and Alto de La Palma on Hwy 220, 1 June 1974, Utley & Utley 892 (DUKE); between San Jeronimo and Aho de La Palma on Hwy. 220, 6 July 1975, Utley & Utley 2584 (F), 2585 (US), 2591 (DUKE) & 2596 (US); between Casca- jal and 8 km northeast on Hwy. 216, 23 August 1975, Utley & Utley 2944 (DUKE); Cedral (Cerros de Escazii), approach to and summit of Cerro Daser about 3-5 km south- west of Aserri, 13 March 1976, Utley & Utley 4264 (DUKE), 4266 (DUKE), 4282 (CR) & 4286 (MICH); 2 April 1976, Utley & Utley 4450 (CAS) & 4451 (DUKE); at Cascajal about 3 km north of Las Nubes on Hwy. 216, 3 July 1976, Utley & Utley 5239 (DUKE), 5240 (US), 5252 (CR) «& 5253 (F). EL SAL- VADOR: DEPARTMENT OF SANTA ANA: moist cloud forest on Cordillera Miramundo, Mountain of Montecristo, 27- 31 January 1966, Molina, A. W. Burger and B. Wallenta 16920 (US). HONDURAS: Montana Uyuca, off Tegucigalpa-Zamorano road, trail starts Km. 25, 10 July 1964, Gil- martin, A.S. 956 (US). PANAMA: PRO- VINCE OF VERAGUAS: NW of Santa Fe, 8.8 km from Escuela Agricola Alto de Piedra. Pacific Slope, 25 February 1975, Mori & Kallunki 4845 (MO). 10. Vriesea ororiensis (Mez) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953. Figs. 11, 16 Guzmani ororiensis Mez in DC. Mon. Phan. 9:917. 1896. TYPE: COSTA RICA: Province not given. Oersted Brom. n. 25, Volcan Irazu, elev. 2200 m; Donnell Smith 4963. [LECTOTYPE: Oersted Brom. n. 25 (C!); SYNTYPE: Donnell Smith n. 4963 (US!)]. Thecophyllum ororiense (Mez) Mez, Bull. Herb. Boiss. ser. II. 3: 131. 1903. Thecophyllum crassiflorum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II 3: 138. 1903. TYPE: COSTA RICA: Province not given, near Cartago, elev. 1200-1400 m, Werckle, Inst. Phys-Geogr. Costar. 16188 Thecophyllum irazuense Mez & Werckle, Bull. Herb. Boiss. ser. II. 3: 138. 1903. TYPE: COSTA RICA: Province not given, on Volcan Irazu near Las Pacayas de Cartago, elev. 1800 m, Werckle, Inst. Phys. -Geogr. Costar. 16206. (HOLOTYPE: B!, photograph US!; ISOTYPE: GH). Thecophyllum standleyi L.B. Smith, Contr. Gray Herb. 117: 30. 1937. 42 Tulane Studies in Zoology and Botany Vol. 24 TYPE: COSTA RICA: SAN JOS^: on tree in paramo thicket, Cerro de las Vueltas, elev. 2700-3000 m, Standley & Valerio 43573 (HOLOTYPE: US!). Thecophyllum cylindraceum Suessen- guth &. Goeppinger, Bot. Jahrb. 72: 292. 1942. TYPE: COSTA RICA: Canaan, elev. 1300 m, Kupper 1252 (HOLOTYPE: M!, photograph US!). Thecophyllum kupperi Suessenguth & Goeppinger, Bot. Jahrb. 72: 292. 1942. TYPE: COSTA RICA: Location not given, Kupper s.n. (HOLOTYPE: M!, photograph US!). Vriesea crassiflora (Mez & Werckle) L.B Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Vriesea irazuensis (Mez & Werckle) L.B Smith & Pitt. J Wash. Acad. Sci. 43: 403. 1953. Vriesea standleyi (L.B. Smith) L.B Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953. Vriesea cylindraceae (Suessenguth & Goeppinger) L.B Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Vriesea kupperi (Suessenguth & Goeppinger) L.B Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Epiphytic or rarely terrestrial, acaulescent to short (4.0-6.0 cm) caulescent. LEAVES 1 7.0-48.0 cm long; /?/at/e5 13.5-36.0 (46.5) cm long and (2.7) 3.3-6.5 (7.3) cm wide, green or green variously suffused with maroon, occa- sionally entirely maroon, usually with fine wavy cross bands, these maroon, dark green or dark green suffused with maroon, com- monly more distinct abaxially or near the blade-sheath junction, ligulate to subligulate or rarely broadly triangular, apex acute or acuminate to attentuate or rounded and acuminate; sheaths (7.5) 10.0-25.0 cm long and 5.0-10.5 cm wide, castaneous abaxially and pale brown adaxially or pale brown on both surfaces, usually with castaneous band near base and with dark green or maroon wavy cross bands distally, elliptic to obovate. SCAPE erect or curved-erect, (32.0) 35.0- 68.0 (74.0) cm long and 0.3-1.0 cm in diame- ter; bracts 2.4-24.5 cm long; blades 1.8-14.0 (19.0) cm long and (I.O) 1.6-3.4 (4.2) cm wide, blades of the lowermost bracts usually green with dark green, maroon or red wavy cross-bands, middle and upper green, pale cream, pink or crimson with cross-bands of pink to crimson, erect to divergent distally, ligulate to subligulate, apex rounded or attentuate rarely rounded and short acumi- nate, sheaths 0.6-5.0 cm long and 2.2-4.5 (5.5) cm wide, ochre or green occasionally suffused with red or pink, drying ochre to pale brown abaxially and pale brown adax- ially, if colored when living occasionally retaining a trace of the color when dry, clasping-erect, ovate to elliptic. INFLOR- ESCENCE erect, (8.0) 19.0-35.0 (39.0) cm long and 5.0-12.0 cm in diameter, cylindric to subcylindric or narrowly conical in out- line, rachis glabrous green to green mottled with maroon or scarlet or entirely scarlet; primary bracts spreading at anthesis, fre- quently drooping-reflexed or spirally re- curved at the blade-sheath junction, blades 1.8-9.5 (13.9) cm long and 1.1-4.0 cm wide, pink to scarlet, frequently with fine, wavy pink to crimson cross bands, triangular to subligulate or rarely ligulate, apex acute to rarely acuminate or attenuate, sheaths (1.1) 1.6-3.8 (4.4) cm long and (2.2) 2.4-4.5 (6.0) cm wide, green to pale cream or ochre occa- sionally suffused with pink to red wavy cross-bands, ovate to elliptic. LATERAL- BRANCHES spreading, to subascending, 2-flowered or rarely with a third vestigial bud, sessile to 1.1 cm pedunculate, rachis absent and the flowers collateral to 0.3 (0.6) cm long and the flowers subcollateral. FLOWERS sessile to 0.5 cm pedicellate, No.l Middle American Vrieseas 43 floral bracts 0.7-1.7 cm long and 1.0-1.2 cm wide, subcoriaceous to membranaceous, keel usually pronounced, frequently inflated, rarely obscure to wanting, green to pale green occasionally weakly suffused with red or maroon, drying brown post anthesis, abaxial surface minutely pitted, usually with castaneous to pellucid punctulae, finely nerved or rarely subeven, scattered lepidote adaxially, ovate or broadly ovate to elliptic or suborbicular, apex acute to obtuse or acuminate or rounded and minutely acumi- nate, at times rostrate, frequently torn at anthesis and the shape not obvious; sepals (1.3) 1.5-2.4 cm long and 1.1-2.0 cm wide, coriaceous, keeled, green drying yellow- green or castaneous, obovate to elliptic, apex round, torn-incised to truncate or acute; petals (2.0) 2.5-3.0 (3.4) cm long and 0.8-1 .3 cm wide, green elliptic to obovate or sub- pandurate; appendages (0.2) 0.6-0.7 cm long and 0. 1-0.3 cm wide, stamens in a hood con- figuration over the gynoecium, filaments (1.2) 1.4-2.0 cm long, anthers 0.6-0.9 cm long, coherent laterally into an androecial hood; ovary 0.4-0.6 cm long and 0.3-0.4 cm in diameter, style (1.4) 1.6-1.9 cm long fre- quently flexed abaxially and the stigma oriented away from the androecial hood; capsule about 2.9 cm long and 1.0 cm in diameter, fusiform. PHENOLOGY: Flowering January through April with a distinct peak in March and April. DISTRIBUTION AND HABITAT: Known from the Cordillera Central and Cordillera de Talamanca in Costa Rica and the Chiriqui region of Panama. DISCUSSION: Although Vriesea ororien- sis was better represented in herbaria than any other thecophylloid vriesea, the speci- mens available were not sufficient to permit a thorough evaluation of the species. Inten- sive collecting of V. ororiensis in Costa Rica has resulted in a much broader concept of this species than previously existed. Vriesea ororiensis is one of the most abundant and conspicuous thecophylloid vrieseas in Costa Rica, being the dominant Vriesea in mon- tane regions. Vriesea ororiensis is an extremely variable taxon in general robustness as well as in the size, color, and markings of the primary bracts. Within a single large population on Volcan Barba, the primary bracts vary in length from 3. 1 cm to 1 1 .4 cm and their color varies from pink with dark red wavy trans- verse lines to totally crimson. There seems to be a correlation between elongate primary bracts and pink color with wavy transverse lines and, conversely, between short primary bracts with deep crimson coloration. More- over, plants with elongate primary bracts are generally larger, with elongate scapes and inflorescences, whereas the forms with short crimson primary bracts tend to be more compact with shorter scapes and inflores- cences. This initially appeared to be corre- lated with exposure to sunlight; however, field observations revealed the two extremes growing side by side in both exposed loca- tions and deep shade. This range in variation is reflected in most other populations that I sampled (Volcan Poas, Volcan Irazu, Cerro Dasser) but the Barba population is the most variable. Collections from the Cordillera de Talamanca exhibit greater robustness and longer primary bracts than those from other areas. In contrast to this variability in color and size of the scape and primary bracts the flowers are consistently greenish with tubu- lar corollas and diurnal anthesis. This ex- treme variation in a small suite of characters has resulted in several populational or geo- graphical variants previously being recog- nized as distinct species. Examination of the type material and the correlation of the specimens with my field observations show that Mez misinterpreted the inflorescence orientation of V. ororien- sis. The inflorescence of this species is erect at anthesis, and not pendent as he stated in 44 Tulane Studies in Zoology and Botany Vol. 24 Monographiae Phanerogamarum (1896) and Das Pflanzenreich (1934-1935). Smith and Downs (1977) also state that the inflo- rescences of V. ororiensis are pendent. In the case of V. balanophora, I believe this mis- understanding is probably due to the speci- men's, pendent appearance which resulted from bending the scape during collecting or pressing. Aside from this factor, Mez's des- cription of V. ororiensis falls well within the range of variation I have observed in popula- tions of the montane, day flowering taxon which I treat as V. ororiensis. The type mate- rial also agrees well with collections from Volcan Irazu. I doubt that Oersted's collec- tion was made at Orosi as indicated on the label because the elevation at Orosi is ap- proximately 1000 m. This elevation is at least 900 m lower than the lowest elevation at which V. ororiensis has ever been encoun- tered. The Oersted specimen was most likely collected on the slopes of Volcan Irazu, which he is known to have ascended. Mez described Vriesea crassiflora from a Werckle collection that was reportedly made in the vicinity of Cartago. However, exami- nation of the type specimen of V. crassiflora revealed a handwritten notation by Werckle on a field tag indicating that the collection was made in the vicinity of Coliblanco on the slopes of Irazu. Moreover, Werckle included several sketches of the flowers of Vriesea crassiflora at anthesis. These sketches, in conjunction with the character Mez (1934- 1935) and Smith & Downs (1977) used to separate Vriesea crassiflora irom V. ororien- sis, penduncle length (which is variable in V. ororiensis), leave little doubt that this taxon is conspecific with V. ororiensis. Based on his misinterpretation of scape orientation in K oronen^w Mez described V. irazuensis, a species with erect inflorescen- ces. Smith and Downs (1977) also distin- guished V. irazuensis, from V. ororiensis on the basis of scape orientation. Other than this alleged difference in scape orientation, V. irazuensis, agrees in all critical characters with V. ororiensis. Vriesea irazuensis sensu L. B. Smith and Downs (1977), not Mez & Werckle ( 1903), is conspecific with V. neph- rolepis, a night flowering taxon with an erect inflorescence. Vriesea standleyi is well within the range of variability observed in V. ororiensis and con- sequently its maintenance as a distinct spe- cies is unsupportable. When Smith described V. standleyi, he misinterpreted its affinities by considering its closest relationship to be with V. stenophylla, a distinctly different taxon. In 1977, Smith and Downs also separated V. standleyi from V. ororiensis on the basis of a primary bract character. They considered the primary bracts of V. stand- leyi, to be densely pale lepidote, whereas they found those of V. ororiensis to be glabrous and even or verrucose. This type of differ- ence is more a reflection of populational variation than it is of species differences. In 1942 Suessenguth and Goeppinger (in Suessenguth, 1942) described two additional taxa from the Costa Rican collections of Kupper which are also synonyms of V. oro- riensis. The first of these, V. cylindracea, was collected by Kupper near Canaan, appar- ently during his ascent of Cerro Chirripo. The type specimen has 2-flowered lateral branches, subcoriaceous floral bracts and stout pedicels. These characters, in conjunc- tion with the marginal pubescence of the primary bracts (a character that I never observed in species of the pedicellate line), indicate that V. cylindracea is closest to V. ororiensis and V. nephrolepis,ralherihan V. leptopoda (Thecophyllum rubrum) and V. latissima to which it was compared by Sues- senguth. Although Smith and Downs (1977) maintained this taxon on the basis of its primary bract development, it is best treated as a synonym of V. ororiensis, rather than V. nephrolepis, because of its relatively short petals and its primary bracts which are not rugose. The second species described by Suessen- guth and Goeppinger, V. kupperi, was based No.l Middle American Vrieseas 45 on a specimen consisting of the upper part of a scape and an inflorescence. Because of the good color preservation, there is no doubt that the scape and primary bracts were origi- nally a shade of red or pink. When this factor is considered with the size of the sepals and petal size and color, there is no doubt that V. kupperi is conspecific with V. ororiensis. This species was first placed in synonymy under V. ororiensis by Smith (1966). In summary, V. ororiensis may be charac- terized by its erect inflorescence, red, crim- son, or rarely pink primary bracts and its green-yellow, tubular, diurnal flowers. V. ororiensis is extremely difficult to distinguish from V. nephrolepis in fruiting or vegetative condition. This problem is discussed under the latter species. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: Volcan Poas, I96I, //o6Zj5 5.A7. (US); 2-4 km south of Volcan Poas on Hwy. 120, 11 March 1975, Utley & Utley 1930 (CAS), 1931 (M), 1932 (DUKE), 1933 (CR), 1934 (F), 1935 (US), 1936 (MO), 1937 (NOLS), 1938 (MICH), 1939 (MEX), 1940 A (GOET), 1941 (DUKE), 1942 (B), 1943 (WIS), 1944 (K), 1945 (L), 1946 (DUKE) & 1947 {C). CAR- TAGO PROVINCE: Volcan Irazu, 24 Feb- ruary 1957, Carlson 3554 (US); 11 March 1894, Donnell Smith 4963 (US); beyond Cartago, 15 December 1948, Foster 2677 (US); 19 miles south of Cartago on Intera- merican Highway, Talamanca Mountains, 10 July 1962, Haines & Haines 697 (US); 22 miles south of Cartago on Interamerican Highway, north slopes of Talamanca moun- tains, north slopes of Talamanca mountains, 17 July 1962; Haines & Haines 724 (US); 21 miles south of Cartago on Interamerican Highway, 19 July 1962, Haines & Haines 731 (US); 29 miles south of Cartago on Intera- merican Highway, 19 July 1962, Haines & Haines 733 (US); 45 miles south of Cartago on Interamerican Highway, 23 July 1962, Haines & Haines 740 (US); 48 miles south of Cartago on Interamerican Highway, 23 July 1962, Haines & Haines 741 (US); 48 miles south of Cartago on Interamerican High- way, 28 July 1962, Haines & Haines 745 (US); 50 miles south of Cartago on Intera- merican Highway, 30 July 1962, Haines & Haines 755 (US); 41 miles south of Cartago on Interamerican Highway, 30 July 1962, Haines & Haines 757(US); 34 miles south of Cartago on Interamerican Highway, 1 1 August 1962, Haines & Haines 770 (US); 33 miles south of Cartago on Interamerican Highway, 1 1 August 1962, Haines & Haines 772 (US); south slope of Volcan Irazu, on highway about 5 km northeast of Finca Robert, 26 June 1949, Holm & litis 437 (US); upper slopes of Volcan Irazu on road from Cartago to summit, 7 April 1953, Moore 6658 (US); southern slope of Volcan Irazu, from native collector, March 1924, Standley 36631 (US); southern and eastern slopes of Volcan Irazu along Hwy. 8, 23 February 1974, Utley & Utley 647 {D\}Y.E, US, CR); 12 March 1975, Utley & Utley 1953 (CR), 1954 (DUKE), 7955 (F), 1956 (CR); on old road off of the Interamerican Highway north of El Empalme, 16 April 1975, Utley & Utley 2174 (DUKE), 2176 (CR); the southern slopes of Volcan Irazu on farm road leading from Hwy. 8 to Finca Coliblanco, 25 January 1976, Utley & Utley 3815 A&B (DUKE, CR), 3816 A&B (US, DUKE), 2817 A&B (F, MO), 3818 (GH), 3819 (NOLS), 3820 (MlCHl 3821 A.B&C (DUKE, US, B), 3824 (K), 3825 (DUKE), 3826 (WIS), 3827 (CAS), 3828 (NY), 3829 (DUKE), 3830 5 (NOLS, CR, MEX), 3831 (US) & 3832 (F); southern slope of Volcan Turrialba, 15 March 1976, Utley & Utley 4309 (US), 4310 (DUKE), 4311 (CR), 4312 (DUKE), 4i7i (NOLS), 4i/4 (US), 4i75(F), 4316 (MO), 4'i77 (MICH), 4318 (GH), 4319 (DUKE), 4322 (IJSF), 4323 (NO), 4324(F), 4325 (DUKE), 4326 (B), 4327 (NY), 4328 (MO), 4329 (NOLS), 4330 (DUKE), 4331 (CAS), 4332 (BR) & 4333 (DUKE); cloud forest area between the paramillo of Cerro de la Muerte and Villa Mills, Cordillera de 46 Tulane Studies in Zoology and Botany Vol. 24 Talamanca, 26 February 1965, Williams et al. 28337 (US). BORDER OF CARTAGO AND SAN JOSE PROVINCES: 2-3 km west of Villa Mills on the Interamerican Highway, 17 March 1975, Utley & Utley 1975 (DUKE), 1976 (CR), 1977 (US), /979(NOLS), 1980 (BM), 1981 (F), 1983 (MO), 1984 (MICH), 1985 (GH), 1986 (CAS), 1988 (NY), 1989 (WIS), 1990 (PMA), /997(C), 7992 (L); 27 April 1975, Utley & Utley 2266 (DUKE);2-3 km west of Villa Mills on the Interamerican Highway, 5 April \976,Utley & Utley 4483 (DUKE) & 4484 (CR); at Villa Mills on the Interameri- can Highway, 5 April 1976, Utley & Utley 4487 (US), 4488 (NOLS), 4489 (MICH), ^^90(DUKE), 4491 (MO) & 4492 (DUKE); between Salispuedes and Ojo de Agua on the Interamerican Highway south of Cartago, 5 April 1976, Utley & Utley 4496 {CR)& 4497 (DUKE). HEREDIA PROVINCE: summit of Volcan Barba within 1 km of Laguna Barba, 16 March 1975, Utley & Utley 1961 (NOLS), 79(52(DUKE),796i (US), 7955(F), 1966 (DUKE), 1967 (NY), 79(5<^(MO), 79<59 (CAS), 7970 (MICH), 7977 (DUKE), & 7972 (NOLS); 12 April 1975 Utley & Utley 2013 (DUKE), 207^ (CR), 2015 (US) 2023 (F), 2024 (MEX), 2039 (MO) & 2042 (DUKE); 25 January 1976, Utley & Utley 3706 (DUKE), 3730 (CR), 3731 (DUKE); 5 March 1976, Utley & Utley 4210 (DUKE), 4211 (CR), ^27i (DUKE), ^27^ (F), ^275 (MO), 4216 (NOLS), 4217 (MICH), 4218 A&B(GH, CAS), ^279 (MEX), 4220 (NY), ^227 (WIS), 4222 (DUKE), 4223 (C), 4224 (M), 4225 (MICH), 4226 (PMA), 4227 (DUKE), 4228 (F), 4229 (DUKE), 4230 (DUKE), 4232 (CAS), 4233 (NY), 4234 (CAS), 4235 (HEID), 4236 (DUKE), 4237 (MICH), 4238 (F), 4239 {VS),4240{DVKE), 4255 (CR), 4256 (F), ^257 (DUKE); 9 April 1976, Utley & Utley 4578 (NOLS), 4580 (DUKE) & ¥5.^7 (CR). SAN JOSE PRO- VINCE: on tree trunk north of San Jose , 5 March 1966, Smith & Berry 15316 (US); on tree cloud forest, Aserri, 19 April 1966, Smith. LB. 15327 (US); Cedral (Cerros de Escazu), approach to and upper slopes of Cerro Daser 3-5 km southwest of Aserri, 30 April 1975, Utley & Utley 2301 (DUKE), 2302 (DUKE), 2303 (CR), 2304 (DUKE) & 2305 (DUKE); 13 March 1976, Utley & Utley 4261 (DVKE), 4262 (CR), 4267 A & B (US, DUKE) 4268 (F), 4269 (MO), 4270 (MICH), 4272 iVS),4273 (NOLS), 4274 (DUKE). 4275 (CR), 4276 (NY), 4277 (C AS), 4278 (GH), 4279 (MO). PAN AM A:- CHIRIQUI PROVINCE: Las Nubes, 7 August, Croat 26471 (MO). 11. Vriesea balanophora (Mez) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Guzmania balanophora Mez in DC. Mon. Phan. 9:918. 1896. TYPE: COSTA RICA. Oersted Brom. n. 22. (HOLOTYPE: C!, photograph US!). Thecophyllum balanophorum (Mez) Mez, Bull. Herb. Boiss. ser. II. 3: 131. 1903. Thecophyllum lineatum Mez & Werckle, Bull. Herb. Boiss. ser. II. 4: 875. 1904. TYPE: COSTA RICA: Werckle "Bromel. Costaric." 114 and 16,260. (LECTOTYPE: Werckle 114, B\, pho- tograph US!; SYNTYPE: Werckle Inst. Phys. -Geogr. Costar. 16260. B!, photograph US!). Thecophyllum balanophorum var. subpictum Suesseng. Bot. Jahrb. 72: 291. 1942. TYPE: COSTA RICA. Kupper 257. (HOLOTYPE: M!, photograph US!). Vriesea lineata {Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Epiphytic or rarely terrestrial, acaulescent to 20.0 cm caulescent. LEAVES in a sub- No.l Middle American Vrieseas 47 speading to subascending rosette, (19.0) 22.0-31.2 cm long; blades 12.0-23.0 cm long and 1.8-2.9 cm wide, green with maroon wavy cross bands and occasionally with maroon longitudinal lines proximally, sub- ligulate to broadly triangular, apex acute to acuminate or attenuate; sheaths (6.8) 7.6-9.8 cm long and 3.2-5.0 cm wide, castaneous, usually darker abaxially, with a coating of cinereous trichomes and frequently long- itudinal maroon lines distally, elliptic or rarely obovate. SCAPE erect or curved-erect (26.0) 33.0-40.0 (48.0) cm long and 0.2-0.3 cm in diameter; bracts 4.0-10.0 (17.7) cm long and 0.4-2.1 cm wide, green or green suffused with maroon, usually with maroon wavy transverse lines and a covering of cinereous trichomes, clasping-erect in sheath area becoming spreading in the area of blade, triangular to subligulate, apex acute and attenuate. INFLORESCENCE erect, 3.0-6.0 (9.0) cm long and 3.0-7.0 cm in diameter, cylindric, glabrous occasionally with brown-margined punctulae; primary bracts 3.0-6.2 cm long and 1.4-2.0 (2.8) cm wide, spreading to subascending, green with maroon wavy cross-bands, occasionally suf- fused with red or maroon, triangular, apex acute to acuminate or attenuate. LATER- AL BRANCHES subspreading or spread- ing, collaterally 2-flowered, peduncles 0.2- 0.5 cm long. FLOWERS sessile to less than 0.1 cm pedicellate; y7ora/ bracts 0.9-1.2 cm long and 1.0-1.5 cm wide, subcoriaceous green frequently suffused with maroon or completely maroon, drying to brown,surface finely nerved, minutely pitted or brown- margined punctulate, keeled, broadly elliptic to suborbicular, apex rounded to obtuse, frequently incised; sepals 1 .0-1 .3 cm long and 0.8-1.1 cm wide, green suffused with maroon, entirely maroon or castaneous, coriaceous, elliptic, apex rounded, frequently minutely incised; petals 1 .9-2.0 cm long and 1 .0- 1 . 1 cm wide, white more or less obovate; appendages 0.3-0.4 cm long and 0.2 cm wide; stamens in hood configuration over the stigma, anthers not laterally connivent, fila- ments 1.2-1.4 cm long, anthers 0.4 cm long; ovary 0.3-0.4 cm long and 0.3-0.4 cm in diameter, style about 1.2 cm long; capsule 1.7-2.5 cm long and 0.7-1.0 cm in diameter, subfusiform to obovate, maroon becoming castaneous when mature. PHENOLOGY: V. balanophora has been collected in flower during the months of August and December. That these two months represent a sample of a flowering season extending through the heavy rain months of September, October and Nov- ember is difficult to believe. Rather, the spe- cies probably flowers sporadically through- out the "dry" season of January through April (May) and the early rainy season of JuneJuly. DISTRIBUTION AND HABITAT: The species is endemic to Costa Rica where it is known from the volcanos of the Cordillera Central and the northern extension of the Cordillera de Talamanca. The species has been collected between 1500-1900 m in the lower montane and premontane rain forest life zones and possibly extends slightly into the lower montane wet forest life zone. DISCUSSION: V. balanophora is easily distinguished from other vrieseas by its cinereously pubescent leaves with wavy maroon transverse lines and erect inflore- scences bearing white, crepuscular flowers with campanulate corollas. Although the type of V. balanophora is poorly preserved, there is no doubt that V. lineata is conspecific with it. Judging from the type, Mez's interpretation of the or- ientation of the scape apparently was in error. At first glance, the scape appears pen- dulous but this is a result of its having been broken and bent to facilitate mounting or pressing. In its natural state, it was obviously erect or curved-erect. Suessenguth and Goeppinger (1942) described T. balanopho- rum var. subpictum on the basis of its shorter leaves which were suffused with red and red 48 Tulane Studies in Zoology and Botany Vol. 24 wavy crossbands. However, the type of T. balanophorum var. subpictum is well within the range of variation that I have observed in V. balanophora and is undeserving of recog- nition at the varietal level. Vriesea balanophora is most similar vegeta- tively to V. leucophylla. Because of their overlapping distributions and almost identi- cal leaf pubescence and markings, the taxa are frequently difficult or impossible to separate in the vegetative condition. In flower or fruit the species are easily separated by a number of characters, the most con- spicuous of which is inflorescence orienta- tion. Inflorescences of V. balanophora are erect, in contrast to the pendent inflorescen- ces of V. leucophylla. The primary bracts of the former taxon are also shorter than those of the latter species. Moreover, the floral syndromes of these species are different. As I indicated prevously, V. balanophora has campanulate, crepuscular flowers; this dif- fers markedly from the chartreuse, more or less tubular, diurnal flowers characteristic of V. leucophylla. Vriesea balanophora may be most closely related to V. luis-gomezii, but their relation- ship cannot be fully interpreted until more material of this latter poorly known taxon is available. These two species share a number of characters including nocturnal or crepus- cular white, campanulate flowers and leaves with cinereous pubescence and wavy trans- verse lines. The two taxa are currently separ- ated on the basis of the longer floral bracts (1.5-1.7 cm) and sepals (1.5-1.6 cm) of V. luis-gomezii as opposed to the shorter floral bracts (0.9-1 .2 cm) and sepals ( 1 .0-1.3 cm) of V. balanophora. SPECIMENS EXAMINED: COSTA RICA: PROVINCE OF ALAJUELA: about 3-4 km west of the junction of high- ways 9 and 120 on Highway 120; the saddle between Volcan Poas and Volcan Barba, 27 March 1976, Utley & Utley 4338 (NOLS). BORDER OF ALAJUELA ANDHEREDIA PROVINCES: on the finca of Mike Canon at Vara Blanca, near the junction of high- ways 9 and 120, 20 July 1975, Utley & Utley 2661 (DUKE). CARTAGO PROVINCE:- Tapanti Hydroelectric Project, slopes over- looking the Rio Grande de Orosi about 5 km beyond the entrance to the Project, 22 March 1976, Utley & Utley 4354 iDVKE)& 4355 (NY). BORDER OF CARTAGO AND SAN JOSE PROVINCES: along the Interamerican Highway southeast of Casa Mata or 10-12 km south of San Isidro de Cartago, 17 March 1975, Utley & Utley 2000 (US), 2001 (DUKE) & 2002 (CR); 3 August 1975, Utley & Utley 2756 (CAS) & 2757 (DUKE); 5 September 1975, Utley & Utley 3086 (DUKE). HEREDIA PROVINCE: Cerro de Las Lajas, north of San Isidro, 7 March 1926, Standley & Valerio51496(VSy, Cerro de Las Caricias north of San Isidro, 1 1 March 1926, Standley & Valerio 52325 (VS); region north of Cerro Chompipe about 15 km northeast of Heredia and 3 km north of Cerro Redondo de La Cruz, 15 October 1974, Utley & Utley 1312 (DUKE). SAN JOSE PROVINCE: region between Casca- jal and about 8 km northeast on Highway 216, 23 August 1975, Utley & Utley 2917 (DUKE), 2918 (CR) & 2945 (NOLS); 10 December 1975, Utley & Utley 3601 (MO), i679(F)&i(520(DUKE);AhosdeTablazo, about 5 km southeast of Higuito on Calle Tablazo, or about 12 km southeast of Desamparados, Utley & Utley 3871 (DUKE)&5<^72(NOLS);9 May 1976, Utley & fy//^v^7i6(DUKE);4kmnorthofCasca- jal or 7 km north of Las Nubes on Hwy 216 near the road terminus, 3 July 1976, Utley & Utlev 5263 (VS), 5269 iDVKE)8i 5270 {CR, F). 12. Vriesea luis-gomezii Utley, sp. nov. Fig. 21. A V. balanophora (Me?) L. B. Smith & Pitt, cui valde affmis, bracteis florigeris et sepalis longioribus differt. No.l Middle American Vrieseas 49 Epiphytic acaulescent to short caulescent. LEAVES in a spreading to subascending rosette, 33.6-43.2 cm long; blades 17.3-27.3 cm long and 3.5-4.0 cm wide, red suffused with areas of green, and with reddish-purple or dark wavy transverse lines, occasionally with longitudinal reddish-purple lines prox- imally, ligulate, apex acute to acuminate or attentuate; sheaths 14.0-16.6 cm long and 6.6-7.1 cm wide, dark castaneous abaxially, brown adaxially, frequently suffused with purple-maroon wavy transverse lines dis- tally, elliptic in outline. SCAPE erect, 48.5- 62.0 cm long and 0.4-0.5 cm in diameter; bracts 5.9-9.1 cm long and 2.8-3.2 cm wide, sheath and blade not strongly differentiated, blades green or green deeply suffused with red and with red wavy transverse lines prox- imally, erect, divergent distally, triangular, apex acute to attenuate, sheaths green with red longitudinal lines proximally and occa- sionally with wavy transverse lines distally, sheathing erect, ovate to elliptic. INFLO- RESCENCE erect, 9.0-1 5.5 cm long and 4.5- 7.0 cm in diameter, cylindric, rachis red; primary bracts 5.0-8.2 cm long, blade and sheath frequently weakly differentiated, blades about 2.7-3.9 cm long and 1.8 cm wide, green frequently with wavy red trans- verse lines proximally, spreading to recurved or occasionally spirally involute, triangular, apex acute to attenuate, sheaths about 2.3- 3.2 cm long and 2.7-3.2 cm wide, green with red longitudinal lines proximally or red wavy transverse lines distally, ovate. LAT- ERAL BRANCHES spreading, to 2.5 cm long, collaterally 2-flowered peduncle 0.2- 0.4 cm long, rachis wanting to 0.2 cm long. FLOWERS sessile to 0.3 cm pedicellate; floral bracts 1.5-1.7 cm long and 1.7-1.9 cm wide, maroon or green suffused with maroon marginally, subcoriaceous, brown- centered lepidote adaxially, glabrous abax- ially, even or keeled, ovate or oblong-ovate to suborbicular, apex obtuse to broadly rounded, occasionally incised or torn medially; sepals 1.5-1.6 cm long and 1.0-1.2 cm wide, maroon or green, coriaceous, brown-centered lepidote adaxially, glabrous abaxially, elliptic incised; petals about 2.9 cm long and 1 . 1 - 1 .2 cm wide, white, obovate, appendages 0.4 cm long and 0.2 cm wide; stamens in hood configuration over the gynoecium, filaments about 2.1 cm long, anthers 0.6 cm long; ovary 0.5 cm long, style 2.7 cm long. TYPE: COSTA RICA: CARTAGO: on foot or beast path off the old Pan American Highway about 1 -2 km north of El Empalme in the Cordillera de Talamanca, elev. 2000- 2100 m, 10 May 1975, Utley & Utley 2460 (HOLOTYPE: DUKE!, ISOTYPES: CR!, F!, DUKE!, US!). PHENOLOGY: Flowering specimens col- lected in May. DISTRIBUTION AND HABITAT: Known only from the type location in Costa Rica at 2000-2100 m in the lower montane rain forest life zone. DISCUSSION: Vriesea luis-gomezii is a poorly known but distinct species, easily recognized on the basis of its green leaves with longitudinal and transverse red-purple to dark red lines and spreading to sub- spreading rosette. The floral bracts which are about equal in length to the maroon sepals, are maroon or green suffused with maroon. Flowers of this taxon are white, campanu- late and nocturnal or crepuscular. Since this taxon is only known from a few collections, any definitive analysis of its affinities must await additional field work. However, V. luis-gomezii appears to be most closely allied to V. balanophora. This is presented in greater detail under the dis- cussion of V. balanophora. ETYMOLOGY: This species is named in honor of Luis Diego Gomez P., pteridologist and former Director of the Museo Nacional de Costa Rica. SPECIMENS EXAMINED: COSTA RICA: CARTAGO PROVINCE: on horse trail northeast of the old Pan 50 Tulane Studies in Zoology and Botany Vol. 24 American Highway 1-2 km north of El Empaime, 16 April 1975, Utley & Ut ley 2184 (NOLS). 13. Vriesea viridis (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953. Thecophyllum viride Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 872. 1904. TYPE: COSTA RICA: near Desengano, Werckle s.n. (HOLOTYPE: B!, pho- tograph US!). Epiphytic or terrestrial, acaulescent or cau- lescent to 20.0 cm. LEAVES in a subspead- ing to subascending rosette, (28.0) 31.0-46.0 (63.0) cm long; blades 16.8-33.0 (46.5) cm long and (3.1) 4.0-5.7 cm wide, green occa- sionally suffused with maroon or red- maroon at the apices and margins, with fine Fig. 21 . Vriesea luis-gomezii. A. Inflorescence; B. Leaf; C. Flower D. Nearly mature fruit with sepals and floral bract. From (Ulley & Utley 2467). No.l Middle American Vrieseas 51 wavy, dark green cross bands, these occa- sionally suffused with red-maroon or en- tirely red-maroon, ligulate to subligulate or broadly triangular, apex acute to acuminate or attenuate; sheaths (9.6) 12.4-18.0 (20.0) cm long and (4.4) 6.0-8.4 cm wide, pale white with a layer of cinereous trichomes on the upper and lower surfaces, frequently with an abaxial castaneous area or rarely both surfa- ces castaneous, elliptic to obovate. SCAPE erect to curved-erect (27.0) 36.0-60.0 cm long and 0.3-0.6 cm in diameter; bracts erect to spreading or the uppermost recurved, 5.5- 15.0(19.0)cm long, blades 3.0-10.4(15.2) cm long and (0.9) 1.2-2.5 (3.0) cm wide, lower- most colored like leaves, middle and upper with red-maroon transverse lines and fre- quently suffused with red-maroon, triangu- lar to broadly triangular or ligulate, apex acute to acuminate orattenuate, sheaths 2.1- 4.0 (4.8) cm long and 2.0-4.2 cm wide, red- maroon becoming yellow near junction with blade but occasionally ochre or entirely red- maroon, ovate or elliptic to obovate or sub- orbicular. INFLORESCENCE erect, 11.0- 31.0 cm long and 4.5-7.5 cm in diameter, cylindric, rachis glabrous occasionally with dark castaneous punctulae; primary bracts 5.0-10.0 ( 1 3.0) cm long, spreading or reflexed at the blade-sheath junction, blades (2.4) 3.0- 7.0 (9.7) cm long and 0.8-2.2 (2.8) cm wide, green to pink, red or maroon, with wavy cross bands, these usually suffused with red or maroon, triangular to broadly triangular, apex acute to attenuate, sheaths (2.2) 2.8-3.6 cm long and (2.2) 2.6-3.6 cm wide, red or red-maroon frequently becoming yellowish near the junction with blade, drying to ochre or brown, elliptic to broadly elliptic or ovate. LATERAL BRANCHES spreading, collat- erally 2-flowered, peduncles 0.0-0.4 (0.9) cm long, rachis wanting. FLOWERS 0.3-1.3 cm pedicellate;y7ora/^rac/5 0.5-l.l cm long and 0.6-1.3 cm wide, green usually suffused with maroon, subcoriaceous, keel frequently inflated or creased, surface even to obscurely nerved or pitted, with castaneous to light brown punctulae especially basally and on the keel, ovate to elliptic, apex acute or acuminate to obtuse or broadly rounded, frequently torn or appearing incised; sepals 1.1-1.6 cm long and 0.8-1.3 cm wide, green suffused with maroon, or completely maroon, coriaceous, elliptic to obovate, apex rounded to obtuse, frequently emargi- nate or torn-incised; petals 1.9-2.5 cm long and 1 . 1 - 1 .4 cm wide, white to greenish-white, obovate to elliptic; appendages 0.4-0.5 cm long and 0.2 cm wide; stamens arranged in a hood over the gynoecium, anthers laterally connivent, filaments 1 .9-2. 1 cm long, anthers 0.4-0.5 cm long; ovary 0.4-0.5 cm long and 0.3-0.5 cm in diameter, style 1 .5-1.8 cm long; capsule 3.0 cm long and 0.8 cm in diameter, castaneous subfusiform to obovate. PHENOLOGY: Flowering specimens have been collected during March and April. DISTRIBUTION AND HABITAT: Endemic to the vicinity of Volcan Barba in the Cordillera Central de Costa Rica between 1800-2100 m in the lower montane rain forest life zone. DISCUSSION: V. viridis is a narrow endemic which can be locally abundant and even weedy in some areas. It is distinguished by an erect inflorescence, maroon sepals and white, crepuscular flowers in which the corollas are geniculate-tubular. This taxon is most similar to V. balanophora and V. luis-gomezii from which it is dis- tinguished by floral bract and inflorescence length as well as its geniculate-tubular corolla. In sterile or fruiting condition, this species can be confused with both V. oro- riensis and V. nephrolepis from which it can generally be distinguished by the cinereous pubescence of the abaxial leaf surface as well as the tendency for the leaves to be broad triangular in outline, in contrast to the ligu- late or subligulate leaf blades of the other two taxa. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: between 52 Tulane Studies in Zoology and Botany Vol. 24 the junction of highways 9 and 120 and the summit of Volcan Poas, 11 March 1975, Utley & Utley 1915 (CR), 1916 (GH), 1917 (DUKE), 1919 (NOLS), 1920 (K), 1923 (CR),7925(F)«&792(5(DUKE).HEREDIA PROVINCE: region north of Cerro Chom- pipe, about 15 km northwest of Heredia and 3-5 km north of Cerro Redondo de La Cruz, 15 October 1974, Utley & Utley 1292 (NOLS); 13 April 1975, Utley & Utley 2086 (CR), 2089 (DUKE), 2090 (NOLS), 2091 (US), 2092(F), 2093 (DUKE) & 209^ (MO); about 1-3 km south of the junction of high- ways 9 and 120, on slopes above the Rio Desengano, 11 May 1975, Utley & Utley 2481 (DUKE) &. 2483 (US); road north of Hwy. 113 connecting with Calle Gallito, saddle area between Cerro Chompipe and the southeast flank of Volcan Barba, 25 October 1975, Utley & Utley 3201 (DUKE); 8 December 1975, Utley & Utley 3580 (DUKE); 2-3 km south of the junction of highways 9 and 1 20 on Hwy. 9, slopes above the Rio Desengano, 27 March 1976, Utley & Utley 4377 (CR), 4380 (NOLS), 4381 (US), 4388 (F), 4389 (MO), 4390 (NY), 4397 (CAS), 4398 (DUKE) 4399 (B), 4400 (CR), 4401 (MICH), 4402 (MO), 4403 (DUKE), 4404 (DUKE) 4405 (US), & 4408 (MEXU); approach to the southeastern flank of Vol- can Barba on the southwest side of Cerro Chompipe, 1 April 1976, Utley & Utley 4421 (CR), 4422 (DUKE), 4423 (DUKE), 4425 (NY), 4426 (CAS), 4427 (US), 4428 (BM), 4429 (MO), 4430 (DUKE), 4431 (NOLS), 4437 (DUKE), 4438 (DUKE), 4440 (F), 4441 (US), 4442 (GH), 4443 (DUKE), 4444 (CR), 4445 (CAS), 4446 (B), 4447 (DUKE). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Heredia along the banks of the Rio Para Blanco and on the lower slopes of Cerro Zurquf, 30 March 1974, Utley & Utley 736 (DUKE), 737 iVS), 81.750 A (CR); 21 October 1975, Utley & Utley 3177 (DUKE); about 5 km northeast of San Isidro de Heredia on Calle Yerbabuena, 3 April 1976, Utley & Utley 4476 (DUKE). 14. Vriesea hainesiorum L. B. Smith, Phyto- logia 8: 498. 1963 TYPE: COSTA RICA: CARTAGO PROVINCE: Interamerican Highway, 6 miles south of Cartago, Talamanca Mountains, elev. 1650 m, Haines & Haines 682 (HOLOTYPE: US!). Epiphytic acaulescent. LEAVES in an ascending rosette, 14.0-21.0 (27.0) cm long; Zj/ac/e5 9.0-1 3.0 (19.0) cm long and 0.8-1. 4 cm wide, green, narrowly triangular, apex attenuate; sheaths (3.5) 4.5-6.5 (8.0) cm long and 2.2-3.4 cm wide, green densely cinere- ously pubescent, margins purple, dark maroon or castaneous, elliptic. SCAPE decurved, usually crimson, (16.0) 20.0-25.0 cm long and about 0.1-0.2 cm in diameter; bracts divergent to spreading (3.5) 4.2-8.4 (10.1) cm long, blades 1.5-8.0 cm long and 0.2-0.6 cm wide, green or green suffused with red, occasionally crimson, narrowly triangu- lar to setiform, apex attenuate to long acum- inate, sheaths 1 .8-3.0 cm long and 0.9-1 .4 cm wide, green suffused with red to crimson, clasping erect, elliptic to ovate. INFLO- RESCENCE pendent, 3.0-6.0 cm long and 3.0-7.0 cm in diameter, more or less cylin- dric, rachis geniculate, crimson, primary bracts spreading 3.4-5.3 (6.5) cm long, blades 1.4-3.2 cm long and 0.2-0.7 cm wide, green suffused with red to crimson, drying to brown, narrowly triangular to setiform, apex attenuate. LATERAL BRANCHES divergent to spreading, 0.5-0.7 cm long, col- laterally 2-flowered, peduncles 0.2-0.3 cm long, rachis 0.2-0.3 cm long. FLOWERS sessile to 0.2 cm pedicellate; Jloral bracts (1.3) 1.5-1.8 cm long and 1.4-1.9 cm wide, pale green drying to brown, subcoriaceous to coriaceous, finely nerved with scattered punctulae, keel prominently folded or in- flated, elliptic to ovate or suborbicular, apex obtuse to rounded, frequently torn-incised; No.l Middle American Vrieseas 53 sepals 1.9-2.1 cm long and 1.1-1.4 cm wide, pale green, drying to brown, coriaceous, elliptic to obovate, apex obtuse to rounded, frequently torn-incised; petals 2.7-2.8 cm long and 0.9-1.0 cm wide, greenish-yellow, obovate; appendages 0.4 cm long and 0.15- 0.2 cm wide; stamens in a hood config- uration over gynoecium, filaments about 2.0 cm long, anthers 0.3-0.35 cm long; ovary 0 A cm long and 0.3 cm in diameter, style about 1.9 cm in diameter, green becoming cas- taneous. PHENOLOGY: Flowering specimens have been collected during the month of May. DISTRIBUTION AND HABITAT: Endemic to the northern extension of the Cordillera de Talamanca in Costa Rica. Col- lected between 1700-1900 m in the lower montane wet and lower montane moist forest life zone. DISCUSSION: Vriesea hainesiorum is a distinctive taxon which is characterized by narrowly triangular leaves with attenuate apices borne in an ascending rosette. Inflo- rescences are decurved to pendent and highly colored. Both the scape and primary bracts are deep red to crimson, and the flowers are diurnal, greenish yellow and tubular. V. hainesiorum is most closely related to V. leucophylla, from which it is easily distin- guished by its ascending rosette, unmarked, short, narrowly triangular leaves and few- flowered inflorescence (ten or fewer flowers). The relationship of V. hainesiorum to V. leucophylla 's discussed in greater detail under V. leucophylla. SPECIMENS EXAMINED: COSTA RICA: CARTAGO: 5-6 km south of San Isidro de Cartago on the Pan American Highway, 1700-1800 m, 28 February 1975, Utley & Utley 1908 (DUKE). SAN JOSE: on road to Tablazo ca. 1-2 km from crest of Altos de Tablazo or 5-6 km SE of Higuito or 12-14 km SE of Desamparados, 1700-1800 m, 5 September 1975, Utley & Utley 3062 (CR), 3063 (US), 3064 (DUKE) & 3065 (NOLS); 30 January 1976, Utley & Utley 3864 (DUKE), 3865 (MO), 3866 (NY) & 3557 (F); 9 May 1976, Utley & Utley 47 38 A (NOLS), 4741 (CR), 4742 (CAS), 4780 (GH),^7«7(F)&^7«5(DUKE):2Julyl976, Utley & Utley 5205 (DUKE). 15. Vriesea leucophylla L. B. Smith, Phyto- logia 8: 498. 1963 TYPE: COSTA RICA: CARTAGO PROVINCE: rock outcrop 11 miles south of Cartago, on Interamerican Highway, Talamanca Mountains, elev. 1860 m, Haines & Haines 693 (HOLOTYPE: US!). Vriesea leucophylla var. subtessellata Utley, Journ. Bromeliad Society 24: 16-18. 1974. TYPE: COSTA RICA: SAN JOSE PROVINCE: southern slopes of Cerro Zurqui, 4.0-4.5 km north of San Isidro de Heredia, alt. 1800 m. John & Kathy Utley 401 (HOLO- TYPE: DUKE!, photographs CR!, US!). Epiphytic or rarely terrestrial, acaulescent. LEAVES in a spreading to subascending rosette, 21.0-41.0 cm long; blades 13.0-29.0 cm long and 2.1-2.8 cm wide, green with dark green or maroon wavy transverse lines, ligulate to long triangular, apex attenuate to acute; sheaths 7.5-15.0 cm long and (3.9) 4.6-5.7 cm wide, pale brown to brown with a mosiac-like cover of brown to castaneous centered trichomes, usually suffused with purple or maroon wavy transverse lines in the distal portion, elliptic to ovate or sub- ovate. SCAPE strongly decurved, 23.0-44.0 cm long and 0.3-0.5 cm in diameter; bracts erect, divergent distally, 6.0-18.0 cm long, blades 3.5-14.5 cm long and 0.8-2. 1 cm wide, lowermost green and foliaceous, middle and upper suffused deep red-pink to crimson, usually with dark red or crimson wavy trans- 54 Tulane Studies in Zoology and Botany Vol. 24 verse bands, subligulate to long triangular, apex acute to attenuate, sheaths clasping- erect, 2.2-4.2 cm long and 1.5-3.0 (3.3) cm wide, deeply suffused red-pink to crimson, ovate to elliptic in outline. INFLORES- CENCE pendent (12.0) 20.0-34.0 cm long and 5.0-14.0 (19.0) cm in diameter, cylindric, to more or less conical, rachis red or crim- son; primary bracts strongly spreading, geniculately reflexed at the blade-sheath junction, 6.0-13.5 cm long, occasionally with wavy transverse bands, subligulate to long triangular, apex acute to attenuate, sheaths 2.4-3.4 cm long and (1.9) 2.6-3.8 cm wide, deep red to crimson, ovate in outline. LATERAL BRANCHES spreading, 1.02.2 cm long, 2-flowered, or rarely 3-flowered, penduncles 0.7-1.5 cm long, rachis 0.2-0.5 cm long, flowers borne subcollaterally or if spikes 3-flowered then apparently pinnate. FLOWERS sessile to 0.3 cm pedicellate; floral bracts 1 .4-2.0 cm long and 1 .8-2.6 cm wide, subcoriaceous, green or rarely suffused with light pink, brown punctate or immersed lepidote abaxially, lepidote abaxially, oblong to elliptic or ovate, apex obtuse to broadly rounded and minutely apiculate, entire to deeply torn-incised along the con- spicuous, frequently inflated keel; sepals 1 .5- 1 .9 cm long and 0.9- 1 .5 cm wide, coriaceous, usually keeled, glabrous abaxially, lepidote adaxially, green to yellowgreen or strongly suffused with red at anthesis, elliptic to obo- vate, apex obtuse to rounded, occasionally torn incised; petals 2.4-2.9 cm long and 0.8- 0.9 cm wide, greenish, elliptic to obovale, frequently subpandurate; appendages (0.2) 0.5-0.6 cm long and 0. 1 -0.2 cm wide; stamens in a hood configuration over gynoecium, filaments 1.8-2.1 cm long, anthers 0.4-0.6 cm long; ovaryQA-0.5 cm long and 0.3-0.4 cm in diameter, style 1.6-1.8 cm long; capsules 3.7- 4.0 cm long and about 0.9 cm in diameter, olive-brown, fusiform. DISCUSSION: The features that charact- erize Vriesea leucophylla are its pendent. multiflowered inflorescence (more than 14 flowers) with red to crimson scape and prim- ary bracts and floral bracts which are about as long as, or only slightly shorter than the sepals. The flowers of this taxon are diurnal, green-yellow and tubular. Vriesea leucophylla is most closely related to V. hainesiorum and V. vietoris. It differs from the former species in leaf size and shape. The relationship of V. leucophylla to V. vietoris is discussed under the latter taxon. DISTRIBUTION: The Cordillera Central of Costa Rica and the Cordillera de Tala- manca in Costa Rica and northwest Panama in the lower montane rain forest. PHENOLOGY: Flowering June-August. SPECIMENS EXAMINED: COSTA RICA: BORDER OF ALAJUELA AND- HEREDIA PROVINCES: Vara Blanca, the fmca of Mike Canon, near the junction of highways 9 and 120, 1900 m, 20 July 1975, Utley & Utley 2658 (F) & 2660 (DUKE). CARTAGO PROVINCE: 6 miles south of Cartago on the Interamerican Highway, Talamanca Mountains, 5500 feet, 10 July 1962, A. Lee & Bruce L. Haines 679 (CR, US); region north of Muneco or about 5.5 km west of Orosi, 1600m, 18 October 1974, Utley & Utley 1440 (DUKE). BORDER- OF CARTAGO AND SAN JOSE PRO- VINCES: about 20 km south of Cartago on the Interamerican Highway in the vicinity of Casa Mata, 1900 m, 17 March 1975, Utley & Utley 7999 (DUKE), 3 August 1975, Utley & Utley 2758 (US), 2759 (DUKE) «fe 2761 (NOLS), 24 August 1975, Utley & Utley 2951 (DUKE). HEREDIA PROVINCE: vicinity of Vara Blanca in pastures along CR. highways 9 and 120, 1900 m, 21 June 1976, Utley & Utley 5146 (VS), 5147 {CRl 5148iNOLSl5149{F),5150iDVKE)J151 (MO) & 5152 (DUKE). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San IsLdro de Here- dia along the banks of the Ri'o Para Blanco No.l Middle American Vrieseas 55 and on the lower slopes of Cerro Zurqui, 1500-1700 m, 10 June 1974, Utley & Utlev 933 (CR) & 955 (DUKE); 18 June 1974, Utley & Utley 967 (DU KE), 962 (US) & 965 (DUKE); 22 December 1974, Utlev & Utlev 1749 (F), 1750 (DUKE), 1751 (US) & 1752 (CR); 12 July 1975, Utley & Utley 2613 (US) & 2614 (DUKE); 21 October 1975, Utley & Utley 3176 (DUKE); 27 December 1975, Utley & Utley 3665 (DUKE). SAN JOSE- PROVINCE: Alto de la Palma, on Finca Porvenir, 18 August 1975, Utley & Utley 2912 (DUKE); between 2 and 5 km sou- theast of Higuito on Calle Tablazo or between 8 and 12 km southeast of Desampa- rados, 1800-1900 m, 5 September 1975, Utley & Utley 3066 A &BiNOLS, DUKE). PANAMA: CHIRIQUI PROVINCE: Bajo Chorro, 13 February 1938, Davidson 281 (F). 16. Vriesea vietoris Utley, Brittonia, 33: 584- 586.1981. Fig. 22 TYPE: COSTA RICA: BORDER OF CARTAGO AND SAN JOSE PROVINCES: pasture along the Inter- american Highway, 200 yards SE of Casa Mata in the Cordillera de Tala- manca S of Cartago, 1800-1900 m Utley & Utley 4747 (HOLOTYPE: DUKE!) Plants acaulescent, epiphytic. LEAVES in a spreading to subascending rosette, 29-40 cm long; blades 19-29 cm long and 1.4-2.6 cm wide, green, occasionally suffused with maroon at margins and tips, densely pubes- cent on abaxial surface, subligulate to long- triangular apex attenuate or acuminate; sheaths 10-12.2 cm long, 3.4-5.2 cm wide, brown with a dense mosaic-like coating of castaneous-centered trichomes usually suf- fused with purple in the distal half, especially near the margins, elliptic to ovate in outline. SCAPE strongly decurved, 25^44 cm long, 0.3-0.5 cm diameter; bracts erect, divergent distally, 11-18 cm long, blades 7.4-14.4 cm long and 0.8-1.7 cm wide, lowermost green, middle and upper suffused deep red-pink to crimson, subligulate to long-triangular, apex attenuate to acute sheaths clasping erect, 2.2- 4.2 cm long and 1.5-2.4 (3.3) cm wide, suf- fused red-pink to crimson, ovate. INFLO- RESCENCE pendent, 20-34 cm long and 10-14 ( 19) cm in diameter, subcylindrical to subconical, rachis deep reddish-pink to crimson; primary bracts scalariform- spreading, 8.5-13.5 cm long and 2.0-2.4 cm wide, deep red or crimson, ovate in outline. LATERAL BRANCHES spreading, 1-1.8 cm long, subcollaterally 2-flowered, pedun- cles 0.7-1.5 cm long, rachis 0.2-0.4 cm long. FLOWERS sessile or pedicels to 0.3 cm \ong, floral bracts 1.6-2 cm long and 1.5-2 cm wide, subcoriaceous, deep pink to crim- son, minutely pitted, or brown-punctate abaxially, lepidote adaxially, oblong to ellip- tic or ovate-truncate, apex obtuse to broadly rounded and minutely apiculate, entire or torn-incised up to one-half the length at anthesis, keel inflated; sepals 1 .6-1.9 cm long and 1.2-1.5 cm wide, coriaceous, keeled, glabrous abaxially, lepidote to rounded; petals 2.8-3 cm long and 0.8-1 cm wide, pale greenish-white, obovate to subpandurate; appendages 0.5-0.6 cm long and 0.2 cm wide; stamens forming a hood over the style, fila- ments 2.0-2.1 cm long, anthers 0.4 cm long; ovary 0.3-0.4 cm long and 0.4 cm in diame- ter, style 1.5-1.7 cm long. PHENOLOGY: Flowering April and May. DISTRIBUTION: Known only from the northern part of the Cordillera de Tala- manca in Costa Rica in the lower montane wet forest at 1800-2100 m elevation. DISCUSSION: The features which char- acterize V. vietoris are its unhanded leaves, pendent, many-flowered inflorescence ( 14 or more flowers) subscalariform primary bracts and floral bracts which are equal or subequal to the brightly colored sepals. The flowers 56 Tulane Studies in Zoology and Botany Vol. 24 Fig. 22. Vriesea vietoris. A. Habit; B. Lateral inflorescence branch; C. Petal; D. Lateral inflorescence branch; E. leaf; F. floral bract excised and flattened. A, B, E, from (Utley & Utley 2I72X C, F, from (Utley & Utley 4784X D, from (Utley & Utiev 2216). No.l Middle American Vrieseas 57 are diurnal, greenish-white, tubular and weakly zygomorphic. Vriesea vietoris is very similar in general appearance to both V. leu- cophylla and V. hainesiorum but differs from them in characters mentioned above in the accompanying key, as well as in ecologi- cal preference. It is known only from the premontane wet forest, whereas V. leuco- phylla and V. hainesiorum are most abund- ant in the premontane rainforest and pre- montane moist forest respectively of the Holdridge life zone system (Holdridge, 1967). In the topographically complex region south of Cartago, where these three life zones are closely adjacent, the taxa occur sympatrically. The only indication of poten- tial introgression in the area is the reduction of banding in the leaves of V. leucophylla\ however the other characters which distin- guish the three species remain constant. This paucity of intermediates may be partially explained by the phenologies of the species in the zone of sympatry. Vriesea vietoris has been observed in flower during April and early May, while in the same areas V. leuco- phylla displays a flowering peak in July and August. This is in contrast to the strong June flowering peak of the latter species in areas of nonsympatry. Vriesea hainesiorum, which is rare in the area of sympatry, typically flowers from May through July. However, despite a deliberate search, flowering specimens of V. hainesiorum were not observed in the con- tact area during the flowering period of V. vietoris or V. leucophylla. ADDITIONAL SPECIMENS EXAMINED: COSTARICA: BORDER OF CARTAGO AND SAN JOSE PROVINCES: 18 miles S of Cartago on Interamerican Highway, N side of Talamanca Mts., 5 August 1962, A.Lee & Bruce L. Haines 7(5i (US). Near the junction of C. R. Hwy. 2 (Interamerican Highway) and 222 about 4-5 km S of El Empalme in the Cordillera de Talamanca, 16 April 1975, Vtley & Utley 2172 (NOLS), 2173 (DUKE); pasture along the Interamer- ican Highway 200 yards SE of Casa Mata, 9 May 1976, Utley & Utley 4744 (CAS), 4745 (NY), 4746 (US), 4748 (DUKE), 4757 (CR) & 4758 (NOLS); along Interamerican High- way W of Cartago, 9 May 1976, Utley & Utley 4782 (US), 4783 (MO), 4784 {D\JKE); along Interamerican Highway S of Cartago, 3 April 1979, A. Meerow, L. Besse & K. Tan 1117 (SEL). SAN JOSE PROVINCE: on CR. Hwy. 222about2kmfromthejunction with the Interamerican Highway and about 3.5 km E of San Cristobal Sur, 17 April 1975, Utley & Utley 2215 {¥), 2216 {\)\]Y.E)\ along CR. Hwy. 12, 6 km from the junction with the Interamerican Highway, 15 April 1976, Utley & Utley 4648 (F). 17. Vriesea uxoris Utley, Brittonia 33:583- 584. 1982. Fig. 23 TYPE: COSTA RICA: BORDER OF CARTAGO AND SAN JOSE PROVINCES: bog and surrounding thickets about 15.9 km S of El Empalme on the Interamerican High- way, 4 July 1976, Wilbur 19892 (HOLOTYPE: DUKE!; photographs CR, US). Epiphytic, acaulescent. LEAVES in a sub- spreading to subascending rosette, 33.0-44.0 (48.0) cm long; blades {\%.Q) 20.0-24.0 (36.0) cm long and 3.6-5.0 cm wide, green or green suffused with maroon to entirely maroon, ligulate to broadly triangular, apex acute to acuminate; 5/jea/;25(12.0)15.0-17.0(I9.5)cm long and 6.0-8.4 cm wide, obovate, castane- ous usually with a coating of cinereous tri- chomes. SCAPE decurved, (34.0) 38.0^W.O cm long and about 0.4 cm in diameter; bracts (13.0) 17.0-23.0 (33.0) cm long and 3.0^.2 cm wide, the blade and sheath portions not well delineated, ascending to erect becoming divergent or spreading near apices, enfolding the scape in the sheath area, usually red- pink, at times suffused with green or rarely entirely green, ligulate to subligulate or 58 Tulane Studies in Zoology and Botany Vol. 24 broadly triangular, apex acute. INFLOR- ESCENCE pendent, 21.0-38.0 cm long and about 17.0-30.0 cm in diameter, pyramidal, rachis glabrous occasionally with numerous castaneous punctulae,green to strongly suf- fused with red or maroon; primary bracts spreading, 1 1.0-15.0 (21.0) cm long and 2.8- 3.6 cm wide, pink to peach or salmon colored, occasionally with fine longitudinal maroon lines, subligulate to broadly triangu- lar, apex acute to acuminate. LATERAL- BRANCHES spreading (0.9) 1.2-1.7 cm long, collaterally 2-flowered, peduncles 0.9- 1 .4 cm long, rachis 0.0-1 .3 cm long. FLOW- ERS 0.3-0.8 cm pedicellate;y7ora/ ^ram 1 .0- 1.4 cm long and 1.2-1.7 cm wide, green or yellow-green, drying to brown, even to ob- scurely nerved, frequently rugulose especially in distal portion, keeled, obovate to subor- bicular. apex rounded or truncate, minutely apiculate or torn-incised; sepals 2.2-3.0 cm long and 1.0-1.5 cm wide, green drying to pale brown with a castaneous base, coria- ceous, obovate, the apex rounded to trun- cate, occasionally minutely apiculate or torn- incised ;/7e/fl/5 3.0-3. 1 cm long and 0.8-0.9 cm wide, green, obovate; appendages 0.6 cm long and 0.2 cm wide; stamens in a hood configuration over the style, filaments 1.6- 1.8 cm long, anthers 0.4-0.5 cm long; ovary 0.4 cm long and 0.3 cm in diameter, style 2.3 cm long; capsules 3.8^.0 cm long and about 0.8 cm in diameter, broadly fusiform, castaneous. PHENOLOGY: Flowering April and May. DISTRIBUTION AND HABITAT: Known only from the Cordillera de Tala- manca in Costa Rica in the montane rain forest life zone. DISCUSSION: Vriesea uxoris is a distinc- tive montane species characterized by green to maroon, unmarked leaves, a lax, pendent inflorescence with a comparatively slender rachis and peach or salmon-colored primary bracts. These characters in conjunction with the long sepals (2.2-3 cm) and floral bracts (1.1-1.4 cm) consistently much shorter than them immediately separate V. uxoris from the remainder of the thecophylloid vrieseas. Vriesea uxoris resembles V. leucophylla, V. vietoris and V. hainesiorum in its pendent inflorescence, brightly colored primary bracts and green, diurnal flowers. However there are numerous differences in leaf shape, inflorescence and floral characters, and habi- tat preference. The leaf blades of V. leuco- phylla, V. vietoris and V. hainesiorum are narrower (1-3 cm) and generally triangular in outline. Moreover, in the latter three taxa the floral bracts are equal or subequal to the sepals at anthesis and the primary bracts are pink to red in color while the floral bracts of V. uxoris are not more than one-half as long as the sepals and the primary bracts are peach to salmon in color. The montane rain- forest habitat of V. uxoris contrasts with the lower and frequently drier pre-montane hab- itats of the other pendent-inflorescenced taxa. Vriesea uxoris does not occur sympat- rically with these species. SPECIMENS EXAMINED: COSTA RICA: BORDER OF SAN JOSE AND CARTAGO PROVINCE: about 17 km SE of El Empalme on the Interamerican High- way, 6 April l914Mtley & Utley 811 (CR), 812 (MO) & 843 (DUKE); 17 March 1975, Utley & Utley 2004 (NOLS), 2005( DUKE), 2006 (NOLS); 27 October 1975, Utley & Utley 3202 (CAS); 9 May 1976, Utley & Utley 4755 (VSl 4756(F). 18. Vriesea pictaiMez & Werckle) L. B. Smith & Pitt. Journ. Wash. Acad. Sci 43: 403. 1953. Thecophyllum pictum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 874. 1904. TYPE: COSTA RICA: Province not given, near Desengano, elevation 1460 m, Werckle Bromel. Costaric. 119 pro No.l Middle American Vrieseas 59 parte. (LECTOTYPE: B!; ISOLECTO- TYPE: B!). Epiphytic, acaulescent to several cm cau- lescent. LEAVES in a subascending rosette, 33.0-45.0 cm long; blades 18.5-30.0 cm long and 3.3-4.8 cm wide, green with darker green wavy transverse lines, these weakly suffused with maroon at the blade-sheath junction, ligulate, apex acute; sheaths 12.8-17.6 cm long and 5.0-7.5 cm wide, castaneous abax- ially becoming green suffused with indistinct wavy maroon transverse lines distally, brown abaxially with distinct wavy maroon trans- verse lines, ovate. SCAPE erect, 41.5-61.5 Fig. 23. Vriesea uxoris. A. Habit; B. Leaf; C. Lateral inflorescence branch; D. Petal; E. Floral bract, excised and flattened; F. Flower. A, B, C, E, From (Utiey & Uiley 4756); D, F, From Wilbur 19892. 60 Tulane Studies in Zoology and Botany Vol. 24 cm long and 0.7-1.0 cm in diameter; bracts 6.9- 1 1 .0 cm long, blade 3.7-7.7 cm long and 1.8-2.9 cm wide, green with wavy maroon transverse lines, erect, divergent distally, lig- ulate to subtriangular, apex acute, sheaths about 3.1-3.3 cm long and 3.5-4.5 cm wide, green with faint wavy transverse lines dis- tally, sheathing erect, ovate. INFLORES- CENCE erect, 13.0-20.0 cm long and 6.0- 11.0 cm in diameter, cylindric; primary bracts 6.8-7.8 cm long, blades about 4.0-5.0 cm long and 2.0 cm wide, green with darker green wavy transverse lines, these occasion- ally suffused with maroon distally, spread- ing, subligulate, apex acute, sheaths about 3.5 cm long and 4.5 cm wide, ovate in out- line. LATERAL BRANCHES spreading to ascending, to 4.5 cm long, collaterally biflo- wered, pedicels wanting or to 0.1 cm long. FLOWERS sessile; floral bracts about 1.6 cm long and 0.5-0.6 cm wide pale green, membranaceous, unkeeled, brown-centered lepidote adaxially, glabrous with brown- margined pellucid punctulae abaxially, lan- ceolate, apex narrowly acute; sepals 1.9-2.2 cm long and 1 .3-1 .6 cm wide, green suffused with maroon near margins and apex, brown- centered lepidote adaxially immersed lepi- dote abaxially, elliptic to oblong elliptic, apex acute to obtuse; petals 4.5-5.0 cm long and 1.6-1.9 cm wide, greenish white, indis- tinctly ovate proximally becoming ligulate distally; appendages 0.5 cm long and 0.3-0.4 cm wide; stamens in a hood configuration over the gynoecium, filaments 4.3-4.5 cm long, anthers 0.6-0.7 cm long; ovary more or less 0.6 cm long 0.5-0.6 cm in diameter, style about 4.2 cm long. PHENOLOGY: Flowering plants collected during May. DISTRIBUTION AND HABITAT: Known from the type location in Costa Rica and from northwestern Panama between 1900 and 2000 m in the lower montane rain forest life zone. DISCUSSION: The type material of Vrie- seapicta must be considered a mixed collec- tion composed of two taxa, V. picta and V. viridis. However, the element which repres- ents V. viridis does not agree with either the diagnosis of V. picta or the illustration of V. picta in Das Pflanzenreich (Mez, 1934- 1935). Specimens of the remaining element, however, agree in all critical characters with both the description and illustration of V. picta. This situation possibly originated sometime after the original circumscription of V. picta, since the discordant element lacks one of Werckle's original tags identify- ing it as "119". The addition may have occurred when Mez's personal collection was accessioned into the Berlin Herbarium. From the scant material of V. picta availa- ble, this species may be recognized by its caulescent habit, its formation of tank-roots, and its castaneous, brittle leaf sheaths. Flow- ers of V. picta are green-white, strongly zygomorphic and nocturnal. Vriesea picta is most similar to V. stenophylla and V. willi- amsii in floral morphology, the formation of tank-roots and brittle leaf bases. A combina- tion of features may be used to distinguish these three taxa from one another and include leaf markings, inflorescence length, floral bract shape and length and flower size. In Costa Rica V. picta is still only known from the type location where I re-collected it. Only three flowering individuals were ob- served {Utley & Utley 4961. 4979. & 4984) and these were all part of a large cluster growing in a mass on a tree trunk. No other plants of V. picta were seen in the area. The Panamanian material of this taxon is unique in that the floral bracts are absent. This con- dition has not been noticed in any other specimen of thecophylloid vriesea. Further field study and collections are needed to crit- ically evaluate the status and relationships of this unusual taxon. SPECIMENS EXAMINED: COSTA RICA: HEREDIA PROVINCE: About 3 km south of the junction of highways 9 and 120 on Highway 9. slopes above the Rio No.l Middle American Vrieseas 61 Desengano, 22 May 1976. Utley & Vtlev 4961 (DUKE) 4979{CK) & 4984 (NOLS). PANAMA: CHIRIQUI PROVINCE: Las Nubes, 7 August 1974, Croat 26448 {UO) & 26468 (MO). 19. Vriesea stenophyllaiMez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci 43: 403. 1953. Thecophyllum stenophyllum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 875. 1904. TYPE: COSTA RICA: location not given, Werckle Bromel. Costaric. n. 112 (HOLOTYPE: B!, photograph US). Epiphytic, or terrestrial, short caulescent. LEAVES in a ascending or subascending rosette, (15.0) 22.0-36.0 (44.0) cm long; blades (9.0) 16.0-28.0 (37.0) cm long and 1 .3-2.4 cm wide, green with distinct maroon longitudinal lines, long triangular to ligulate, apex attenuate to acute; sheaths (5.6) 6.6-8.6 cm long and 2.0-3.4 cm wide, castaneous abaxially, suffused with maroon longitudi- nal lines distally and adaxially elliptic to ovate in outline. SCAPE erect, (16.0) 23.0- 39.0 cm long and 0.2-0.4 cm in diameter; bracts 4.-12.0 (20.0) cm long, blades 2.0-17.0 cm long and 0.7-1.3 cm wide, green with maroon longitudinal lines, erect, divergent distally, ligulate to triangular, apex attenuate or rarely acute, sheaths 1 .6-2.3 (2.9) cm long and 1 .4-1 .7 cm wide, green with longitudinal lines, sheathing erect, ovate. INFLORES- CENCE erect, 3.5-6.0 cm long and about 4.0-6.0 cm in diameter, laxly short cylindric rachis green or green with longitudinal maroon lines; primary bracts 3.3-5.7 cm long, blades 1.9-3.6 cm long and 0.3-0.9 cm wide, green or green suffused with maroon, marked with distinct maroon longitudinal lines, spreading, subligulate to triangular or long triangular, apex attenuate to narrowly acute, sheaths 1.9-2. 2 cm long and 1.0-1.8 cm wide, green with longitudinal maroon lines, ovate to elliptic; LATERAL BRANCHES spreading sessile, the two flowers borne col- laterally in the axil of the primary bracts. FLOWERS sessile to rarely 0.3 cm pedicel- late;/7ora/6rarac/5 1.1 -1.4 cm long and 1.0-1.5 cm wide, green, submembranous to coriace- ous, brown-centered lepidote adaxially, im- mersed brown-centered lepidote with brown- margined pellucid excrescences abaxially, unkeeled or infrequently keeled, broadly obovate, oblong-obovate to elliptic or sub- orbicular, apex obtuse to broadly rounded or suborbicular, usually minutely acumi- nate; sepals 1.1-1.6 cm long and 0.8-1.2 cm wide, green, coriaceous, brown-centered lep- idote adaxially and immersed brown-cen- tered lepidote abaxially, obovate to elliptic, apex obtuse to broadly rounded; capsules about 2.0 cm long and 0.7-1.9 cm in diame- ter, fusiform dark brown to castaneous in color. PHENOLOGY: Although flowering spec- imens have been collected only during August, this species probably flowers throughout the late dry season and early rainy season (June-September). DISTRIBUTION AND HABITAT: Costa Rica and Panama between 700-1600 m in the premontane rain forest life zone and possibly extending into the premontane basal belt transition of the tropical wet forest life zone. DISCUSSION: Vriesea attenuata is an in- frequent species distinguished by the cinere- ous abaxial surfaces of the leaf blades and the dark castaneous sheaths in conjunction with the floral bracts which are about equal to to shorter than the sepals. The flowers are white and nocturnal. Although the exact form of the corolla is not known, the general shape is probably tubular to narrowly campanulate. Vriesea attenuata is most closely related to and frequently grows sympatrically with V. comata and is difficult to distinguish from this taxon in vegetative condition. In flower or fruit, they are readily separable from each other. In contrast to the 2.5-3.0 cm long, obovate, olive-green to brown capsules characteristic of V. comata, those of V. atten- uata, are shorter (about 2 cm fusiform and castaneous, these species may also be separ- ated by the degree of development of the sterile coma and floral bract length. Al- though only a few flowering specimens have collected of V. attenuata, this taxon appears to lack the gelatinous matrix on its inflores- cences which is commonly observed in V. comata. Vriesea attenuata also superficially resembles V. kathyae, from which it differs in sepal length, leaf length and coloration and floral bract length. Smith and Downs (1977) placed V. atte- nuata in synonymy with V. stenophylla on the basis of the congruence of the taxa in floral bract length relative to sepal length and the presence of longitudinal lines in the leaves of both taxa. After an evaluation of the type material and my personal collec- tions in conjunction with my field observa- tions, I am unable to agree with their disposi- tion of this taxon. Vriesea attenuata can be separated from V. stenophylla consistently on the basis of its longer floral bracts (1.1-1.4 cm vs. 0.7-0.9 cm) and its cinereous abaxial leaf surfaces. Although the inflorescences of these taxa appear similar, they are subtly different in form. Moreover, the longitudinal lines which are present in the leaves of V. attenuata are narrow and the color is con- fined to the veins, while those of V. steno- phylla are much broader and the color is not 66 Tulane Studies in Zoology and Botany Vol. 24 confined to the veins. SPECIMENS EXAMINED: COSTA RICA: PROVINCE OF ALAJUELA: North of San Ramon between Los Angeles Norte and about 7 km north, 15 December 1974, Utley & Utley 1638 (DUKE); north of San Ramon between La Balsa and about 4.5 km north,26 February 1975, Utley & Utley 1852 (DUKE), & 1854 (CR); north of San Ramon between La Balsa and the Rio Cata- ratas or about 12 km north of La Balsa, 27 January 1976, Utley & Utley 3738 {D\]¥.E\ 3747 (CR), 3748 (NOLS), 3749 (F), 3759 (NY), 3764 (MO) 3765 (K), 3766 (DUKE), 3749 (US); about 4 km north of la Balsa de San Ramon, 8 February 1976, Utley & Utley ^7/5 (DUKE). BORDER OF ALAJUELA AND HEREDIA PROVINCES: Vicinity of Colonia Virgen del Socorro on and around the finca of Sr. Carlos Molina or about 3 to 6 km east of Cariblanco, 10 August 1975, Utley & Utley 2850 (NOLS) 2860 (US), 2863 (DUKE). SAN JOSE PROVINCE: La Hondura, 9 March 1926, Standley & Valerio 51909 (US). PANA- MA: COCLE PROVINCE: north of El Valle de Anton, trail to Las Minas, 2 December 1941, Allen 2884 (US). VER- GUAS PROVINCE: northwest of Santa Fe, 8.8 km from Escuela Agricola Alto de Piedra, 25 February 1975, Mori & Kallunki 4854 (US). 22. Vriesea comata (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Thecophyllum comatumMez 8l Werckle in Mez, Bull. Herb. Boiss. ser. 11.4:871. 1904. TYPE: COSTA RICA: near Turrialba, alt. 900 m, Werckle s.n. (HOLOTYPE: B!, photograph US!). Vriesea hygrometrica var. angustifolia L.B. Smith Phytologia 8: 222. 1962. TYPE: COSTA RICA: without further locality, Robert G. & Catherine Wil- son 32 HOLOTYPE: US!). Epiphytic or terrestrial, to 10 cm cau- lescent. LEAVES in a spreading to sub- ascending rosette, 16.0-23.0 (26.0) cm long: Mfl^e59.0-I4.0 (16.5) cm long and 1.5-3.2cm wide, green, usually with longitudinal purple- castaneous lines proximally, ligulate to long triangular, apex acute to acuminate or at- tenuate; sheaths 5.0-9.5 cm long and 2.7-4.5 cm wide, castaneous abaxially, occasionally with longitudinal purple-castaneous lines distally, plumbeous adaxially, usually with longitudinal purple-castaneous lines, elliptic to ovate or obscurely obovate. SCAPE erect, (16.0) 19.0-23.0 (25.0) cm long and 0.3-0.5 cm in diameter; bracts 4.5-10.0 (14.0) cm long, blades 3.0-6.5 (9.5) cm long and 1 .2-2.4 cm wide, green, erect divergent distally, ligu- late to subligulate or triangular, apex acute to attenuate, sheaths (1.8) 2.8-4.0 (5.2) cm long and 2.2-3.2 cm wide, green, sheathing erect ovate. INFLORESCENCE erect, 4.5- 9.5 cm long and 5.0-10.0 cm in diameter, short, dense cylindric to subcapitate; prim- ary bracts 5.0-7.5 cm long, the blade and sheath merging gradually, subligulate to tri- angular, apex acute to attenuate, sheaths (2.0) 2.6-3.4 cm wide, green ovate. LATER- AL BRANCHES spreading to ascending, sessile to 0.3 cm pedicellate, the 2 or rarely 3 flowers borne collaterally or subcollaterally. FLOWERS sessile to 0.2 cm pedicellate; floral bracts 2.5-3.0 cm long and 1 .7-2.2 (2.7) cm wide, green, subcoriaceous, brown- centered lepidote on both surfaces, strongly keeled, the bracts usually conduplicately folded along the keel, ovate to oblong-ovate, apex broadly acute to rounded, occasionally asymmetric; sepals 1.3-1.6 cm long and 0.8- 1.0 cm wide, green coriaceous, brown-cen- tered lepidote adaxially, brown-centered, frequently immersed lepidote abaxially ellip- tic or rarely obovate, apex obtuse to broadly acute, petals about 2.5-2.8 cm long and 1 . 1 No.l Middle American Vrieseas 67 cm wide, white; appendages 0.2 cm long and 0.15-0.2 cm wide; stamens in a hood con- figuration over the gynoecium, the filaments flexed and the anthers weakly connivent by their lateral margins, filaments about 2.1 cm long and the anthers 0.4 cm long; ovary about 0.35 cm long and 0.4 cm in diameter, style 1.7-1.8 cm long; capsules 2.2-2.5 cm long and about 0.8 cm in diameter, obovate, olive-green to brown. PHENOLOGY: A single flowering speci- men has been collected in January. DISTRIBUTION AND HABITAT: Known only from Costa Rica where the species has been collected between 700-1500 m in the premontane rain forest life zone. Consider- ing the abundance of this species at middle elevations in Costa Rica, more intensive col- lecting in Panama presumably would result in the collection of this taxon. DISCUSSION: Vriesea comata is notable for its subcapitate to short cylindric inflor- escence terminated by a conspicuous sterile coma. The flowering inflorescence is usually covered with a gelatinous matrix similar to that observed in V. capitata and V. brac- teosa. As a result, the white, campanulate nocturnal flowers which are borne in the axils of the usually spreading primary bracts emerge from the exudate at anthesis. The spreading rosettes of V. comata are relatively small and formed of leaves with castaneous sheaths and green blades which are long- itudinally lined with purple proximally. Vriesea comata is readily distinguished from other thecophylloid vrieseas in fruit by its obovate, olive-green to brown capsules. The type of V. hygrometrica var.angustifo- lia is very similar in its critical characters to V. comata. The taxa were tacitly separated using leaf pubescence (Smith and Downs, 1977). In V. comata the leaves were des- cribed as cinereously pubescent while those of V. hygrometrica var. angustifolia were thought to be glabrous or brown-punctulate. However, these taxa share a number of important characters including leaf shape, color, and marking as well as primary bract shape and orientation, inflorescence shape and floral bract length. These features coupled with a striking congruence in overall appearance are compelling reasons for plac- ing V. hygrometrica var. angustifolia in syn- onomy under V. comata. Vriesea comata is most closely related to V. attenuata with which it is frequently found growing sympatrically. Although few flow- ering specimens of either taxon have ever been collected, V. comata and V. attenuata appear to flower at different times of the year. However, more collections of flower- ing material are needed to verify these obser- vations. Like V. attenuata, V. comata also superficially resembles V. kathyae, but it is immediately separable from this taxon on the basis of floral bract and sepal length, leaf coloration and capsule characters. The spe- cies relationships of these three taxa are further discussed under V. attenuata . SPECIMENS EXAMINED: COSTA RICA:ALAJUELA PROVINCE: Roadside and pasture trees between Hwy. 9 and Lag- una Hule on a semi-maintained secondary road, between 0.5 and 5.0 km northwest of Cariblanco, 900-1000 m, 16 October 1974, Utley & Vtley 1370 (CR, DUKE, F, K, NOLS, US); 7 December 1975, Utley & Utley 3561 (DUKE); North of San Ramon on road to Los Angeles Norte, between Los Angeles Norte and 7 km north 1000-1200 m, 15 December 1974, Utley & Utley 1680 (DUKE) & 1684 (MO); North of San Ramon between La Balsa and 4.5 km north, 1000 m, 26 February 1975, Utley & Utley 1854 (DUKE) & 1863 (CR); North of San Ram6n between Los Angeles Norte and 7 km north of La Balsa, 900 m, 8 August 1975, Utley & Utley 2801 (DUKE); North of San Ramon, between La Balsa and the Rio Cata- ratas, 550-1 100 m, 27 January 1976, Utley & Utley 3750 (US), 3770 (F), 3771 (DUKE), i772(NOLS), 3773 (CAS), 3790 (NOLS), 68 Tulane Studies in Zoology and Botany Vol. 24 3791 (CR), & 3792 (GH); 7 to 9 km north of San Ramon, 1 100 m, 27 January 1976, Utley & Utley 3798 (US), 3800 (DUKE) & 3801 (F); about 13 km north of La Balsa de San Ramon on roadbanks and hillsides above the Rio Catartas, 500-600 m, 8 February 1976, Utley & Utley 4061 (DUKE); about 4 km north of La Balsa de San RamOn, 1000 m, 8 February 1976, Utley & Utley 4109 (DUKE). BORDER OFALAJUELA AND HEREDIA PROVINCES: 3-6 km east of Cariblanco in the vicinity of Colonia Virgen del Socorro, 900 m, 10 August 1975, Utley & Utley 2857 (DUKE) & 2858 (CR). CAR- TAGO PROVINCE: 19 km south-south- east of the town of Turrialba, in the region southwest of Platanillo, 990-1200 m, 23 July 1975, Utley & Utley 2684 (US), 2685 (DUKE), 2686{CK), 2687 (NOLS), 2<5«<^ (B), 2689 (MO), 2690 (MICH) & 2691 (GH). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Heredia on Calle Yerbabu- ena, 1400 m, 3 April 1976, Utley & Utley 4474 (DUKE). SAN JOSE PROVINCE: Alto de La Palma, Finca Porvenir, in secon- dary growth and remnant trees and pastures, 1500 m, 18 August 1975, Utley & Utley 2901 (DUKE); vicinity of Bajo la Hondura, between the plateau and the steel bridge over the Rio Hondura, 1050 m, 23 October 1975, Utley & Utley 3183 (DUKE); about 7 km north of San Jerdnimo in the vicinity of La Palma on CR. Hwy. 220, 1500 m, 27 December 1975, Utley & Utley 3686 (DUKE). 23. Vriesea singuliflora (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953. Thecophyllum singuliflorumMez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 870. 1904. TYPE: COSTA RICA: Werckle 89 (HOLOTYPE: B!, photograph US!). Probably epiphytic, slightly caulescent. LEAVES in a ascending rosette, to about 22.0 cm long, drying to brown; sheaths abax- ially castaneous with pale margins, adaxially light brown. SCAPE erect, about 18.0 cm long, 0.1 cm in diameter or slightly less; bracts about 5.5 cm long, blades erect, spreading distally, drying brown with traces of purple, apex acute to attenuate, sheaths clasping-erect, drying brown. INFLORES- CENCE erect, 3.0-7.0 cm long and 2.0-3.0 cm in diameter, loosely cylindric; pirimary bracts spreading, blades 2.0 cm long and about 0.3 cm wide, dried brown, filiform, apex attenuate, sheaths broadly ovate to sub- orbicular, 1.0 cm long and 1.0 cm wide. LATERAL BRANCHES spreading, bear- ing a single, secund flower and a minute vestige of a second flower. FLOWERS with pedicels 0.3-0.4 cm long; floral bracts 0.6 cm long and 0.6 cm wide, chartaceous-mem- branaceous, finely nerved, ecarinate or with an indistinct longitudinal fold near the right margin, broadly elliptic to subrotund, apex broadly obtuse; sepals 1.5 cm long and 1.1- 1.2 cm wide, coriaceous, broadly elliptic to suborbicular; pe/fl/5 2.0 cm long and 0.7 cm wide, elliptic; appendages 0.2-0.3 cm long and 0.15 cm wide; stamens probably in a hood configuration over the gynoecium, fil- aments 1 .4 cm long, anthers 0.4 cm long and connivent by their lateral margins into an androecial hood; ovary 0.4 cm long and about 0.2 cm wide, style 1.3 cm long, curved and the stigma probably oriented away from the androecial hood. PHENOLOGY: Unknown, but the type specimen was collected in January and appears to have been flowering or near flowering. DISTRIBUTION AND HABITAT: Known only from the type location, Carillo, in Costa Rica at 300 m elevation in the tropical wet forest life zone. DISCUSSION: Although Vriesea singuli- flora is known only from the type and one No. Middle American Vrieseas 69 other collection, it is distinct among the the- cophylloid vrieseas in consistently having one-flowered lateral branches. From my examination of the material, the flowers appear to be either campanulate or tubular. Although the flowers of this taxon perhaps are diurnal, they are more likely either cre- puscular or nocturnal. During my field work, I have never observed diurnal flower- ing thecophylloid vrieseas at elevations lower than 1 500 meters; these taxa appear to be restricted to montane and premontane regions. In contrast, nocturnal flowering taxa are abundant at lower elevations in premontane and lower montane life zones. Although thecophylloid vrieseas have been collected frequently at lower elevations (700- 1500 m), V. singuliflora is unique among these taxa in apparently growing at low ele- vations (300 m) in tropical wet forests. Considering the green, unmarked leaf blades and castaneous leaf sheaths with pale margins that are found in V. singuliflora and its ecological preference, this taxon probably has its closest affinities with the comata species group. However, this species lacks large floral bracts that are characteristic of several species in the group. Because I was never able to collect V singuliflora, 1 am by no means certain of its exact nature and affinities. Further collections and field ob- servations are needed to resolve the status of this anomalous species. SPECIMENS EXAMINED: COSTARICA: PROVINCE OF SAN JOSE (?): Vicinity of Carillo, January 1908, Werckle 144 (=Museo Nacional de Costa Rica #17423, (B). 24. Vriesea hygrometrica (Andre) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Fig. 13 Caraguata hygrometrica Andre, Enum. Bromel. 6. 13 December 1888. TYPE: COLOMBIA: NARINO: Alto de San Antonio near Cali, Andre 2638 (HOLOTYPE: K!, photography US!). Guzmania hygrometrica (Andre) Andre ex Mez, DC. Monogr. Phan. 9: 919. 1896. Thecophyllum hygrometricum (Andre) Mez, Bull. Herb. Boiss ser. 1 1. 3: 131. 1903. Epiphytic or terrestrial to 8 cm caulescent. LEAVES in a subspreading to ascending rosette, 21.0-31.0 (52.1) cm long; blades (13.5) 17.5-21.0 (34.5) cm long and 2.0-3.8 cm wide, green with fine wavy transverse lines, these ranging from dark green and relatively inconspicuous to deep red-maroon, frequently most pronounced proximally, ligulate to subligulate, apex acute to acumi- nate or rarely attenuate; sheaths 7.4-13.5 (17.6) cm long and 3.7-6.2 cm wide, castane- ous with pale margins abaxially and grey to greyish-brown adaxially, frequently with wavy transverse lines distally, elliptic to oblong or oblong-ovate. SCAPE erect or curved-erect, 33.5-45.0 (72.7) cm long and 0.4-0.8 cm in diameter; bracts (3.1) 3.8-7.6 (9.3) cm long, the blades ( 1 .0) 1 .5-3.6 (6.0) cm long and 1 .0- 1 .9 cm wide, green occasionally suffused with wavy transverse lines, erect to divergent apically, triangular to ligulate or subligulate, apex acute to acuminate or attenuate, sheaths 2.1-4.0 cm long and 1.9- 3.3 (3.9) cm wide, drying to ochre adaxially sheathing erect, ovate to elliptic in outline. INFLORESCENCE erect, 5.0-6.5 (12.0) cm long and 5.5-7.0 (9.5) cm in diameter; pr/m- ary bracts 3.1-5.8 cm long, blades 1.0-2.8 (3.7) cm long and 1.0-1.7 cm wide, green occasionally with wavy transverse lines, spreading at anthesis, triangular, apex acute to acuminate or attenuate, sheaths 2.1-3.3 cm long and 2.0-3.7 cm wide, ovate or broadly ovate to elliptic. LATERAL BRANCHES spreading or divergent 3.0-4. 1 cm long collaterally 2-flowered, peduncle wanfing to 0.4 cm long, rachis wanting to 0. 1 cm long. FLOWERS sessile; floral bracts 70 Tulane Studies in Zoology and Botany Vol. 24 (2.1) 2.5-3.3 (3.8) cm long and 1.8-3.0 cm wide, green, subcoriaceous, brown-centered lepidote to subglabrous, keeled, and the bracts usually conduplicate along the keel, oblong to oblong-ovate, apex obtuse to rounded, frequently short acuminate or torn-incised; sepals 1.4-2.0 cm long and 0.9- 1.3 cm wide, green drying pale castaneous, coriaceous, brown-centered lepidote adax- ially, frequently immersed lepidote abax- ially, elliptic to obovate, apex obtuse to rounded, frequently minutely apiculate; petals about 2.3-3.4 cm long and 1 .3- 1 .8 cm wide, white, elliptic to obovate with a medial constriction on the lateral margins; appen- dages 0.4-0.5 cm long and about 0.2 cm wide; stamens in a hood configuration over the style, anthers connivent by their latral mar- gins, filaments 1.9-2.6 cm long, anthers 0.5- 0.8 cm long; ovary 0.4-0.6 cm long and 0.3- 0.4 cm in diameter, style 2.0-2.7 cm long; capsules about 1.9 cm long and 0.8 cm in diameter, castaneous. PHENOLOGY: Flowering plants have been collected in February, April, May and July. Time of flowering apparently is pop- ulation dependent, and for the species as a whole probably spans several months. DISTRIBUTION AND HABITAT: This species is the widest ranging thecophylloid, extending from southern Mexico to Ecua- dor. The species ranges from 700-1500 m in elevation and is found in the premontane rain forest and lower montane rain forest and possibly extends into the premontane wet forest. DISCUSSION: Among the thecophylloid vrieseas, Vriesea hygrometrica has the most widespread distribution, extending from southern Mexico through Central America and into Ecuador. Considering its distri- bution, this taxon remains poorly collected outside of Costa Rica. Vriesea hygrometrica is a variable species with green leaves that are frequently suffused proximally with wavy transverse lines ranging from green to red- maroon in color. Inflorescences of V. hygrometrica are short, dense cylindric to subcapitate at anthesis and frequently elon- gate during and after anthesis so that fruiting inflorescences are sublaxly cylindric. More- over, the inflorescences are typically covered with a gelatinous exudate at anthesis. The white to greenish white campanulate flowers are nocturnal. In one collection, Utley & Utley 877, the flowers had a sweet, spicy odor in the early evening. Vriesea hygrometrica is very similar to V. notata and , at times, is difficult to distin- guish from this latter taxon. The only char- acter that separates these taxa consistently is the presence of longitudinal lines in the leaves, scape bracts and primary bracts of V. notata and their absence in V. hygrometrica. Populations of V. notata from Tapanti region exhibit a pronounced elongation of the inflorescence post anthesis. I doubt that an acyanic individual of V. notata in which the inflorescence had elongated post anthesis could be reliably distinguished from V. hygrometrica. Further field collections and observations, coupled with a study of breed- ing systems, are necessary to determine if the current delimitation of these taxa is biologi- cally sound. For the present, I have taken the course which causes the fewest taxonomic changes. Vriesea hygrometrica and V. notata are most similar to V. kathyae in habit, habitat and overall appearance. V. kathyae is regu- larly sympatric with these two taxa and may be distinguished from them by its shorter floral bracts, sepals and petals. SPECIMENS EXAMINED: COSTA RICA: ALAJUELA PROVINCE: north of San Ramon between Los Angeles Norte and about 7 km north of La Balsa, 26 February 1975, Vtley & Utley 1858 (US) & 1862 (DUKE); shore of Laguna Hule west of Cariblanco, 18 April 1975, Vtley & Utley 2238 /i (DUKE); 8 August 1975, Utley & Utley 2816 (DUKE); about 5 km west of No.l Middle American Vrieseas 71 Cariblanco on the road to Laguna Hule, 7 December 1975, Utley & Utley 3564 (DUKE); between La Balsa de San Ramon and the Rio Cataratas, 8 February 1976, Utley & Utley 4062 (DUKE), 4110 (US), 4111 (CR), 4112 (F), 4113 (NOLS), 4114 (DUKE)^779(MO)^/27(K),^72<^(DUKE) & ^75/ (NY). BORDER OF ALAJUELA AND PUNTARENAS PROVINCES: evergreen forest and wet wind-gap forma- tions on and near the Continental Divide about 2-5 km east and southeast of Monte Verde, 17-20 March 1973, Burger & Gentry 8618 (CR, F). CARTAGO PROVINCE:- mountains south of El Muneco, 7 April 1974, Utley & ^//ev<^77(DUKE); road from Tapanti to Taus and Tausito, 27 May 1976, Utley & Utley 5037 (DUKE), & 5039 (NOLS); between Tausito and Selva, 4-6 km northeast of the Rio Grande de Orosi at Tapanti; 27 May 1976, Utley & Utley 5067 (CR), 5068 {\5S), 5069(C ASl 5070 {¥), 5071 (DUKE), 5072 (GH), & 5073 (DUKE). SAN JOSE PROVINCE: Alto de La Palma, 29 June 1974, Utley & Utley 982 A, B & C(DUKE, US, CR); 6 July 1975, Utley & Utley 2573 (DUKE), 257^(F), 2575 (NOLS), 2589 (MEXU); 18 August 1975, Utley & Utley 2907 A,B & C (DVKE, MICH, MO); vicinity of Bajo la Hondura, between the Finca on the plateau and the steel bridge over the Rio Hondura, 23 October 1975, Utley & Utley 3184 (DUKE). MEXIC6: OAXACA: Vista Hermosa, 50 miles south of Valle Nacional, 28 April 1960, Van Hyn- ing 6051 (US). PANAMA: CHIRIQUI PROVINCE: Cerro Colorado, along road above San Felix, 14 July 1976, Croat 37107 (US). 25. Vriesea kathyae Utley, sp. nov. Fig. 24 A V. hygrometricaiAndri) L.B. Smith & Pitt, et V. notata L.B. Smith & Pitt., quibus verisimiliter affmis, bracteis florigeris, sepa- lis et petalis minoribus, laminis foliorum rus- sis differt. Epiphytic or terrestrial, to about 4.0 cm caulescent. LEAVES in a subspreading to ascending rosette, 17.0-32.0 cm long; blades 10.7-22.5 cm long and 1 .6-2.0 cm wide, green suffused with pink or russet-brown margi- nally or abaxially, with fine longitudinal maroon or purple lines, these frequently more pronounced proximally, ligulate, apex attenuate; sheaths 7.2-9.3 cm long and 2.6- 3.8 cm wide, dark castaneous abaxially fre- quently with pale margins and fine longitud- inal maroon lines distally, pale brown or greyish brown with fine longitudinal maroon lines adaxially. SCAPE erect, 27.0-35.0 cm long and 0.3-0.4 cm in diameter; bracts 4.1- 7.9 cm long, blades 1 .9-5.7 cm long and 0.8- 1 .8 cm wide, green with maroon longitudinal lines, drying to brown, erect to divergent distally, subligulate to long triangular, apex acute to narrowly acute or attenuate, sheaths 1.9-2.8 cm long and 1.8-2.4 cm wide, shea- thing erect, broadly ovate. INFLORES- CENCE erect, 4.5-6.0 (7.5) cm long and 4.0- 6.5 cm in diameter, short, cylindric in outline; primary bracts 3.7-4.5 cm long, blades 1 .6-2.7 cm long and 0.6-1 .0 cm wide, green frequently weakly suffused with maroon near margins, spreading, triangular to long triangular, apex narrowly acute to attenuate, sheaths 1.8-2.2 cm long and 2.0- 2.5 cm wide, ovate in outline. LATERAL- BRANCHES spreading 1.7-2.4 cm long, collaterally 2-flowered, rarely with a minute vestigial third bud, peduncle 0.2-0.4 cm long, rachis wanting to 0.2 cm long. FLOWERS sessile;y7ora/ bracts ( 1 .2) 1 .6- 1 .8 cm long and 1 .4- 1 .7 cm wide, green drying to brown, sub- coriaceous, lepidote adaxially, subglabrous to immersed lepidote abaxially, ovate to oblong and usually conduplicate along the keel, apex obtuse to rounded or rarely acute; sepals 0.7-1.0 cm long and 0.6-0.8 cm wide, green or olive green, drying brown to cas- taneous, lepidote adaxially, immersed lepi- 72 Tulane Studies in Zoology and Botany Vol. 24 dote abaxially, elliptic to broadly ellipitc or rarely suborbicular, apex obtuse to broadly rounded or rarely acute; petals 1.7-1.9 cm long and 0.9-1.0 cm wide, white, ovate to obovate constricted medially; appendages 0.3 cm long and 1.2 cm wide; stamens in a hood configuration over the gynoecium, anthers connivent laterally, filaments 1 .2- 1 .4 cm long, anthers 0.3-0.4 cm long; ovary 0.4 cm long and 0.3 cm in diameter, style 1 .2 cm long; capsules about 1.8 cm long and 0.7 cm in diameter, light castaneous, fusiform. TYPE: COSTA RICA: SAN JOSE PROVINCE: Alto de la Palmade San Jose on Finca Porvenir in uncut primary forest near streams and in secondary growth and remnant trees in pastures, elev. ca 1500 m, 18 August 1975, Utley & Utley 2888 (HOLO- TYPE: DUKE!, isOTYPES: CR!,F!and US!). PHENOLOGY: Flowering specimens have been collected in June. DISTRIBUTION AND HABITAT: Endemic to Costa Rica between 1400-1900 m in the premontane rain forest life zone and possibly extending slightly into the lower montane rain forest life zone. DISCUSSION: The most notable features of V. kathyae are its relatively short leaves (10.7-22.5 cm) which are green suffused with russet brown abaxially and castaneous sheaths, its floral bracts which exceed the sepals in length, and its extremely short Fig. 24. Vriesea kathyae. A. Inflorescence and upper scape; B. Leaf; C. Flower; D. Postanthesis flower with floral bract. From (Utley & Utley 2568). 73 Tulane Studies in Zoology and Botany Vol. 24 sepals (0.7-1.0 cm). The inflorescence of V. kathyae is erect and short cylindric, bearing white, nocturnal campanulate flowers. Vriesea kathyae is most similar to V. hygrometrica and V. notata in habit, its hab- itat requirements and the general appear- ance of its inflorescence. The charateristics which distinguish V. kathyae from these two taxa are discussed under V. hygrometrica. In addition to a general similarity to V. hygrometrica and V. notata, V. kathyae also has affinities with V. attenuata. This is reflected in similarities in leaf size and mark- ings, inflorescence shape and size and prim- ary bract orientation. The differences be- tween these taxa are presented under V. attenuata. ETYMOLOGY: This species is named in honor of Dr. Kathleen Burt-Utley who first brought this taxon to my attention. SPECIMENS EXAMINED: COSTA RICA: CARTAGO PROVINCE: Tapanti Hydroelectric Project, between 9 and 14 km southeast of Orosi, slopes above the Rio Grande de Orosi, 13 December 1974, Utley & Utley 1620 (DUKE); 20 December 1974, Utley & Utley 1728 (NOLS); 14 April 1975, Utley & Utley 3416 (DUKE); 22 March 1976, Utley & Utley 4350 (DUKE) & 4368 (MO). SAN JOSE PROVINCE: between San Jerbnimo and Alto de La Palma on Hwy. 3320, 19 June, 1975, Utley & Utley 2568 (NOLS): region between Cascajal and 8 km northeast on Hwy. 216, 10 December 1975, Utley & Utley 3616 (DUKE). 26. Vriesea notata L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 403. 1953. Thecophyllum pittieri Mez, Bull. Herb. Boiss. ser. 11. 3: 137. 1903, not Vriesea pittieri Mez, 1903 TYPE: COSTA RICA: SAN JOSE PROVINCE: La Palma, Tonduz 12526 (HOLOTYPE: B?, n.v. ISO- TYPE: US!). Epiphytic or terrestrial, to 9.0 cm caules- cent. LEAVES in a ascending rosette, 21.0- 3 1 .0 (39.0) cm long: blades 1 4.0-29.0 cm long and 1 .8-4.2 cm wide, green, usually suffused with purple or maroon, with wavy transverse lines or longitudinal stripes, these frequently maroon or brownish maroon, when occur- ing concomitantly giving a tessellate appear- ance, maroon suffusion of markings at times alternating with green in a regular pattern resulting in interrupted transverse and longi- tudinal markings; 5/7ea//25 (7.1) 1 1.5-13.8 cm long and 3.4-5.4 cm wide, castaneous adax- ially, greyish brown abaxially, becoming colored and marked like the blade distally, elliptic to oblong. SCAPE erect to curved erect, 24.0-42.5 (54.0) cm long and 0.3-0.5 cm in diameter; bracts 4.6-10.2 cm long, blades 2.0-7.1 cm long and 1.3-2.4 cm wide, green, frequently suffused with maroon or maroon markings, erect to divergent, triangular to ligulate, apex acute to acuminate, sheaths 2.5-3.7 (4.4) cm long and 2.3-4.2 cm wide, ochre, abaxially, frequently with lateral, longitudinal maroon lines ovate or broadly ovate to elliptic. INFLORESCENCE erect, 4.0-9.0 cm long and 5.5-7.0 (lO.O) cm in diameter, capitate to subcapitate or short cylindric, frequently elongating in fruit; primary bracts 4.5-7.0 cm long, blades 1.9- 3.2 cm long and 1 .3-2.0 cm wide, pale green, usually with maroon or purple longitudinal lines especially near the margins and fre- quently with maroon or purple wavy trans- verse lines, sheaths 2.1-3.5 cm long and 2.3- 4.2 cm wide, ovate to elliptic. LATERAL BRANCHES spreading to ascending, (2.5) 3.0-3.8 cm long, collaterally 2-flowered, peduncles 0.0-0.5 cm long, rachis 0.0-0. 1 cm long. FLOWERS sessile; yZora/ bracts 2.5- 3.5 cm long and 1 .9-3.2 cm wide, green usu- ally suffused with maroon or red, subcori- aceous, brown-centered lepidote adaxially, immersed lepidote or pitted abaxially, ovate to oblong-ovate or suborbicular, apex broadly acute to obtuse or rounded, occa- sionally minutely acuminate or apiculate; 74 Tulane Studies in Zoology and Botany Vol. 24 sepals 1.2-1.7 cm long and 0.9-1.4 cm wide, green occasionally suffused with red or maroon near the margin, coriaceous, brown- centered lepidote adaxially, immersed lepi- dote abaxially, elliptic to oblong-elliptic or obovate, apex obtuse to rounded; petals about 2.0-3.4 (4.0) cm long and 0.9-1.9 cm wide, white, occasionally weakly suffused with-maroon near the margins, ovate to ellip- tic or oblong-obovate, frequently subfalcate and slightly constricted medially; append- ages 0.4 cm long and 0.2 cm wide; stamens in a hood configuration over the gynoecium, anthers usually connivent by their lateral margins, filaments 1.7-3.0 cm long, anthers 0.3-0.7 (1.0) cm long; ovary 0.4-0.5 cm long and 0.3-0.5 cm in diameter, style 1.4-3.1 cm long; capsules 1 .9-2.7 cm long and 0.7-1 .0 cm in diameter, fusiform to subobovate, cas- taneous. PHENOLOGY: Flowering specimens have been collected in March through August with an apparent peak in August. DISTRIBUTION AND HABITAT: Endemic to Costa Rica between 900-2000 m in the premontane and lower montane rain forest life zones, possibly extending into the lower montane wet forest. DISCUSSION: Vriesea notata is charac- trized by leaves that are usually suffused with red or maroon and have red to maroon wavy transverse lines and longitudinal lines and nocturnal, campanulate corollas. In one population, however, individuals lacked the wavy transverse lines and only possessed colored longitudinal lines. The inflorescence of V. notata is short, densely cylindric or subcapitate at anthesis and often elongate post-anthesis. This elongation is so pro- nounced in some populations that, if the phenomenon were not observed in the field, the elongate and non-elongate forms might well be described as separate taxa. This variability in inflorescence form in V. notata is further aggravated by variation in leaf coloration and markings as well as corolla size. Further studies of variation as well as breeding systems are needed before a realistic evaluation of the relationship of V. notata and its closest relative, V. hygrome- trica, can be made. This problem is discussed further under the latter species. SPECIMENS EXAMINED: COSTARICA: ALAJUELA PROVINCE: 3-5 km west of Cariblanco on the road to Laguna Hule, 16 October 1974, Utley & Utley 757^ (DUKE); north of San Ramon, between Los Angeles Norte and 7 km north, 15 December 1974, Utley & Utley 1650 (DUKE), & 1652 (CR); 3-5 km west by west-southwest of Volcan Viejo in the vicinity of Finca Pozo Verde, 24 December 1974, Utley & Utley 1767 (DUKE, US); between Los Angeles Norte and about 7 km north of La Balsa de San Ramon, 8 August 1975, Utley & Utley 2776 (DUKE), 27«7(US), & 2785 (CR). BOR- DER OF HEREDIA AND SAN JOSE PROVINCES: on hwy. 9 between Cinco Esquinas and the junction of highways 9 and 120, 31 December 1973, Utley & Utley 583 (DUKE); vicinity of Colonia Virgen del Socorro 3-6 km east of Cariblanco, 10 August 1975, Utley & Utley 2859 (T>\}¥.E). CARTAGO PROVINCE: Tapanti Hydro- electric Project, between 9 and 14 km sou- theast of Orosi, 5 January 1974, Utley & Utley 593 (DUKE); 13 December 1974, Utley & Utley 1593 (DUKE); 20 December 1974, Utley & Utley 7729 (DUKE. MICH), 1730 (DUKE) & 7757 (F); 14 April 1975, Utley & Utley 2115 (DUKE); on the road from Tapanti to Tausito/ Selva area, 25 August 1975, Utley & Utley 3009 {D\}Y.E), 3010 (F) & 3011 (NOLS); Tapanti Hydroe- lectric Project 7-12 km south of the bridge over the Rio Grande de Orosi at Tapanti, 5 December 1975, Utley & Utley 3411 (DUKE), 3419 (CR), 3423 (MO); the north- western slopes of Volcan Irazu on Hwy. 216 beyond Cascajal, 10 December 1975, Utley & Utley 3572 A. B. & C(DUKE, MO, NY), 3600 (GH), 3618 (CAS); Tapanti Hydroe- lectric Project 7-12 km from the bridge over No.l Middle American Vrieseas 75 the Rio Grande de Orosi at Tapanti, 22 March 1976, Utley & Utley 4349 (NOLS), 4i57(DUKE),¥i25( DUKE); 7 April 1976, Utley & Utley 4544 (DU KE), & 4545 (F); 25 June 1976, Utley & Utley 5165 (CR), 5166 (DUKE), 5168iF) & 5170 (DUKE). BOR- DER OF CARTAGO AND SAN JOSE PROVINCES: near the junction of the Interamerican Highway with Hwy. 22 in the Cordillera de Talamanca south of Cartago, 16 April 1975, Utley & Utley 2180 (DUKE); about 15 km south of Cartago on the Inter- american Highway, 24 August 1975, Utley & Utley 2949 (DUKE), 2950 (DUKE), 2952 (US), 2953{CR), & 2954 (NOLS). HERE- DIA PROVINCE: region north of Cerro Chompipe or 1 5 km northeast of Heredia, 1 5 October 1974, Utley & Utley 13 14 {DVKE); between Vara Blanca and Cinchona on Hwy. 9, 16 October 1974, Utley & Utley 1332 (DUKE). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Heredia along the banks of the Rio Para Blanco and the lower slopes of Cerro Zurqui, 10 June 1974, Utley & Utley 957 (DUKE); 18 June 1974, Utley & Utley 972 (DUKE); 28 May 1976, Utley & Utley 5102 (DUKE). SAN JOS6 PRO- VINCE: between San Jerdnimo and Alto de La Palma on Hwy. 330, 15 December 1973, Utley & Utley 524 (DUKE); 16 December 1973, Utley & Utley 534 (DUKE); 12 Janu- ary 1974, Utley & Utley 607 (DUKE); 4 March 1974, Utley & Utley 685 (DVKE); 19 June 1975, Utley & Utley 2565 (DUKE), 25<5(5(DUKE), 2567 (CAS); 6 July 1975, Utley & Utley 2588 (US), 2590 (MO), 2592 (DUKE), 2594iF) 8i 2595 (DUKE); 18 August 1975, Utley & Utley 2895 (VS), 2896 (CR); between Cascajal and 8 km northeast on Hwy. 216, 23 August 1975, Utley & Utley 2913 (DUKE), 2939 (GH), 2940 (DUKE), 2941{MO), 2942 (F) & 2943 (NOLS); between 2 and 5 km southeast of Higuito on Calle Tablazo or 8 to 12 km southeast of Desamparados, 5 September 1975, Utley & Utley 3034 (DUKE), 4 km north of Cascajal on Hwy. 216, southwestern slopes of Volcan Irazu, 3 July 1976, Utley & Utley 5273 (DUKE). 27. Vriesea bracteosa (Mez & Werckle) L.B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Thecophyllum bracteosum Mez & Werckle in Mez, Fedde Rep. Spec. Nov. 14: 246. 1916. TYPE: COSTA RICA: Province not given, near La Palma, elevation 1500 m Werckle 17921 (HOLOTYPE: B!, ISOTYPE: GH, photograph US!). Epiphytic, acaulescent. LEAVES in a sub- spreading to ascending rosette, 18.0-39.0 (50.5) cm long; blades 15.0-19.0 (29.5) cm long and (3.3) 4.2-5.5 cm wide, green, fre- quently suffused with purple marginally and apically, occasionally with obscure wavy purple lines or mottled areas, ligulate or sub- ligulate to long triangular, apex acute to attenuate or rounded and acuminate, fre- quently abruptly recurved or spirally invo- lute; sheaths 15.5-23.7 cm long and 6.8-9.5 cm wide, pale brown with distal purplish areas or scattered areas of charcoal grey, elliptic to oblong-ovate. SCAPE erect or curved-erect, 36.5-48.0 cm long and 0.7-1.0 cm in diameter; bracts 5.9-9.0 cm long, blades 0.8-2.0 cm long and 0.5-1 .5 cm wide, green, frequently suffused with purple dis- tally, erect, to recurved or spirally involute, triangular, at times long, narrow triangular, apex acute to attenuate, sheaths 4.4-8.0 cm long and 3.3-3.5 cm wide, green, sheathing erect, ovate to elliptic. INFLORESCENCE erect, 10.5-13.0cmlongand (6.8)8.0-10.5 cm in diameter, densely cylindric; primary bracts 5.1-7.3 cm long, blades 0.9-1.5 cm long and 0.3-1.3 cm wide, green occasionally suffused with maroon distally, ascending, triangular to cuniform in outline, apex acute to narrowly acute or attenuate, sheaths 4.2- 6.2 cm long and 3.9-6.0 cm wide, green ovate 76 Tulane Studies in Zoology and Botany Vol. 24 to broadly ovate. LATERAL BRANCHES ascending, 3.5-5.0 cm long, collaterally or fasciculately, 2- or 3-flowered, peduncle 0.0- 0.2 cm long, rachis 0.0-0.2 cm long. FLOW- ERS sessile to 0.2 cm pedicellate; floral bracts 3.8-4.6 cm long and 3.4-5.4 cm wide, green, subcoriaceous, brown-centered lepi- dote adaxially and immersed lepidote abax- ially, broadly elliptic or ovate to orbicular frequently conduplicately folded along the pronounced keel, apex rounded; sepals 2.0- 2.5 cm long and 1 .5-2.4 cm wide, olive-green, coriaceous, brown centered lepidote adax- ially immersed lepidote to glabrous abax- ially, elliptic to oblong-elliptic or obovate, apex broadly rounded; petals 3.8-4.3 cm long and 1.6-1.8 cm wide, white, obovate; appendages 0.5-0.7 cm long and 0.2-0.3 cm wide; stamens in a hood configuration over the gynoecium, usually weakly connivent by the lateral margins of the anthers, filaments 3.0-3.5 cm long, anthers 0.9-1.0 cm long; ovary about 0.6 cm long and 0.5-0.6 cm in diameter, style 2.6-2.9 cm long; capsules 4. 1- 5.3 cm long and 1 .3-1.6 cm in diameter, cas- taneous, fusiform. PHENOLOGY: Flowering specimens have been collected in June, July and October. DISTRIBUTION AND HABITAT: Endemic to Costa Rica between 1500-1900 m in the lower montane and premontane rain forest life zones. DISCUSSION: The inflorescence of Vrie- sea bracteosa is short, dense cylindric or sub- capitate with large, coriaceous primary bracts which are more or less imbricate at anthesis. The unmarked leaves typically form an ascending rosette. Like V. capitata, the inflorescence of V. bracteosa is usually covered with a gelatinous matrix. The white, campanulate, nocturnal flowers of this taxon are exserted beyond the primary bracts. Vriesea bracteosa was originally des- cribed by Mezand Werckle(in Mez, 19 16) as having three flowers in the axils of the lower primary bracts. However, specimens have also been collected which have two flowers in the axils of the lower primary bracts. Vriesea bracteosa is undoubtedly most closely related to V. capitata, which it super- ficially resembles. These taxa share a series of common characters, including rosette form, leaf shape, inflorescence form, floral morphology and gelatinous matrix. Beyond these characters, V. bracteosa and V. capi- tata have a distinctive leaf anatomy which has not been observed in other thecophylloid vrieseas. In both species, the adaxial hypo- dermis is interrupted by a well developed layer of extrafascicular sclerenchyma. Although both taxa occur in the pass area between Volcan Barba and Volcan Irazu in the Cordillera Central, they have not been encountered growing sympatrically. Within this area V. bracteosa is typically found at slightly higher elevations ( 1 500- 1 900 m) than V. capitata (900-1400 m). Although V. bracteosa and V. capitata may be difficult to distinguish on the basis of vegetative morphology alone, they are easily distinguished from each other in flower or fruit. In addition to subtle differences in the nature of their inflorescences, V. bracteosa and V. capitata are separated by floral bract shape and length and petal length. More- over, the adaxial leaf surfaces of V. capitata are densely, cinereously pubescent, whereas those of V. bracteosa are not. Although this may be an artifact of leaf weathering which resulted in trichome loss, this character is constant in all my collections of V. bracteosa. SPECIMENS EXAMINED: COSTA RICA: CARTAGO PROVINCE: 5-6 km south of San Isidro de Cartago on the Inter- american Highway, 28 February 1975, Utley & Utley 1907 (DUKE); on the road from Tapanti to Taus and Tausito, between 1 and 4 km beyond the bridge over the Rio Grande de Orosi, 31 July 1976, Utley & Utley 5588 (MO), 5608 (DUKE) & 5609 (US). BOR- DER OF CARTAGO AND SAN JOSE PROVINCES: along the Interamerican No.l Middle American Vrieseas 11 Highway about 20 km south of Cartago, in the vicinity of Casa Mata, 17 March 1975, Utley & Utley 1996 (DUKE), 1997 (CR), 1998 (F). BORDER OF HEREDIA AND SAN JOSE PROVINCES: along the Rio Para Blanco in the vicinity of Cerro Zurqui, 30 March, 1974, Utley & Utley 72i (DUKE); 15 October 1974, Utley & Utley 1284 (NOLS). SAN JOSE PROVINCE: between San Jeronimo and Alto de La Palma on Hwy. 220, 29 June 1974, Utley & Utley 979 (DUKE); 19 June 1975, Utley & Utley 2562 (DUKE); between Cascajal and 8 km nor- theast on Hwy. 216, 23 August 1975, Utley & Utley 2924 (NOLS); 7-10 km southeast of Desamparados in the Altos de Tablazo, 2 July 1976, Utley & Utley 5233 (DUKE); at Cascajal about 3 km north of Las Nubes on Hwy. 216, the northwest slopes of Volcan Irazu, 3 July 1976, Utley & Utley 5248 (NY) & 5272 (DUKE). 28. Vriesea capitata (Mez & Werckle) L. B. Smith & Pitt. J. Wash. Acad. Sci. 43: 402. 1953. Thecophyllum capitatum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 873. 1904. TYPE: COSTA RICA: no location given, Werckle Bromel. Costaric. 86 (HOLOTYPE: B!, photograph US!). Epiphytic, acaulescent to 5.5 cm caulescent. LEAVES in an ascending to suberect rosette, 23.7^0.0 cm long; blades 11.5-17.5 cm long and 3.6-4.5 cm wide, green fre- quently suffused with purple or maroon dis- tally, ligulate apex acute to rounded and apiculate; sheaths 1 2.0-1 7.0 cm long and 5.5- 7.0 cm wide, castaneous abaxially, brown adaxially, margins suffused with purple to maroon distally. SCAPE erect, 22.5-35.0 cm long and 0.4-1.0 cm in diameter; bracts 4.2- 7.6 cm long, blades 1 .2-2.6 cm long and 1 . 1- 2.0 cm wide, green drying to brown, erect or frequently recurved, triangular or short ligu- late, apex attenuate to acute or rounded and abruptly acuminate, sheaths 2.6-5.0 cm long and 2. 1-3.3 cm wide, drying ochre or brown castaneous proximally, sheathing erect, ovate to elliptic or oblong elliptic. INFLO- RESCENCE erect, 3.5-6.5 cm long and 4.0- 6.5 cm in diameter, subcapitate; primary bracts 2.3-3.8 cm long, blade 0.7-1. 4 cm long and 0.5-1 .4. cm wide, green becoming brown at anthesis, spreading, triangular, apex acute, sheaths 1 .9-2.9 cm long and 2.2-2.6 cm wide, green elliptic to ovate in outline. LAT- ERAL BRANCHES ascending, 3.8-5.2 cm long, subcollaterally 2-flowered, peduncle 0.1-0.3 cm long rachis about 0.4 cm long. FLOWERS sessile, yZora/ bracts 2.0-2.2 cm long and 1.3-1.6 cm wide, green subcoriace- ous, brown-centered lepidote adaxially, gla- brous abaxially, oblong to obovate, un- keeled or very weakly keeled distally, apex obtuse; sepals 2.0-2.2 cm long and 1 .3-1 .6 cm wide, olive green, coriaceous, brown-cen- tered lepidote adaxially, glabrous abaxially, elliptic to obovate, apex obtuse;/7e/a/5 about 3.1-3.4 cm long and 1.5-1.6 cm wide, white, obovate; appendages 0.4 cm long and 0.2- 0.25 cm wide; stamens in a hood config- uration over the gynoecium, filaments about 2.3 cm long anthers about 0.7 cm long; ovary about 0.7 cm long and 0.5 cm in diameter, style 1 .7 cm long; capsules 4. 1 cm long and 0.8-1 .0 cm in diameter, casteneous, fusiform. PHENOLOGY: Flowering plants have been collected only during August. DISTRIBUTION AND HABITAT: Known from Costa Rica and a few recent collections from Cerro Jefe in Panama. Col- lected between 900-1400 m in premontane rain forest life zone possibly extending slightly into the premontane wet forest life zone. DISCUSSION: Vriesea capitata remains a poorly collected taxon. In locations where I have collected this species, 1 have never seen more than one or two individuals in flower or fruit. It is characterized by its rigidly cori- aceous leaves in a subascending to suberect 78 Tulane Studies in Zoology and Botany Vol. 24 rosette, subcoriaceous primary bracts which are divergent to subspreading at anthesis, and its subcapitate inflorescence. The inflo- rescence of V. capitata is often totally covered with a gelatinous exudate. The developing floral buds are immersed in this gelatinous matrix until anthesis, at which time the white, campanulate nocturnal flow- ers emerge. When the atmospheric humidity is low, this exudate will dry on the flower buds and inhibit anthesis. Vriesea capitata is most closely related to V. bracteosa and the characters which distin- guish these taxa are discussed under the lat- ter species. SPECIMENS EXAMINED: COSTA RICA. ALAJUELA PROVINCE: along road between Hwy. 9 and Laguna Hule between 0.5-5.0 km northwest of Carib- lanco, 16 October 1974, Utley & Utley 1368 (DUKE); north of San Ramon, between Los Angeles Norte and about 7 km north, 15 December \914,Utley & Utley 1635 (CR) & 7<55/ (DUKE). BORDER OF ALAJUELA AND HEREDIA PROVINCES: vicinity of Colonia Virgen del Socx)rro about 3-6 km east of Cariblanco, 10 August 1975, Utley & Utley 2878 (DUKE). BORDER OF HEREDIA AND SAN JOSE PROVINCES: about 5 km northeast of San Isidro de Here- dia on Calle Yerbabuena, 3 April 1976, Utley & Utley 4475 (NOLS) & 4477 (DUKE). PANAMA: PANAMA PROVINCE: Cerro Jefe, December 1974, Mori & Kal- lunki 3619 (MO), 18 December 1974, Mori & Kallunki 3800 (MO). DOUBTFUL AND EXCLUDED SPECIES Vriesea discolor (Mez & Werckle) L.B. Smith & Pitt. J. Wash. Acad. 43: 402. 1953. Thecophyllum discolor Mez & Werckle in Mez, Fedde, Rep. Spec. Nov. 14: 246. 1916. TYPE: COSTA RICA: without further location, "Werckle." Mez (1916) did not cite a specific collection when he described Thecophyllum discolor. Moreover, no specimen of this taxon is on deposit at the Berlin-Dahlem Herbarium, which acquired Mez's personal collection of Bromeliaceae in 1936. The original diagnosis of Thecophyllum discolor agrees with the description and type collection of the earlier Thecophyllum latis- sima, but the taxa reportedly differ in the number of flowers per lateral inflorescence branch (four or five in Th. latissimum versus up to 12 in Th. discolor). The only putative material of V. discolor I have seen is Werckle 72 (US) which has 4 to 6 flowers per lateral inflorescence branch and is undoubtedly V. latissima. In all likelihood V. discolor is a synonym of V. latissima but until authentic material is available the former should be maintained as a doubtful species. Vriesea spectabilis (Mez & Werckle) L.B. Smith 8l Pitt. J. Wash. Acad. 43: 403. 1953. Thecophyllum spectabile Mez & Werckle in Mez, Bull. Herb. Boiss. ser. II. 4: 873. 1904. TYPE: COSTA RICA: without further locality, Werckle s.n. (HOLOTYPE: B!, photograph US!). Recent flowering collections have shown that this taxon has connate sepals and petals and the species has been transferred to Guz- mflma (Utley, 1978). Vriesea splitgerberi (Mez) L.B. Smith & Pitt. J. Wash. Acad. 43: 403. 1953. Guzmania splitgerberi Mez in DC. Monogr. Phan. 9: 930. 1896. TYPE: SURINAME: Blauwe Berg, upper Suriname River, Splitgerber966 (HOLOTYPE: L!, photograph US!). Thecophyllum splitgerberi (Mez) Pitt. Evolution 2: 60. 1948. No.l Middle American Vrieseas 79 This species displays the thecophylloid syn- drome of reduced lateral branches and en- larged primary bracts. However, V. splitger- beri differs from all other thecophylloid vrieseas in its polystichous lateral inflores- cence branches. Moreover, the stigmatic sur- face consists of three, solid, cylindrical branches (Fig. 15a) rather than the 3-cupped stigmatic structure typical of the thecophyl- loid vrieseas and their allies. In light of this I feel that this species is not closely allied to the thecophylloid vrieseas and I have excluded it from this treatment. Vriesea vittata (Mez & Werckle) L.B. Smith & Pitt. J. Wash. Acad. 43: 403. 1953. Thecophyllum vittatum Mez & Werckle in Mez, Bull. Herb. Boiss. ser. 11.4:871. 1904. TYPE: COSTA RICA: without exact locality, Werckle Brom. Costar. 79 (HOLOTYPE: B!, photograph US!). A through examination of the type material of this species failed to reveal any evidence of two bracts subtending the single flowers. Moreover, numerous collections which are essentially identical to the type of V. vittata also have simple inflorescences with flowers subtended by single bracts rather than two bracts as stated by Mez. The affinities of this species seem to be with the thecophylloid allies (cf.p. 18 ) and not the taxa with com- pound inflorescences considered in this treatment. ACKNOWLEDGEMENTS This paper is based in part on a Ph.D. dissertation submitted to the Department of Botany, Duke University, Durham, North Carolina. The study was supported by the National Science Foundation Grant No. GB-23200 to the Department of Botany, Duke University, a Smithsonian Research Fellowship and by a University of New Orleans Research Council Grant. The Field Museum of Natural History, Missouri Botanical Garden, Museo Nacional de Costa Rica and The Organization for Tropi- cal Studies provided invaluable logistic sup- port during field work. My thanks are also expressed to the following individuals who contributed to the success of this study: Dr. Frank Almeda, Janice Bittner, Dr. Tom Croat, Sue Dickerson, Janet Leslie Forbes, Dr. Amy Jean Gilmartin, Luis Diego Gomez, Marie LeBlanc, Dr. James Luteyn, Dr. Joan Nowicke, Dr. Robert Read, Dr. Harold Robinson, Dr. Lyman B. Smith, Dr. F. Gary Stiles, Dr. D. E. Stone and Dr. R. L. Wilbur. Special thanks are due Dr. Kathleen Burt-Utley who was my constant field com- panion and who prepared the illustrations for this work. I am indebted to the curators of the follow- ing herbaria who loaned material for this study: Botanischer Garten und Botanisches Museum Berlin-Dahlem (B); Botanical Museum and Herbarium, University of Copenhagen (C); Duke University Herba- rium, Durham (DUKE); Gray Herbarium of Harvard University, Cambridge (GH); Systematisch-Geobotanisches Institut, Uni- versitat Gottingen (GOET); The Herbarium and Library, Royal Botanic Gardens, Kew (K); Rijksherbarium, Leiden (L); Botanische Staatssammlung, Munich (M); Herbarium of the University of Wisconsin, Madison (WIS). In addition to the loan of specimens I am indebted to the following herbaria and their curators for generously making their facilities available to me: Herbarium, Cali- fornia Academy of Sciences, San Francisco (CAS); Herbario Nacional, Museo Nacional de Costa Rica, San Jos^ (CR), Field Museum of Natural History, Chicago (F); Herbario Nacional, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico DF (MEXU); Herbarium of the Missouri Botanical Garden, St. Louis (MO); U.S. National Herbarium, Smithsonian Institution Washington DC (US). 80 Tulane Studies in Zoology and Botany Vol. 24 RESUMEN Los thecophylloid vrieseas habian sido mantenidos por mucho tiempo en un gdnero distinto, Thecophyllum, por razdn de sus brdcteas agrandadas y ramas florescentes secundarias reducidas. Ahora, estas especies forman una taxa de alianza cohesiva geogrd- ficamente y morfdlogicamente dentro de las seccidn Xiphion de Vriesea. Aparte de las flores nocturnales que generalmente se encuentran en la seccidn Xiphion, varios the- cophylloid vrieseas demuestran un periodo crepuscular y diurno y sindromes florales que sugieren una gran adaptacidn de vec- tores de polen. Claves, descripcidn y sino- nomias se d^ para la taxa que pertenece a Mexico y Am^ica Central y su afinidad es discutida. Vriesea greenbergii, V. kathyae, V. luis-gomezii y V. lyman-smithii han sido ilustradas y descritas como especies nuevas. LITERATURE CITED Andre, E, 1889. Bromeliaceae Andreanae. Description et histoirede bromeliacees. Librarie Agricole, Paris. Baker, H. G. 1961. The adaptation of flowering plants to nocturnal and crepuscular pollinators. Quart. Rev. Biol. 36: 64-73. Benzing, D. H. 1976. Bromeliad trichomes; structure, function and ecological significance. Selbyana 1: 330-348. Brewbaker, J. L. and D. D. Gorrez. 1967. Genetics of self-incompatibiality in the monocot genera. Ana- nas (pineapple) and Gasieria. Amer. J. Bot. 54: 611-616. Dolzmann, P, 1964. Elektronenmikroskopische unter suchunger an den saughaaren von Tillandsia us- neoides (Bromeliaceae) 1. feinstruktur der kuppel- zelle. Planta 60: 46M72. Downs, R. J. 1974. Anatomy and physiology. In: Pit- cairnioideae (Bromeliaceae) (Flora Neotropica, monograph no. 14). L. B. Smith and R. J. Downs, pp. 2-28. Hafner Press, New York. Ehler, N. 1977. Bromelienstudien II. Neueentersuchun- gen zur entwicklung, struktur und funktion der bromelien trichome. (Tropische und subtropische Pflanzenweh, 20). Akademie der Wissenschaften und der literatur, Mainz in Kommission Bei Franz Steiner Verlag GMBH, Weisbaden. pollination ecology, 2nd rev. ed. Pergamon Press, New York. Gilmartin, A. J. 1973. Transandean distribution of Bromeliaceae in Ecuador. Ecology 54: 1389-1393. Grant, K. and V. Grant. 1968. Hummingbirds and their flowers. Columbia University Press, New York. Holdridge, L. 1967. Life zone ecology. Tropical Science Center, San Jose, Costa Rica. Holdridge, L., W. C. Grenke, W. H. Hatheway, T. Liang and J. A. Tosi. 1971. Forest environments in tropical life zones: a pilot study. Pergamon Press, New York. Mc Williams, E. L. 1974. Evolutionary ecology. In: Pit- cairnioideae (Bromeliaceae) (Flora Neotropica, monograph no. 14). L. B. Smith and R. J. Downs, pp. 40-55. Hafner Press, New York. Mez, C. 1896. Bromeliaceae. In: C. de Candolle, Monographic phanerogamarum 9: 1-990. Sumpti- bus Masson & Cie., Paris. 1903a. Additamenta monographica 1903. Bull. Herb. Boiss. ser. II 3: 130-146. 1903b. Additamenta monographica 1903. Bull. Herb. Boiss. ser. II 3: 224-228. 1904a. Additamenta monographica 1904. Bull. Herb. Boiss. ser. II 4: 619-634. 1904b. Additamenta monographica 1904. Bull. Herb. Boiss. ser. II 4: 863-878. 1904c. Additamenta monographica 1904. Bull. Herb. Boiss. ser. II 4: 1121-1136. 1905a. Additamenta monographica 1904. Bull. Herb. Boiss. ser. II 5: 100-116. 1905b. Additamenta monographica 1904. Bull. Herb. Boiss. ser. II 5: 232-233. 1916. Additamenta monographica 1916. Repert. Spec. Nov. Regni. Veg. 14: 241-256. 1934-1935. Bromeliaceae. In: Engler, Das Pflanzenreich IV (32): 1-667. Wilhelm Engelmann, Berlin. Pittendrigh, C. S. 1948. The bromeliad-anopheles- malaria complex in Trinidad. 1-the bromeliad flora. Evolution 2: 58-89. Porsch, O. 1932. Das problem fledermausblume. Akad. Wiss.Wien, mth. naturwiss. Kl., Anz. 69: 27-28. Robinson. H. 1969. A monograph on foliar anatomy of the genera Connellia, Cottendorfia and Navia (Bromeliaceae). Smithsonian Contrib. Bot. no. 2. Salas, D. S. 1973. Una Bromeliaceae costarricense poli- nizada por murcielagos. Brenesia 2: 5-10. Faegri, K. and L. van der Pijl. 1971. The principles of Skog, L. 1976. A study of the tribe Gesnerieae with a No.l Middle American Vrieseas 81 revision of Gesneria (Gesneriaceae: Gesnerioideae). Smithsonian Contrib. Bot. no. 29. Smith, L. B. 1932. Studies in the Bromeliaceae, III. Contr. Gray Herb. 98: 3-35. .1938. Bromeliaceae. In: North America Flora 19 (2): 61-228. .1941. Studies in the Bromeliaceae. XII. Lilloa6:38M17. .1963. Notes on Bromeliaceae, XX. Phyto- logia 9: 242-261. .1966. Notes on Bromeliaceae, XXIII. Phytologia 13: 84-161. .1974. Notes on Bromeliaceae, XXXVl. Phytologia 28: 319-333. and R. J. Downs. 1977. Tillandsioideae (bromeliaceae) (Flora Neotropica monograph no. 14, part 2) Hafner Press, New York. and C. Pittendrigh. 1955. Realignments in the Bromeliaceae subfamily Tillandsioideae. J. Wash. Acad. 43: 63 & 401-404. Susessenguth, K. 1942. Neue pflanzen aus Costa Rica, insbesondere vom Chirripo Grande 3837 m. Bot. Jahrb. 72: 270-302. Szidat, L. 1922. Die samen der bromeliaceen in ihrer anpassungan den epiphtismus. Bot. Arch. 1: 29-46. Tomlinson, P. B. 1969. III. Commelinales-Zingiberales. In: Metcalfe, C.R., Ed. Anatomy of the monocoty- ledons. Oxford, Clarendon Press. Utley, J. F. 1978. A new combination in Guzmania (Bromeliaceae). Phytologia 40: 55-57. Walter, H. 1971. Ecology of tropical and subtropical vegetation. Oliver and Boyd, Edinburgh. Volume 24, No. 2 $4.00 July 20, 1983 FISHES OF THE BAYOU SARA DRAINAGE, LOUISIANA AND MISSISSIPPI, WITH A DISCRIMINANT FUNCTIONS ANALYSIS OF FACTORS INFLUENCING SPECIES p. 83 DISTRIBUTION JAMES M. GRADY , ROBERT C. CASHNER AND JAMES S. ROGERS PLANTS AND PLANT COMMUNITIES OF JEAN LAFITTE NATIONAL HISTORICAL PARK, LOUISIANA. DAVID A. WHITE, STEVEN P. DARWIN, AND LEONARD B. THIEN pp. 101 OMP. ZOOL LfBRARY TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. 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Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University." Subscription rates: Volume 24, $8.50 domestic, $9.50 foreign Copies of Tulane Studiesin Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 24 , No. 2 July 20, 1983 FISHES OF THE BAYOU SARA DRAINAGE, LOUISIANA AND MISSISSIPPI, WITH A DISCRIMINANT FUNCTIONS ANALYSIS OF FACTORS INFLUENCING SPECIES DISTRIBUTION JAMES M. GRADY', ROBERT C. CASHNER AND JAMES S. ROGERS University of New Orleans Department of Biological Sciences, New Orleans, Louisiana 70148 Abstract A fish survey of Bayou Sara, a southward flowing tributary to the lower Mississippi River in Mississippi and Louisiana, was con- ducted between June 1976 and May 1979. Eighty species, representing 17 families are known from the system as a result of survey work in the lower reaches by Louisiana State Univer- sity, Baton Rouge, and our work in the upper and middle sections of Bayou Sara. Bayou Sara has fewer species than the adjacent Thompson Creek and nearby westward flow- ing Buffalo Bayou. The species composition is notable in comparison to the other two systems by the absence of common species of petromyzontids, ictalurids and cyprinodon- tids. Of the families shared by the three drainages. Bayou Sara most closely resembles Thompson Creek. Distributional data from six permanent collection stations were supplemented with measurements for 17 environmental para- meters. The combined data were subjected to a multiple discriminant functions analysis to identify significant environmental variables accounting for the separation of fish dis- tributions. A single discriminant function, principally contributed to by stream position, accounted for 79% of the variance among fish distributions. 'Present address: Department of Zoology, Southern Illinois University Carbondale, Illinois 62901 Introduction The eastern embayment of the lower Mississippi River receives several small to moderate-sized tributaries in southwest- ern Mississippi and southeastern Louisi- ana. The major westward flowing tribu- taries are Big Black River, Bayou Pierre, Coles Creek, and Homochitto and Buffalo rivers. The lowermost tributaries are Tunica Bayou, Bayou Sara and Thompson Creek (Fig. 1). This region of the lower Mississippi River drainage, the tributaries to the Lake Pontchartrain estuary, and the Pearl River drainage have been extensively surveyed by R.D. Suttkus (Tulane Uni- versity), N.H. Douglas (Northeast Louisi- ana University), J.V. Conner (Louisiana State University), and their students, on either a regular or sporadic basis for the past 30 years. In addition, intensive fish faunal surveys have been conducted in Thompson Creek (Guillory, 1981, 1982), Buffalo Bayou (Cashner et al., 1976), Tchefuncte River (Saul, 1974) and Amite Editorial Committee for this Paper: Dr. Andre M. Landry, Associate Professor, Department of Marine Biology, Texas A & M University at Galveston, Galveston, Texas 77553 Dr. William J. Matthews, Assistant Professor, University of Oklahoma Biological Station, Star Route B, Kingston, Oklahoma 73439 83 84 Tiilane Studies in Zoology and Botany Vol. 24 River (Laiche, 1980). Douglas (1974) provided comprehensive distributional data for many species in the state. Zoogeographically the lower Missis- sippi embayment eastward to the Pearl River yields a number of distinctive distributional patterns for its indigenous species, as first noted by Guillory and Conner (1973), and discussed in detail by Guillory (1982). Bayou Sara is one of the southernmost tributaries to the eastern embayment of the Mississippi River (Fig. 1) and is cen- trally located in this area of engimatic distributions. In an effort to clarify some species distributions, a study of the ichthyofauna of the Bayou Sara system was initiated in June 1977 and included monthly sampling from the headwaters to the mouth until March 1979. Although numerous collections had been made in the system by R.D. Suttkus and J.H. Caruso, a comprehensive survey of the Bayou Sara fishes has not been previously reported. Figure 1. Bayou Sara study area and adjacent drainages. A second aspect of this study centered on a physico-chemical characterization of Bayou Sara and an attempt to determine associations between environmental para- meters and observed patterns of fish dis- tribution. A number of statistical pro- cedures have been utilized in attempts to produce reliable quantitative associations between physical and chemical factors and the distribution of fishes (Smith and Fisher, 1970; Smith and Powell, 1971; Moyle and Nichols, 1973; Wallace, 1973; Stevenson et al., 1974; Heins and Clemmer, 1975; Rogers et al., 1976; Sobczak, 1976; Baker and Ross, 1981). Discriminant functions analysis was selected to identify significant environmental parameters influencing instream fish distributions in Bayou Sara. Description Of Study Area Bayou Sara originates in Wilkinson Co., Mississippi, west of Woodville. The stream flows in a southward direction for 56.6 km before entering the Mississippi River near St. Francisville, Louisiana, at river mile 266. It drains approximately 493 km^ including portions of Wilkinson Co., Mississippi and West Feliciana Parish, Louisiana. Bayou Sara traverses several geologic formations along its course. In headwater regions, the stream cuts through and exposes the Citronelle formation, a Plio- cene-Pleistocene alluvial formation char- acterized by coarse sand, silt, and gravel (Fisk et al., 1938; Fisk, 1944; Murray, 1961; Sibley, 1972). Partial erosion of the Citronelle has exposed brightly colored clays and coarse gravels of older Tertiary formations at some headwater locations. Sand substrates predominate in the headwater section of Bayou Sara, but gravel and hard packed clay are conspi- cuous in riffle areas. Turbidity is the lowest in the headwaters, although or- ganic debris is abundant during several No. 2 Bayou Sara Fishes 85 months of the year. Terrestrial vegetation providing stream cover and allocthonous detritus is heaviest in this region of Bayou Sara. A mixed hardwood and coniferous forest with sweetgum {Liquidambar styra- ciflua), basswood {Tilia sp.), water oak {Quercus nigra), short leaf pine (Pinus echinata), and loblolly pine {P. taeda) is the dominant community type (Caplenor et al., 1968). Deep cut banks are prevalent in headwater areas where Bayou Sara occupies a narrow channel (0.5-1.0 m) with alternating pools and riffles. Further downstream, Bayou Sara encounters four or five broad step-like depositional surfaces, the Pleistocene Terraces (Fisk et al., 1938; Fisk, 1944; Murray, 1961). Soft, shifting sand sub- strates with patches of fine gravel are characteristic of the terrace areas. Current velocity decreases and tubidity increases in this region of alternating sand bottomed runs and pools. A small portion of lower Bayou Sara cuts through the Recent Mississippi River flood plain (Fisk et al., 1938). This Recent flood plain is separated from the Pleisto- cene Terraces by the eastern escarpment of the Mississippi River. Here water levels and other environmental conditions reflect the influence of the Mississippi River. The lower reaches are character- ized by large accumulations of silt, slight current, and highly turbid waters. Bayou Sara occupies a broad, shallow basin with depths ranging from 15 cm to 1.5 m and widths ranging from 25 to 60 m during periods of low flow in the Mississippi River. In the extreme headwaters many small tributaries are occasionally intermittent during summer and early fall. The main channel, however, is spring-fed and has permanent flow year round. Water levels along the watercourse fluctuate with the seasonality of rainfall and prevailing conditions in the Mississippi River. Stream flow is heaviest during winter and spring, which are the usual periods of heavy rainfall (Snows, 1970). Materials And Methods A. Physico-chemical Analysis Water quality determinations, physical measurements and fish collections were taken at selected stations along the main channel of Bayou Sara from February, 1978 through March, 1979. Five stations (Stations 1-5 in Fig. 2) were initially selected for monthly physico-chemical measurements. However, time limita- tions, due to concurrent fish sampling, dictated an alternative sampling pro- cedure. Three stations were visited one month and the remaining two the next. This system was followed for three months, but then modified so that 17 environmental parameters and fish collec- tions were taken at all five stations bi- monthly. Due to high water and flooding, Station 2 became inaccessible after April, Figure 2. Bayou Sara study area, indicating loca- tions of the 6 permanent stations (triangles) and the 27 supplemental sampling stations (circles). 86 Tulane Studies in Zoology and Botany Vol. 24 1978. The close proximity of Station 3 lessened the loss of this sampling site. A downstream site, Station 6, was added in December, 1978. Conductivity, dissolved oxygen, and water temperature were measured in the field with a YSI Model 33 S-C-T Meter and a YSI Model 5 IB Oxygen Meter. Additional field measurements included the titrimetric determination of free carbon dioxide and alkahnity, and the colorometric measurement of pH utilizing a Rascher and Betzold Colorometric kit. The remaining chemical analyses, turbity, suspended soHds, and total sohds, were conducted in the laboratory. Turbidity measurements were made with a Hellige Turbidimeter and evaporation and ashing techniques were used for total and sus- pended sohds determinations. U.S. Geological Survey topographic maps (1:24,000) were used to determine stream position, stream order, gradient and altitude. Stream order was calculated according to Horton's (1945) method as modified by Strahler (1957). Measure- ments of mean depth, mean width and current velocity were made according to Lagler (1956). Cross-sectional area was computed following Sobczak (1976). B. Fish Collections A survey of the Bayou Sara system was begun in June, 1977 and continued until December, 1977. Fish collections were made concurrently with physico-chemical measurements at five or six of the permanent stations from February, 1978 to March, 1979. Supplemental collec- tions, in addition to those made in 1977, were made at localities throughout the Bayou Sara system to establish as com- plete a faunal hst as possible. Collection sites are shown in Fig. 3. Fishes were collected with 3.3 m, 4.8 mm nylon mesh seines and originally preserved in 10% formalin. C. Data Analysis A multiple discriminant functions analysis subroutine of Nie et al. (1975) was employed to identify environmental parameters contributing to the spatial separation of species. The value of this technique was discussed by Green (1971). Both direct and stepwise methods of discriminant analysis were utilized in this study. The direct method enters all of the independent variables into the analysis concurrently. The discriminant functions are then fitted directly from the entire set of variables. Stepwise discriminant func- tions analysis permits the exclusion of variables containing little information. Environmental variables are selected for inclusion into the analysis on the basis of their discriminating power. The criterion for establishing discriminating power in this analysis was Rao's V, a generalized distance measure. Those variables which contributed to the largest increase in V, when added to the previous variables, were selected. Forty-three groups representing the 43 species collected at the permanent stations were designated for input into the analysis. Mean discriminant scores were generated for each of these groups based on the seventeen parameters (13 field and lab measurements plus 4 stream features calculated from topographic maps) measured at the time each species was " 386^3271116 12 3 3 2 37 I 21 10 23 29 19 18 36U33I541 34 28 204235 9 26 40 24 22 , T ^\JVJ V W^rr T T T , WMT ^^^l^ \T ttt Figure 3. Mean discriminant scores for the first discriminant function (DF I) of the stepwise analysis. No. 2 Bavou Sara Fishes 87 captured. Several modifications were introduced following the initial analysis. A minimum inclusion level for each group was estab- lished following Green (1971). Only those species present in at least two collections at permanent stations were included in sub- sequent analyses, thereby reducing the number of groups to 33. Because a uniform sampling technique was not em- ployed at each site for the collection of fish, data sets were not weighted by species abundance. Results A. Physico-chemical Analysis Means, ranges and standard deviations of thirteen of the physical and chemical measurements from each of the six stations are presented in Table 1. Because stream position, gradient, altitude and stream order are relatively static, only the observed values are presented in Table 2. B. Distribution of Fishes A total of 42,867 specimens was taken in 65 collections at 33 localities in the Bayou Sara system (Fig. 2). Forty-three species, representing 13 families and 27 genera, were collected during the two year study and records for nine more species were ob- tained from Douglas (1974) and from collection records at the Tulane Museum of Natural History. John Van Conner kindly provided records of an additional 28 species taken primarily in the lower part of the system during surveys by personnel in the Department of Fisheries, Louisiana State University, Baton Rouge. These rec- cords include four species represented only by larvae and at least eight species that could potentially range into the main por- tion of our study area. A complete list of all 80 species from 17 families documented from the Bayou Sara system is presented in Table 3. Nomenclature given follows Robins et al. (1980). C. Data Analysis A maximum of 17 discriminant functions (DF) was extracted during the direct analy- sis, five of which accounted for 70% of the among-species variance (Table 4). Stan- dardized DF coefficients for the 17 environmental variables for these five DF are also presented in Table 4. The magni- tude of the coefficients indicates the relative contribution of each variable to the formation of the DF. Thus altitude and stream position provided the major contri- butions toward the separation of groups in DF I, which accounted for slightly over 30% of the among-species variance. These two physical parameters remained the most significant factors contributing to the overall separation of the groups (species) in the remaining 16 DF. Total solids and stream order were the only other variables contributing to group separation in all of the 17 DF. Comparisons of the magnitude of the DF coefficients for these para- meters, with those for altitude and stream position, reveal, however, that their con- tributions are only minor. None of the DF extracted during the direct analysis reached a high degree of statistical significance (p>.05). The distributional data and associated environmental parameters were also sub- jected to a stepwise DF analysis. Two variables, water temperature and stream position, were selected for inclusion into the analysis based on their discriminating power as indicated by a significant increase in Rao's V (Table 5). Discriminant function I accounted for almost 79% of the among - species variance and was significant (p< .01). The analysis weighted most heavily on stream position. Although DF II accounted for over 21% of the among-species variance, it failed to attain as high a degree of significance (p> .05). 88 Tulane Studies in Zoology and Botany Vol. 24 TABLE 1. Means, Ranges, and Standard Deviations of 13 Physico-Chemical Parameters at the 6 Permanent Stations in Bayou Sara Station Total Alkalini' (mg/1) ty Free CO, (mg/1)^ PH Conductivity 2 ( mhos/cm ) Total Solids (mg/l) Suspended Solids (mg/1) Turbidity (mg/1) 1 35^ 9.00 7.01 75 91.04 24.45 2.63 20*^ 7.14 0.16 61 87.80 25.39 4.77 14-72^^ 1.60-22.0 6 .80-7.30 20-215 28.15-293.46 1.31-60.11 0.00-14.00 2 37 1.40 7.13 53 48.65 14.35 0.33 29 0.17 0.06 5 28.29 1.80 0.29 20-71 1.20-1.50 7 .10-7.20 50-59 32.30-81.32 13.31-16.43 0.00-0.50 3 53 1.19 8.03 82 74.41 29.17 2.84 21 1.58 0.55 26 36.52 40.57 4.38 22-78 0.00-3.50 7 .00-8.60 40-117 42.23-148.52 3.31-103.31 0.00-11.70 4 46 5.70 7.33 95 180.29 90.30 17.91 16 5.05 0.16 40 180.40 123.10 17.91 28-58 0.00-10.90 7. ,00-7.50 40-180 60.00-582.10 7.50-362.45 0.30-55.30 5 43 8.93 7.28 98 84.27 18.09 5.55 11 7.86 0.15 27 11.76 24.58 5.87 33-60 2.40-23.90 7. ,10-7.50 71-143 66.66-94.72 3.37-67.42 0.00-12.40 6 59 11.25 7.50 91 92.84 28.34 0.20 13 1.77 0.14 1 17.81 26.64 0.28 50-68 10.00-12.50 7. 40-7.60 90-92 80.25-105.43 9.50-47.18 0.00-0.40 a-mean; b-standard deviation; c-range Discussion A. Physico-chemical Factors Progressional changes in the 17 environ- mental parameters are evident in Table 1. Values for pH, alkalinity, conductivity, total and suspended solids, turbidity, tem- perature, mean depth, mean width and cross-sectional area increase downstream. These successively higher values can be at- tributed to the expected reduction of cur- rent velocity and greater accumulations of allocthonous material. Additionally, the progressive downstream rise in temperature is also influenced by the predominance of sand substrates and a reduction of cover associated with reduced riparian vegetation. Spatial differences in some of the variables C£in be attributed to the influence of localized conditions. High levels of free CO2 and correspondingly lower pH values at Station 1 are due in part to increased deposition of organic material deposited in this area. Turbidity, conductivity, and dis- solved and suspended solids at Station 1 were relatively high for an upstream loca- tion. Heavy detrital input partially accounts for these high values, but a fallen tree dam acted to further increase the readings. The obstruction also caused higher values for mean depth and width, and cross-sectional area, while decreasing flow rates. Localized conditions also caused high readings for turbidity, suspended and total solids, and conductivity at Station 4, where a gravel washing operation dramatically increased the suspended particulate matter. At Station 3 the possible presence of lime deposits upstream could explain the con- sistently high pH and low CO2 values. Overall, the deviations from the observed progressional trends from stations 2-6 are probably the result of reduced sampling effort. Still, despite the influence of No. 2 Bavoii Sara Fishes 89 TABLE CONT. Dissolved Oxygen (mg/1) Temperature (°C) Current (m/sec) Mean Depth (m) Mean Width (m) Cross-sectional Area (m2) 9.73 18.86 0.38 0.13 5.49 0.72 2.62 7.26 0.27 0.07 1.13 0.41 7.20-14.40 10.20-28.00 0.12-0.77 0.07-0.27 3.53-6.82 0.3-1.59 13.23 14.27 0.21 0.10 11.80 1.19 0.95 7.39 0.09 0.01 2.69 0.35 12.50-14.30 10.00-22.80 0.16-0.31 0.10-0.11 10.25-14.90 0.99-1.60 9.89 20.63 0.60 0.15 22.98 3.56 1.76 8.75 0.34 0.04 7.39 1.46 7.70-12.10 10.30-29.50 0.18-1.23 0.11-0.20 7.73-31.25 0.91-5.10 8.51 26.76 0.24 0.34 23.98 8.26 2.18 11.58 0.16 0.11 10.15 5.74 6.50-11.50 10.10-37.80 0.14-0.55 0.18-0.45 17.63-46.15 4.05-20.94 8.75 27.15 0.27 0.24 32.00 7.34 1.48 7.75 0.10 0.09 7.10 2.54 7.70-10.80 14.40-34.00 0.16-0.42 0.12-0.32 23.50-40.50 3.97-10.31 9.45 19.75 0.44 0.12 32.82 3.98 1.77 6.72 0.06 0.03 0.78 1.20 8.20-10.70 15.00-24.50 0.39-0.48 0.10-0.14 32.27-33.37 3.13-4.82 localized conditions and sampling error, measurements for the 17 variables fall within the ranges for those reported from comparable stations in adjacent Buffalo Bayou (Rogers et al., 1976) and Thompson Creek (Guillory, 1974). B. Fish Distributions Intersystem Distribution. Bayou Sara, like Thompson Creek, has a more depauperate fauna than the larger, westward flowing tributaries to the eastern Mississippi Em- bayment and adjacent Lake Pontchartrain TABLE 2, Stream Position, Gradient, Altitude, and Stream Order at Six Physico-chemical Stations. Station Stream Position Gradient Altitude Stream Order (km) (m/km) (m) 59.29 5.33 79.26 3.00 42.00 1.54 54.86 4.00 36.35 1.54 42.67 5.00 16.29 0.76 18.29 5.00 10.50 0.76 12.19 5.00 7.93 1.14 9.14 5.00 90 Tiilane Studies in Zoology and Botany Vol. 24 TABLE 3. List of Species known to occur in Bayou Sara Drainage, Mississippi and Louisiana. (X= collected during present study; D= record from Douglas, 1974; TU= Tulane University Museum of Natural History Rec- ord; L= LSD record; *= likelv to occur in study area). SPECIES SOyRCE__ LEPISOSTEIDAE Lepisosteus oculatus - spotted gar X Lepisosteus osseus* - longnose gar L Lepisosteus platostomus - shortnose gar X AMIIDAE Ami a ca1 va - bowfin L ANGUILLIDAE Anquilla rostrata* - American eel TU CLUPEIDAE Alosa chrysochloris* - skipjack herring L Dorosoma cepedianum - gizzard shad X Dorosoma petenense - threadfin shad X HIODONTIDAE Hiodon alosoides - goldeye L ESOCIDAE Esox americanus - grass pickerel L CYPRINIDAE Cyprinus carpi o - common carp X Hyboqnathus hayi cypress minnow D Hyboqnathus nucha lis - Mississippi silvery minnow X Hybopsis aestivalis - speckled chub X Nocomis leptocephalus - bluehead chub X Notemiqonus crysoleucas - golden shiner X Notropis atherinoides - emerald shiner X Notropis blennius - river shiner X Notropis buchanani - ghost shiner L Notropis camurus - bluntface shiner X Notropis chrysocephalus - striped shiner X Notropis emilae - pugnose minnow TU Notropis fumeus - ribbon shiner TU Notropis lonqirostris - longnose shiner X Notropis lutrensis - red shiner X Notropis shumardi* - silverband shiner L Notropis texanus - weed shiner TU Notropis umbratilis - redfin shiner X Notropis venustus - blacktail shiner X Notropis volucellus - mimic shiner X Pimephales notatus - bluntnose minnow X Pimephales vigilax - bullhead minnow X Semoti lus atromaculatus - creek chub X CATOSTOMIDAE Carpi odes carpio - river carpsucker X Cycleptus elongatus - blue sucker (larvae only) L Erimyzon oblongus - creek chubsucker X Erimyzon sucetta - lake chubsucker (larvae only) L Hypentel ium nigricans* - northern hogsucker L Ictiobus bubalus - smallmouth buffalo X Ictiobus cyprinellus - bigmouth buffalo L Minytrema melanops - spotted sucker (larvae only) L No. 2 Bavou Sara Fishes 91 TABLE 3. CONT. SPECIES SOURCE ICTALURIDAE Ictalurus furcatus blue catfish Ictalurus melas - black bullhead Ictalurus natalis - yellow bullhead Ictal urus puntatus - channel catfish Pylodictus olivaris - flathead catfish APHREDODERIDAE Aphredoderus sayanus* - pirate perch CYPRINODONTIDAE Fundulus oli vaceus - blackspotted topminnow POECILIIDAE Gambusia affinis - mosquitofish Poecilia lati pinna - sailfin molly ATHERINIDAE Labidesthes sicculus - brook silverside Menidia beryl lina* - inland silverside PERCICHTHYIDAE Mo rone chrysops - white bass Mo rone mississippiensis - yellow bass CENTRARCHIDAE Centra rchus macropterus - flier Elassoma zonatum - banded pygmy sunfish Lepomis cyanellus - green sunfish Lepomis qulosus - warmouth Lepomis humi lis - orangespotted sunfish Lepomis macrochirus - bluegill Lepomis marginatus - dollar sunfish Lepomis meqalotis - longear sunfish Lepomis microlophus - redear sunfish Lepomis punctatus - spotted sunfish Lepomis symmetricus - bantam sunfish Micropterus punctulatus - spotted bass Micropterus salmoide^ - largemouth bass white crappie black crappie Pomoxis annularis Pomoxis niqromaculatus PERCIDAE Etheostoma asprigene Etheostoma caeruleum mud dater rainbow darter Etheostoma chlorosomum* - bluntnose darter Etheostoma gracile - slough darter Etheostoma parvipinne - goldstripe darter Etheostoma proeliare* - cypress derter Etheostoma whi pplei - redfin darter Percina ouachitae - saddleback darter Stizostedion canadense - sauger (larvae only) SCIAENIDAE Aplodinotus grunniens - freshwater drum L X X TU TU X L X L L L X X X X drainages (Guillory, 1981). Unfortunately, there is little published on the ichthyofauna of the streams in this region, except for Douglas (1974), Guillory (1981, 1982) and several Master's and Doctoral theses (Saul, 1974; Sobczak, 1976; Laiche, 1980). The number of species known from the two southward flowing lower Mississippi River tributaries, Thompson Creek and Bayou Sara, is similar with 89 and 80, respectively 92 Tulane Studies in Zoology and Botany Vol. 24 TABLE 4. Standardized Discriminant Function Coefficients FUNC 1 FUNC 2 FUNC 3 FUNC 4 FUNC 5 PH -0.07780 0.39875 -0.72813 -0.28597 0.97805 D.O. -0.15755 -0.32399 0.25823 -0.60156 -1.09422 ALKALINITY -0.06090 -0.16229 0.36258 -0.62893 -0.05940 CONDUCTIVITY 0.25892 0.02589 0.13170 -0.19767 -0.22383 TOTAL SOLIDS -0.43664 2.09234 0.28884 1 . 58946 3.10853 SUSP. SOLIDS 0.20762 -0.88644 -0.22385 -0.37946 -1.70610 TURBIDITY 0.17703 -1.60905 -0.02340 -1.76112 -1.07093 FREE CO2 0.00197 0.00790 -0.79007 0.14661 -0.08678 DEPTH (M) 0.06194 1.25672 1.09893 -0.73320 0.44940 WIDTH (M) 0.19755 1.39467 -0.35713 -0.84978 1.01854 CURRENT -0.01765 0.13450 0.35781 -0.67831 0.29226 TEMPERATURE 0.06749 -1.23164 0.13863 -0.43415 -0.18932 CROSS SEC. AREA 0.06456 -1.96236 -0.46527 0.69922 -1.21293 POSITION -6.56132 51.30540 -38.94077 20.61264 15.37281 ALTITUDE 6.77611 -59.50041 43.58128 -24.48588 -15.89091 GRADIENT -0.12318 -2.78645 2.82273 1.00519 -3.95386 STREAM ORDER 0.78970 -11.90598 8.33457 -2.24487 -4.94176 Percent of Total Variance Accounted For 30.07 12.21 11.60 8.63 7.33 (Guillory, 1981). Lists for both systems include at least 20 invaders from the Mis- sissippi River and another 4 or 5 species lo- calized in swamps or lower tributaries. Eighty-five species have been documented from Buffalo Bayou, but collecting efforts were concentrated in the upper and middle sections of that system (Cashner et al., 1976; Rogers et al., 1976). We assume that the species total for Buffalo Bayou would be appreciably greater than either Thomp- son Creek or Bayou Sara if an intensive survey of the lower reaches was conducted. Bayou Sara is perhaps more noteworthy for the species that do not occur within the system than for those that are recorded. Poor representation of three families, Petromyzontidae, Ictaluridae, and Cy- prinodontidae, is the most significant dif- ference between the fauna of Bayou Sara and that of the two adjacent drainages. No lampreys have been taken in Bayou Sara; two species, Ichthyomyzon gagei and Lampetra aepyptera, were reported from Buffalo Bayou and one, Ichthyomyzon gagei, from Thompson Creek. A combined total of ten ictalurids, Ictalurus furcatus, I. melas, I. natalis, I. nebulosus, I. punctatus, Noturus leptacanthus, N. miurus, N. noc- turnus, N. phaeus, and Pylodicitus olivaris, has been recorded in Thompson Creek and Buffalo Bayou, with 7 species found in each system. Although 5 cat fishes, Ictalurus furcatus, I. melas, I. natalis, I. punctatus, and Pylodictus olivaris have been collected in Bayou Sara, madtoms (Noturus) were notably absent. Three cyprinodontids, Fundulus catenatus, F. notatus, and F. olivaceus, ha.\e been taken in Buffalo Bayou and three, Fundulus chrysotus, F. notti, and F. olivaceus have also been reported from Thompson Creek. Only one of these, Fundulus olivaceus, was collected in Bayou Sara. In regard to the remaining families that the three drainages share, Bayou Sara is slightly less diverse than either of its two No. 2 Bayou Sara Fishes 93 adjacent drainages with a composition most similar to that of Thompson Creek. There are several plausible explanations for the lower species diversity in Bayou Sara. Bayou Sara drains an area approxi- mately 495 km\ whereas Buffalo Bayou and Thompson Creek each drain more than 800 km^ This represents a significantly smaller surface area and, consequently, a smaller volume of water in Bayou Sara. Al- though there is not a direct correlation between habitat diversity and water volume, an increase in available habitat in terms of both extent and diversity would be expected with larger volumes of water (Hanson and Campbell, 1963; Sheldon, 1968). Perhaps the two larger systems support more diverse faunas because of increased habitat availability. Thus Bayou Sara may lack suitable habitat or provide only marginal habitat for those species present in Buffalo Bayou and Thompson Creek that it does not share. Additionally, species extinction would be more likely in the smaller Bayou Sara system due in part to the reduction of habitat size associated with smaller volumes of water and correspond- ingly smaller populations of certain fish species (MacArthur and Wilson, 1967). Lack of suitable habitat may explain the absence of such species as Moxostoma poecilurum, Noturus phaeus and N. nocturnus, which have been reported from Buffalo Bayou or Thompson Creek (Guil- lory, 1981). These species are typically associated with firm substrates in contrast to the shifting sand bottoms characteristic of upper Bayou Sara. Taylor (1969) re- ported Noturus phaeus to be commonly encountered under vegetation in fast flowing streams and that N. nocturnus avoids streams with shifting sand bottoms. Paucity of aquatic vegetation, predomi- nance of loose sand substrates, and the absence of moderately deep, swift riffles with firm gravel substrates in Bayou Sara strongly suggest that a number of species are excluded from the system due to lack of suitable habitat. Habitat requirements alone cannot account for the exclusion from Bayou Sara of several species including Notorus miurus and N. leptacanthus which appear in adjacent drainages. Taylor (1969) noted that N. miurus often occurs in lowland or base level streams with soft substrates and N. leptacanthus is common in riffles over sand or gravel substrates. Habitats appro- priate for the above species thus appear to be available in Bayou Sara. Therefore, their absence in the system cannot be explained solely on the basis of lack of suitable habitat. A more complex explanation of Bayou Sara's low species diversity involves environmental changes associated with the geological history of the lower Misissippi River and its tributaries. Primary considera- tion was given to the effects of Pleistocene glaciation on conditions in the lower Mis- sissippi by Guillory (1978) in a discussion of modes of dispersal of upland species. TABLE 5. Standardized Discriminant Function Coefficients from Stepwise Analysis. TEMPERATURE STREAM POSITION Percent of Total Variance Accounted For FUNC 1 -0.15230 0.93651 78.65 FUNC 2 1.05648 0.51216 21.35 94 Tulane Studies in Zoology and Botany Vol. 24 During Pleistocene interglacial periods the lower Mississippi River and associated trib- utaries experienced an increase in stream load, volume and deposition, followed by periodic inundations by the Gulf of Mexico. Subsequent rejuvenation of these tributary systems occurred in response to sea level lowering during glacial periods (Fisk, 1944). At these times the streams were character- ized by lowered baselevels, increased gra- dients, clearer water and coarser sediments. Guillory (1978) suggested that glacial periods would have presented more favor- able conditions for the survival and dis- persal of freshwater fishes in the lower Mississippi Valley. Because periods of favorable and unfavorable conditions would have existed in all the major trib- utaries to the lower southeastern Mississippi River, these events would not account for existing differences in interdrainage species composition. Recent environmental changes compa- rable to those mentioned above have been reported by Fisk et al. (1938). Although inundations of the lower Mississippi by the Gulf have not occurred, the meandering of the Mississippi River has produced condi- tions similar to those during the Pleistocene. Westward movements of the Mississippi bring about a local lengthening of its eastern tributaries, causing an overall decrease in average gradient. Eastward migrations of the river would conversely shorten these tributaries and effectively lower the baselevel. Consequently, tribu- taries to the eastern embayment experience gradient increases and trench their former floodplains to reach the lower baselevel (Fisk et al., 1938; Guillory, 1976). Environmental changes associated with decrease in gradient, reduced flow, decreased oxygen tensions, increased tem- perature and increased siltation may have resulted in marginal, intermittent habitat for many existing species. Local extinction of species would occur if favorable habitat was not available for survival until reverse migration of the Mississippi River occurred. Guillory (1976) explained the disappearance of the blackbanded darter (Percina nigro- fasciata) from the Thompson Creek system as a result of such recent changes. The fact that species were eliminated from Bayou Sara and not Buffalo Bayou and Thompson Creek may result from differential retention of favorable habitat during variations in the course of the Mississippi River. Bayou Sara, Buffalo Bayou and Thomp- son Creek have relatively steep gradients (Fisk et al., 1938). Steep gradients in Thompson Creek and Bayou Sara result from the localized uplift along the Mobile- Tunica flexure in response to sediment overloading in the Mississippi River delta (Howe, 1936; Fisk et al., 1938). This flexure and the steepest gradient occur along a line through the headwaters of Bayou Sara. A large portion of the upstream sections of Thompson Creek and all of Buffalo Bayou lie above this line. Conceivably, some species survived unfavorable conditions occurring downstream during westward migrations of the Mississippi by moving into higher gradient sections of the tribu- taries. Only a small portion of the Bayou Sara system shows the high gradient associated with the Mobile-Tunica flexure. Discharge in this area is consistently low and the extreme headwaters become inter- mittent during dry periods. Sufficient habitat may not have been available for extended periods to support a major influx of downstream species. Therefore, elimina- tion of species could have been more pro- nounced in Bayou Sara. Intrasystem Distribution. A pattern of pro- gressive downstream addition of species has been observed in many drainage systems (Thompson and Hunt, 1930; Burton and Odum, 1945; Kuehne, 1962; Hanson and Campbell, 1963; Larimore and Smith, 1963; Harrel et al., 1967; Jenkins and No. 2 Bavou Sara Fishes 95 Freeman, 1972; Whiteside and McNatt, 1972; Lotrich, 1973; Cashner and Brown, 1977; Evans and Noble, 1979; Guillory, 1981). The observed longitudinal distribu- tion of fishes in Bayou Sara (Table 6) does not strictly follow previously described patterns. There is a progressive downstream addition of species from 1-4, reaching a high of 33 at Station 4. However, species number declines further downstream with only 28 species at Station 5 and 21 species at Station 6. The most abundant components of the headwater fish fauna were Nocomis lepto- cephalus, Notropis chrysocephalus, N. longirostris, and Etheostoma caeruleum. Downstream fauna was characterized by two very abundant cyprinids, Notropis venustus and Pimephales vigilax, the addition of several species, Hybognathus nuchalis, Notropis atherinoides and N. blennius, and the presence of Gambusia affinis. Among the 43 species collected during this study, only four, Nocomis leptoce- phalus, Semotilus atromaculatus, Ictalurus natalis and Lepomis cyanellus, were absent from lower reaches of the stream. Nineteen species were restricted to the middle and lower reaches, and 12 are continuously dis- tributed along the main channel or are absent from only a single locality (Table 6). All 21 species recorded at Station 1 were taken from at least one other locality. Twenty-two species were added from Sta- tions 2-6, four at Station 3, nine at Station 4, three at Station 5 and two at Station 6. Ten species are represented by single speci- mens. Variation in discharge and the associated physical and chemical changes may explain, in part, the pattern of longitudinal distri- bution of fishes in Bayou Sara. Extremely low flow occurred between June and De- cember 1978. Permanent flow did persist at the six stations during this period, but many smaller tributary creeks were intermittent or dry. Hynes (1972) reported that species sur- vive drought conditions as isolated popula- tions in favored downstream locations, and Larimore et al. (1959) observed that the re- colonization of upstream habitats occurred rapidly following restoration of flow. Downstream displacement of species in response to low water or drought conditions would obscure any tendency for zonation and distort the pattern of species addition. Similar displacement would be expected in small creeks and tributaries to the lower reaches of Bayou Sara. Such downstream shifts may account for the infrequent occur- rence of Erimyzon oblongus, Etheostoma caeruleum, E. whipplei, and Percina ouachitae at Stations 5 and 6 and Nocomis leptocephalus, Semotilus atromaculatus and Ictalurus natalis at Station 4. C. Data Analysis Multiple discriminant functions analysis was employed to assess the importance of 17 environmental variables in regard to spatial separation of the groups (species) in Bayou Sara. Seventeen discriminant func- tions (DP) were extracted during the direct analysis. Standardized DP coefficients for each of the 17 variables in these functions (Table 4) indicate the dominance of two factors, altitude and stream position. The high positive coefficient for gradient in DP I, in conjunction with the prominant negative one for stream position, indicates a marked restriction of species to the head- waters and lower reaches of Bayou Sara. According to the analysis, the downstream variation in altitude and stream position accounts for the longitudinal variation in species distribution in the Bayou Sara system. Because, however, the 17 DP extracted during the direct analysis failed to reach a high degree of statistical significance (p>.10), sufficient variation between species apparently does not exist in these two variables and the other 15 parameters to account for the observed species separa- tions. 96 Tulane Studies in Zoology and Botany Vol. 24 =3 a; o oo >1 •1 — (O 2 CO s- OJ c x: •1- -i-> I I I I I I I I I I I I I 1 c\j 1X1 c\j a~i O) 1/1 u ra ZJ 3 • f— C r— to > J2l 1 ^ 2: <-3 No. 2 Bavou Sara Fishes 97 ro <^ 1 — r^ isD r-^ r^ ro oo c\j CO I — ro 1 — 0-) un CO IT) I — n ro in oo :3 O s- 3 CL E i- ro rO O O o (— §; 98 Tiilane Studies in Zoology and Botany Vol. 24 Similar results were obtained with the stepwise DF analysis, with stream position and temperature selected for inclusion into the analysis. One statistically significant (p<0.01) DF, accounting for 78.707o of the among - species variance, was extracted during the stepwise analysis. As was the case in the direct analysis, stream position provided the most significant contribution to the formulation of this function. The high positive standardized coefficient for this variable (Table 5) indicates once again a distinct separation of species at upstream localities. The negative coefficient for temperature signifies species separation in the cooler waters of the upstream stations. However, the magnitude of the coefficient for temperature indicates a minor contribu- tion to this function. Discriminant functions scores for each of the 33 species were also computed during the analysis. These scores were based on physical and chemical measurements asso- ciated with the time of capture of each species. A discriminant score is thus obtained for a species for each instance it was collected. Mean discriminant scores were tabulated to facilitate evaluation of species distribution in relation to important environmental variables. Large discrim- inant scores indicate the presence of a species where significant environmental variables of that function are also cor- respondingly large or small depending on the signs of the DF coefficients (Nie et al., 1975). Figure 3 is a plot of the mean discrim- inant scores for 33 species on DF I of the stepwise analysis. The underlined numbers correspond to one of the 43 species collected during this study. A key to the species codes appears in Table 6. The distribution of mean discriminant scores (Fig. 3) appears to corroborate the observed pattern of longi- tudinal distribution of species in Bayou Sara (Table 6). Three species, Semotilus atromaculatus (22), Erimyzon oblongus (23), and Etheostoma caeruleum (40), were restricted to upstream areas where water temperatures were the lowest. Although not restricted to the most upstream stations, two species, Nocomis leptocephalus (9) and let alums natal is (26), appear to have been limited to more upstream localities with cooler water temperatures. A large number of species were either restricted to the mid- stream sections of Bayou Sara or were continuously distributed along the water- course (-0.5x amevioanus, 90 Etheo stoma asprigene, 91 oaeruleum, 91, 95-96, 98 ahlorosomum, 91 gvaoile, 91 parvipinne, 91 proliare, 91 whipplei, 91, 95-96 Eupatorium aapilli folium , 129 ooelestinvm , 129 rugosum, 129 sevotinim, 129 Euphorbia oordi folia, 123 hirta, 123 nutans, 123 prostrate , 123 supina, 123 Euphorbiaceae, 112, 123 Fagaceae, 119 Fimbristylis oastanea. 111 Fraxinus spp., 99, 102 oaroliniana, 125 pensylvaniaa , 125 profunda, 106, 108, 125 Fundulus catenatus , 92 ohrysotus , 92 notatus , 92 notti, 92 olivaaeus, 91-92, 96 Galium aparine, 128 tinotorium, 128 Gambusia af finis , 91, 95-96 Gentianaceae, 126 Geraniaceae, 122 Geranium oarolinianum , 112 Geum canadense. 111 Gleditsia trioanthos, 104-105, 122 Graminieae, 112, 114 Guzmania , 2 balanophora , 46 hygrometrioa , 69 ororiense, 2 ororiensis, 41 splitg erberi , 78 Gymnospermae, 112, 114 Habenaria repens , 112, 118 Hamamelidaceae, 112, 114 Heliotropium ourassaviaum , 126 Hibiscus lasiooarpus, 124 Hiodon alosoides, 90 Hordeum pusillum, 115 Hybognathus hayi, 90 nuchalis, 90, 95, 97 Hybopsis aestivalis, 90, 97 storeriana, 97 Hydrocharitaceae, 114 Hydroaotyle ranunauloides , 125 imbellata, 125 vertioillata , 125 Hydrophyllaceae, 126 Hygrophioa laaustria, 127 Hymenooallis occidentalis , 118 Hyper caceae, 124 Hypentelium nigricans, 90 Hypericum walteri, 124 Ichthyomyzon gagei, 92 Ictalurus furcatus, 91-92 melas, 91-92 natalis, 91-92, 95-96, 98 nebulosus, 92 punctatus, 91-92, 97 Ictiobus bubalus , 90, 97 cyprinellus , 90 iJea; deoidua, 104, 198, 123 vomitoria, 123 Ipomoea x teuaantha, 126 sagt-ttata, 126 trichooarpa, 126 Iridaceae, 118 Iris fulva, 105, 111, 118 gigantioaerulea , 105, 118 Iva annua, 109, 129 frutesoens, 112 Juglandaceae, 119 Juncaceae, 118 Junaus effusus , 118 tenuis, 118 Justioia lanoeolata, 127 Koetreutevia paniculata, 123 Kosteletzkya virginioa, 124 Labiatae, 126 Labidesthes siaaulus , 91, 97 Lactuaa canadensis, 129 floridana, 129 Lampetra aepiptera, 92 Lauraceae, 121 Leevsia tenticutaris , 115 virginiaa, 115 Leguminosae, 112, 121 Lemna spp. , 108 minor, 117 Lemnaceae, 117 Lentibulariaceae, 127 Lepidium virginicum, 121 Lepisosteus oaulatus, 90, 97 ossews, 90, 97 platostomus, 90 Lepomis cyanellus , 91, 95-96 gulosus, 91 humilis, 91 macroohirus, 91, 97 marginatus, 91 megalotis, 91, 97 miarolophus, 91 Lepomis punotatus , 91 symmetrious , 91 Leptoohloa fasaioularis , 112 Ligustrum japonicum, 125 lucidum, 125 Lilaeopsis aarolinensis , 125 Liliaceae, 118 Limnohium spongia, 114 Liquidamhar styraciflua, 85, 103, 105-106 Lithospermum tuberosum, 126 Lobelia cardinalis , 128 Lobeliaceae, 128 Loganaceae, 126 Lonioera japonica, 128 Ludwigia deourvens, 124 leptooarpa, 124 oatovalvis, 124 Uruguay ensis , 125 Lycopus rubellus, lib virginious , 126 Lygodium japonicum, 113 Lythraceae, 124 Lythrum lane eolation , 124 linear e, 112 Malvaceae, 124 Malvastrum ooromandelianum , 124 Matelia gonocarpa, 126 Mazus japonicus, 127 Mecardonia acuminata, 127 prooumbens. 111 Melia azedarach, 123 Mellaceae, 124 Melilotus indicus. 111 Melotkria pendula, 128 Menidia beryllina, 91 Micropterus punctulatus, 91, 96 salmoides, 91 96 Mikania scandens, 104, 129 Mimosa strigillosa , 111 Mimulus alatus. 111 Minytrema melanops, 90 Modiola aaroliniana, 124 Monocotyledonae, 112 Moraceae, 119 Morone chrysops, 91, 97 mississippiensis , 91 Morus rugra, 119 Moxostoma poecilurum, 99 Myosotis macro spermwv, 126 Myrica cerifera, 105-106, 108, 119 Myricaceae, 119 Myriophyllum brasiliensis , 112 Najadaceae, 114 Najas guadalupensis , 108, 114 Navia lopezii, 5 Nemophila aphylla, 126 Nocomis leptocephalus , 90, 95-96, 98 Notemigonus crysoleucas, 90, 96 Nothoscordum bivalve, 118 Notropis atherinoides, 90, 95, 97 blennius, 90, 95 buchanani, 90 camurus, 90, 97 chrysocephalus, 90, 95-96 emilae, 90 fumeus, 90 longirostris, 90, 95-96 lutr ensis, 90, 97 shumardi, 90 texanus, 90 umbratilis, 90, 97 yenustus, 90, 95-96 voluoellus, 90, 96 ..j; ffo turus ,92 leptacanthus , 92-93 ..iw . miurus, 92-93 noetiir'nus, 92-93 phaeus, 92-93 Nymphaea odorata, 120 Nympheaceae , 121 Nyssa aquatica, 105, 125 Oenothera speoiosa, 125 Oleaceae, 125 Onagraceae, 124 Oplismenus hirtellus, 115 Orchidaceae, 122 Oxalidaceae, 122 Oxalis oorymhosa, 122 dillenii, 122 Palmae, 117 Panicum spp., 107 anoeps, 115 oorrmutatum , 107, 115 diahotomiflorum, 115 gymnooarpon , 115 hemitomim, 115 rigidulum, 115 virgatwn, 115 Fanterpe insignis , 15 Papaveraceae, 121 Parthenium hysterophorus , 129 Parthenocissus quinque folia , 104, 124 Paspalum distiahum, 115 urvillei, 115 vaginatwn, 112 Passi flora incarnata, 124 lutea, 124 Passif loraceae, 124 Persea borhoniaa, 109, 121 Persiaaria spp., 107 densi flora , 119 hydropiperoides , 120 punctata, 120 setacea, 120 Percina nigra fasoiata, 94 ouachitae, 90, 95, 96 Petunia parviflora, 127 Phalaris oaroliniana, 115 Phoradendron tomentosum, 119 Pkragmites australis, 109, 115 Phyllanthus urinaria, 123 Phyla nodi flora, 126 Physalis angulata, 127 Phytolaaa oleraoea, 120 Phytolaccaceae, 120 Pilea pumila, 119 Pimephales notatus, 90, 96 vigilax, 90, 95, 97 Pistia stratiotes, 117 Pinus eohinata, 85 taeda, 85 Plantago major, 128 Plantaginaceae, 124 Pluahea odorata, 129 Poa annua, 115 Poecilia latipinna, 91 Polygonaceae, 112, 119 Polygonum ramossissimum, 120 virginianum, 120 Polymnia uvedalia, 129 Polypodiaceae, 114 Polypodium polypodioidea, 114 Polypogon monspeliensis , 115 Pomoxis annularis, 91, 97 nigromaculatus , 91, 97 Pontederia cordata, 118 Pontederiaceae, 118 Populus deltoides, 119 Portulaoa oleraoea, 120 Portulacaceae, 120 Primulaceae, 125 Ptilirrmium oapillaceum, 125 Pueria lohata, 122 Pylodiatus olivaris, 91-92 Pyrrhopappus oarolinianus , 129 Queraus nigra, 85, 103, 105-106, 108, 119 nuttalli, 104, 108, 119 virginiana, 103, 105-106, 108, 119 Ranunculaceae, 120 Ranunculus platensis , 121 pusillus , 121 sceleratus, 121 trilobus, 121 Rhamnaceae, 123 P/ius toxicodendron var. vulgaris, 104, 123 Rhynchosia minima, 122 Rhynchospora corniculata, 117 Rivina humilis , 120 Rosaceae, 121 Rubiaceae, 128 Rubus trivialis, 109, 121 Ruellia nudi flora, 127 Rumex crispus , 120 pulcher, 120 verticillatus , 120 Rutaceae, 122 Sabal minor, 104, 117 Sabatia calycina, 126 Sacciolepis striata, 115 Sagittaria lanciofolia, 107, 114 platyphylla, 114 Salicaceae, 119 Salix nigra, 109, 119 Salvia oocoinea, 126 Salvinaceae, 113 Sambucus canadensis, 104, 128 Samolus parviflorus, 125 Sanioula canadensis, 125 Sapindaceae, 123 Sapium sebiferum, 109, 123 Sapotaceae, 125 Saururaceae, 119 Saururus cernuus, 119 Scirpus calif ornicus , 107, 117 cyperinus , 117 lineatus , 111 olneyi , 112 validus, 112 Schizaeaceae, 113 Scrophulariaceae, 112, 127 Semotilus atromaculatus , 90, 95-96, 98 Senecio globellus, 129 Sesbania macrocarpa, 122 punicea, 122 Setaria geniculata, 115 glauaa, 115 magna, 115 Sida rhomhi folia , 124 Sisyvinohiwri rosulatum, 118 Smilax bona-nox, 118 hispida, 118 rotundi folia, 118 Solanaceae, 127 Solanum ameviaanwri, 127 aavolinense, 127 Solidago altissima, 109, 129 senrpervirens , 107, 129 Spartina oynosuroides, 116 patens, 107, 116 Sphenopholis ohtusata, 116 Spilanthes americana, 129 Spiranthes cernua var. odorata, 112, 118 Spirodela polyrhiza, 117 Sporobolus indicus , 116 Stachys cvenata, 127 tenuifolia, 127 Stellavia media, 120 Stenotaphnm seaundatum, 116 Stizostedion oanadense, 91 Stvophv styles helvola, 122 Taxodiaceae, 114 Taxodiwn distiahwn, 105, 108, 114 Thalictrvuv dasyaarpum var. hypoglauaum , 121 Theaophyllum , 2, 12, 14, 18 aaiminata, 64 acuminatum, 64 balanophorwri , 46 balanophorim var. siXbpiatvm, 46-48 bracteosum, 75 capt ta tw/^ , 7 7 aomatvm, 66 arassiflorum, 41 aylindraoevm, 42 discolor, 78 hygrometriwn , 69 irazuense , 12, 14, 41 latissima, 78 latissimim, 33, 78 laxvm , 2 1 lineatyjn, 46 kupperi, 42 montanim, 37, 40 oror-iense, 41 panniaulatum , 28 pedicillatwn , Ik-lb piatim, 58 pittieri, 73 poortmani, 2 rubrvm, 22, 44 singuli forum, 68 speatabile, 78 splitgerberi , 78 standleyi, 41 stenophyllum , 61 turbinatum, 24 violascens, 24-25 viride, 50 werokleanum, 14, 36, 39-40 wittmaokii, 2 Thelypleris kunthii, 114 Tilia, 85 Tillandsia, 2 usneoides, 105, 116 Tradesaantia ohiensis, 118 Trifolium repens, 122 resupinatum, 122 Trisetum pensylvaniaum, 116 Typha spp. , 107 domingensis, 114 latifolia, 114 Ulmaceae, 119 f/Zmws americana, 104-106, 108, 119 rubra, 106, 119 Umbelliferae, 112 Urticaceae, 119 Utricularia bi flora, 112 gibba, 127 yer'fcena brasiliensis , 126 rigida, 126 urtici folia, 126 Verbenaceae, 126 Ferfcesina virginica, 129 7er?i<9nt(2: gigantea, 129 7ercnica peregrina, 127 persica, 127 Vicia ludoviciana, 122 Fig'TM: luteola, 122 Viola papilionacea , 124 septemloba, 124 Violaceae, 124 Viscaceae, 119 Vitaceae, 124 Vitis cinerea, 124 rotundifolia , 104, 124 l/rtesea, 2, 10, 14-15, 18 attenuata, 4, 17, 21, 62, 64-67, 73 balanophora , 4, 13, 17, 22, 44, 46-49, 51 balanophora group, 16-17 braateosa, 4. 17, 20, 67, 75-78 bracteosa group, 17 capitata, 4-7, 17, 20, 67, 76-78 comata, 17, 19, 65-68 comata group, 16-17 crassiflora, 42, 44 cylindraceae , 42, 44 diffusa, 4, 6, 9, 17-19, 27-29 discolor, 78 gigantea , 15 greenbergi, 4, 10, 17, 19, 30-34 hainesiorum, 4, 12-13, 17, 21, 52-54, 57-58 hygrometrica , 3-4, 12, 17, 20, 69-71, 73 hygrometrica var. angusti folia, 66-67 hygrometrica group, 16-17 irazuensis , 39, 42-44 Zanniohellia palustris, 113 kathyae , 4. 17, 19, 65, 70-73 Zizaniopsis miliaoea, 107, 116 kupperi , 42, 44-45 latissima, 4, 14, 16-17, 19, 23, 32-35, 44 latissima group, 17 Z-cuca , 2 7 leptopoda, 4-5, 17, 19, 22-23, 26, 28, 34, 44 leuoophylla, 4-5, 12-13, 17, 21, 48, 53-55, 57-58 leucophylla group, 16-17 lineata , 46-47 luis-gomezii , 4, 17, 22, 48-51 lyman-smithi , 4, 17, 22, 35-37 montana , 38, 40 nephrolepis, 3-4, 11, 14, 17, 22, 25, 35-41, 44-45, 51 notata, 4, 17, 20, 70-71, 73-75 ororiensis, 3-5, 10, 12, 14-15, 17, 21-22, 25, 39, 46, 51 ororiensis group, 10-17 panicutata , 28 pedioellata, 304, 14, 17-19, 23-29 pedicellata group, 16-17 piota, 4-5, 17, 21, 58-62 pittievi ,73 rainifeva , 33,35 rubra , 22 saroolepis , 62-64 singuli flora, 4, 10, 17, 19, 68-69 splitgerheri , 78 standleyi , 42, 44 stenophylla, 4, 14, 17, 21, 44, 60-62, 65-66 stenophylla group, 17 tri flora, 4, 6-7, 9, 17, 19, 28-30 triflora group, 16-17 turbinata , 24, 26 umbrosa, 4, 14, 17, 19, 30, 32-34 umbrosa group, 16-17 unilateris , 15 lucoris , 4, 12-13, 16-17, 20, 57-58 uxoris group, 16-17 vietoris , 4, 17, 21, 54-58 violascens, 24-25 :;iHdis, 4, 13, 17, 22, 50-52, 60 vittata, 79 werkleana, 36 -, o-, ^n ijtlZiamst, 3-4, 12, 14, 17, 21, 60, 62-64 Wolffia oolvmbia-m. 111 ¥olfiella, 113 Xanthium strumariiov, 129 Xiphion, 2, 15, 17 Youngia japoniaa, 129 2 0 8 ■/ 055 3 2044 ''i3"'361 087