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Manual de Plantas de Costa Rica

Main | Family List (MO) | Family List (INBio) | Cutting Edge
Draft Treatments | Guidelines | Checklist | Citing | Editors

The Cutting Edge

Volume II, Number 1, January 1995

News and Notes | Recent Treatments | Leaps and Bounds | Germane Literature

MANUAL DEADLINES. Our original plan called for publishing the Manual all at once. We have recently decided to change that plan and prioritize the monocot volume, publication of which is now slated for 1997. MONOCOT CONTRIBUTORS TAKE NOTE: the Manual deadline for all monocot treatments is now 1 January, 1996, slightly less than one year from today. Most of you will already have heard from us individually about this matter. The new deadline is later, not earlier, than previously indicated deadlines; the difference is that this time it is an absolute deadline, not some nebulous target date. We will have no choice but to reassign any family treatments not submitted by the first of next year. DICOT CONTRIBUTORS: Publication of the Manual dicot volume is now scheduled for 1999. Our previously stated deadline of 1 January, 1996 remains in effect for dicots, with an absolute deadline of 1 January, 1997. Although reassignment of dicot treatments will not be considered until the latter date, we would prefer to have them delivered as expeditiously as possible. ALL CONTRIBUTORS: Should you decide that, for whatever reason, you will be unable to meet your absolute deadline, please let us know immediately; we prefer to keep the inevitable last-minute surprises to a minimum.

WHAT ABOUT NEW TAXA? Some contributors have indicated that their family treatments have been delayed because of a need to prepare formal descriptions of new taxa for prior publication. This, in our view, is putting the cart before the horse. Although we want the Manual to be as comprehensive as possible, it is not intended to be revisionary. If time is a consideration and you worry about meeting your deadline, we would urge you toprioritize your treatment over the publication of new taxon names. Nonetheless, we DO want the new taxa to be treated in full in the Manual whenever possible. There are two options for doing this: 1. Provide informal names ("Species A," "Species B," etc.) for undescribed taxa; Latin names can be substituted up until the last moment if they are published in time, or else cross-reference to the informal Manual names can be effected upon publication of the Latin names. 2. Publish new taxon names in the Manual; this option was explicitly proscribed in our original guidelines, but we have had a change of heart and now see no problem with this. Those who opt for publishing new names in the Manual should understand that space will be limited to the minimal requirements for valid publication (type citation, brief Latin diagnosis, etc.). In rare cases, an illustration might be feasible.

TO AND FRO. INBio biologist and Manual stalwart José González arrived at MO on 23 January for a visit of two months, during which time he will concentrate on polishing up his Moraceae treatment, particularly as regards Ficus. Barry Hammel and family arrived from Costa Rica in late December, and will remain through March. José and Barry will travel together to F and US during their stay. Manual contributor Jim Luteyn (NY; Ericaceae pro parte) worked in the herbarium at MO during the last week of October, and OTS's Chuck Schnell paid a return visit during the first week of November to continue his revisionary work on Conostegia (Melastomataceae). Manual co-PI Nelson Zamora returned to Costa Rica in mid-December following nearly a year at MO.

NEW STAFF AT INB. Fern specialist Alexander Rojas, of the Universidad Nacional Autónoma de Heredia, has been hired as a half-time curator at INB on a three-month, provisional basis. Bryologist Gregorio Dauphin, of the Universidad de Costa Rica, will be working full-time for four months at the Estaciones de Altamira and Santa María de Pittier, Cordillera de Talamanca.

NEW DIGS FOR CR. The Herbario Nacional de Costa Rica, shuttled around like an orphaned child for the past four years, now has a spanking new home. The new facility, built in the "hueco" at the northwest corner of the Museo compound, was inaugurated on 5 December, 1994, during a ceremony featuring President José María Figueres and Museo director Melania Ortiz Volio. Also honored was Luis Diego Gómez, a former director of the Museo, who currently administers OTS's Wilson Botanical Garden; Luis Diego had spearheaded the construction of the former natural history building at the Museo, which was razed to accommodate the Plaza de la Democracia. The CR herbarium probably will not be moved into its new quarters before April of this year. Barry Hammel reports that the new facility looks nice and solid, and we all hope it will provide more and better working space.

CONGRATULATIONS to Manual co-PI Nelson Zamora, who was awarded the 1994 Rupert Barneby Award by the New York Botanical Garden. The award will fund travel for Nelson to NY to continue his studies on Costa Rican Fabaceae. He is considering an April departure.

FIRE ON HIGH. We were all shocked and saddened to learn of the complete destruction by fire of OTS's research complex at the Wilson Botanical Garden (Las Cruces). The laboratories, library, dormitories, and dining area were all consumed on 23 November, 1994, in a conflagration possibly caused by an electrical problem in a caretaker's apartment. The cost to rebuild has been estimated at ca. $500,000, only a fraction of which will be covered by insurance.

DATA-BASE DEVELOPMENTS. The Manual data-base has been synonymous with INBio's plant data-base since the inception of the last-mentioned institution. Programming and maintenance of the data-base (VARIOS) have always been shouldered by Manual personnel, particularly co-PI Barry Hammel. These functions have now been turned over entirely to INBio, more specifically to Allan Prendas, who has been hired on a full-time basis as data-base manager/programmer for the Department of Botany. This will relieve Barry of what has become an unacceptably burdensome responsibility. We will continue, as long as practicable, to transfer new accessions from the Manual/INBio data-base to the TROPICOS data-base at MO. Sometime during 1995 the Manual data-base information will all be transferred into the INBio institution-wide data-base system (BIMS), and data-entry will be directly to BIMS rather than through VARIOS.

By the time this bulletin is distributed, INBio data-base manager Werner Böhl will have a version of the entire Manual data-base accessible through the World Wide Web. In fact, it was already briefly available at the beginning of January, but then was taken off while department heads at INBio discussed the tangled question of whether or not to display complete locality information. A decision, which we find quite acceptable, was made to round-off coordinates and to leave off specific ("500 m N of the junction between...") locality information, but to include all political divisions and broad geographic regions. The address is:


Let us know if you find this more or less useful than the version we have on the MO gopher (which presently includes only the pteridophytes and gymnosperms). We suspect some users won't yet have access to WWW, so we will continue to update the version on the gopher.

HARD COPY. The latest wood-wafer edition of our checklist of Costa Rican vascular plants from the Manual data-base was generated on 12 November, 1994. This is a vouchered list based on actual collections and, as such, does not pretend to be comprehensive; those species not yet accounted for are for the most part rare and not represented by specimens in Costa Rican herbaria. The printout runs to 222 pages (not including the index), single-spaced, with one line per species entry. Represented among the ca. 60,000 records now comprising the data-base are 249 families of indigenous or naturalized vascular plants (plus 12 families only in cultivation), with 1877 genera (plus 170 in cultivation) and 8311 species (plus 407 in cultivation). The checklist can also be conveniently accessed via the Internet gopher.


MORACEAE. José González (INB).
Eighteen genera are treated for Costa Rica, of which one (Artocarpus) is known solely from cultivation. The total number of species is 93 (eight of which are cultivated); Ficus is by far the largest genus, with 53 species (including six cultivated). Olmedia is tentatively maintained as distinct from Trophis. Additions to Burger's (1977) Flora costaricensis treatment include: Clarisia mexicana (Liebm.) Lanj. and C. racemosa Ruiz & Pav.; Ficus garcia-barrigae Dugand, F. glaucescens (Liebm.) Miq., F. petenensis Lundell, and F. zarzalensis Standl.; Morus celtidifolia Kunth; Naucleopsis ulei (Warb.) Ducke and two undescribed species of Naucleopsis; Perebea guianensis Aubl.; and two undetermined species of Pseudolmedia.

ORCHIDACEAE (pro parte). John T. Atwood (SEL).
This contribution, recently submitted to Manual Orchidaceae coordinator Bob Dressler, comprises treatments of the genera Trigonidium and Maxillaria. These are "lite" out-takes (the way we like 'em) from a more detailed work that will be published separately. Just three species of Trigonidium occur in Costa Rican, compared with 101 of Maxillaria. About 19 species of Maxillaria are ostensibly endemic to Costa Rica, but many others are shared only with western Panama.

Only the genus Valeriana is represented in Costa Rica. Nine species are treated, although V. sorbifolia Kunth (known from Nicaragua and South America) is hypothetical. No species are endemic to Costa Rica.


APOCYNACEAE. Francisco Morales, who recently completed the Manual Apocynaceae treatment, reports the addition of Forsteronia peninsularis Woodson, based on a collection from Buenos Aires de Osa by Gerardo Herrera.

BROMELIACEAE. Pitcairnia lyman-smithiana H. Luther, recently described from Panama, can now be reported from Costa Rica on the basis of collections made by Francisco Morales and Eduardo Lépiz from Tapantí. Francisco also collected Tillandsia flexuosa Sw. on the Osa Peninsula; this species was not previously known from north of Panama.

BURSERACEAE. Haber 11322 from 1200 m elevation on the Pacific slope of Monteverde has been identified by specialist Doug Daly (NY) as Bursera grandifolia (Schltdl.) Engl. This is the first Costa Rican record of this basically Mexican species.

CONVOLVULACEAE. Hammel 19575 from the Santa Elena Peninsula, Guanacaste Province, has tentatively been identified as Ipomoea ternifolia Cav. The specimen is sterile, but this is a vegetatively distinctive species, disjunct (as far as we can determine) from Mexico.

CYCLANTHACEAE. Chorigyne cylindrica R. Erikss. can now be reported from Costa Rica on the basis of Hammel & Perry 19720, collected at the teleférico site at ca. 500 m elevation in Parque Nacional Braulio Carrillo. This was previously considered a Panamanian endemic.

FABACEAE/MIMOSOIDEAE. Nelson Zamora reports that a collection at DUKE made by Robert L. Wilbur from La Cruz de Guanacaste is the first Costa Rican record of Pithecellobium furcatum Benth., a northern species. This species (a Pithecellobium sensu stricto) was recently recollected from Estación Pitilla, Volcán Orosí, by parataxonomist Elba López.

FABACEAE/PAPILIONOIDEAE. Rivera 1173, from 500 m elevation in Parque Nacional Rincón de la Vieja, is the first Costa Rican record of Lonchocarpus velutinus Benth. ex Seem., known previously from Panama and Colombia.

HYDRANGEACEAE. Francisco Morales, who has completed the Manual treatment of this family, reports that Hydrangea steyermarkii Standl. can be added on the basis of a collection from the Cordillera de Tilarán. This species had not previously been found south of Nicaragua.

LOGANIACEAE. MO specialist Alan Brant has identified a collection (A. Fernández 121) from 200 m elevation in Parque Nacional Braulio Carrillo as Strychnos brachistantha Standl. This is a widespread species known from both Nicaragua and Panama, but we can find no previous Costa Rican record.

ONAGRACEAE. Ludwigia helminthorrhiza (Mart.) H. Hara, a peculiar floating aquatic with inflated nodal roots, was collected by Barry Hammel (19652) in the vicinity of Siquirres. About a week thereafter, a specimen (A. Rodríguez 370) of the same species were brought to INBio for identification by two students working in Parque Nacional Tortuguero. We can find no other record of this species between southeastern Honduras and the Canal Zone. We have no idea whether this is a weird coincidence, or if this species has recently become established.

PTERIDOPHYTA/PTERIDACEAE. Haber & Zuchowski 10113, from 900 m elevation on the Pacific slope near Monteverde, has been identified by INBio pteridologist Alexander Rojas as Adiantopsis chlorophylla (Sw.) Fée. This is the first Costa Rican record of this widespread species.

RUBIACEAE. Hammel 19211 from Cerro Anguciana in the southwestern Pacific region has been identified by Manual contributor Charlotte Taylor (MO) as Joosia umbellifera H. Karst. (incl. J. panamensis Dwyer), not previously reported from north of Panama.


Atwood, J. T. 1994. Two new species of Maxillaria (Orchidaceae) from southern Central America and a new name for a well-known taxon. Lindleyana 9: 227-231.

_____. 1994. Two new Costa Rican species of Maxillaria (Orchidaceae) related to Maxillaria flava and Maxillaria microphyton. Lindleyana 9: 232-237.

_____. 1994. Two new species of Costa Rican Maxillaria (Orchidaceae), and a clarification of Maxillaria bracteata. Lindleyana 9: 239-242.

In the first of these three articles, Maxillaria tubercularis J. T. Atwood is described from the high Talamancas of Costa Rica and Panama, and the new name M. schlechteriana J. T. Atwood is provided for the taxon lately known as M. wrightii (Schltr.) Ames & Correll var. imbricata (Schltr.) Ames & Correll. The Costa Rican novelties described in the second article are: M. burgeri J. T. Atwood, from the Cordillera Central and near Santa María de Dota; and M. muscoides J. T. Atwood, from the Tapantí/Pejivalle vicinity of Cartago Province. In the final article, Maxillaria vagans Ames & C. Schweinf. is relegated to synonymy under the less familiar M. bracteata (Schltr.) Ames & Correll, based on a critical examination of a recently discovered isotype; M. amabilis J. T. Atwood is described from the San Ramón/Zarcero region of Alajuela Province, and M. monteverdensis J. T. Atwood & G. Barboza from the Monteverde Reserve.

All new taxa are illustrated in all three articles, and a key is provided (in the second article) to the Costa Rican species of the Maxillaria flava/M. microphyton complex.

Dressler, R. L. 1994. (1142) Proposal to conserve Scaphyglottis against Hexisea (Orchidaceae). Taxon 43: 665-666.

Hexisea, with 2-5 species, has priority over Scaphyglottis, with nearly 50 species. If conservation is rejected, the proposed merger of these genera will require ca. 40 new combinations.

Gómez-Laurito, J. & B. E. Hammel. 1994. New species in the Acanthaceae of Costa Rica. Novon 4: 350-361.

Six new species are described in this article, all endemic (as far as presently known) to Costa Rica: Habracanthus tilaranensis Gómez-Laur. (Pacific slope of Cordillera de Tilarán), Justicia aurantiimutata Hammel & Gómez-Laur. (Osa Peninsula), J. deaurata Hammel & Gómez-Laur. (Osa Peninsula), J. peninsularis Gómez-Laur. & Hammel (Golfo Dulce region), Kalbreyeriella rioquebradasiana Gómez-Laur. & Hammel (southern slope of Cerro de la Muerte at 1,700 m), and Razisea villosa Gómez-Laur. & Hammel (Sarapiquí region and Osa Peninsula). This report of Kalbreyeriella is the first for Costa Rica; however, the distinctness of this genus from Razisea is called into question. SEM photos are provided for all six species; this and the following article feature fine line drawings by our own Alvaro ("Bary") Fernández.

_____ & Q. Jiménez Madrigal. 1994. A new species of Galipea (Rutaceae_Cuspariinae) from Costa Rica and Nicaragua. Novon 4: 347-349.

Galipea dasysperma Gómez-Laur. & Q. Jiménez is probably most closely related to the largely or exclusively South American G. trifoliata Aubl. and G. jasminiflora Engl., but differs in having longer corollas, 2-seeded carpels, and hairy seeds. Differences from the sympatric G. granulosa Kallunki are not discussed. The new species ranges widely in Costa Rica, from 100-450 m elevation on both slopes; it also extends into southeastern Nicaragua.

Grant, J. R. 1994. The Tillandsia adpressa assemblage: a review and new combinations in Racinaea (Bromeliaceae: Tillandsioideae). Novon 4: 362-364.

The entire "assemblage" of five species is transferred to the splinter genus Racinaea (formerly Tillandsia subg. Pseudocatopsis), based on a traditional morphological analysis. The following changes are pertinent to the Costa Rican flora: Tillandsia schumanniana (Wittm.) Mez becomes Racinaea schumanniana (Wittm.) J. R. Grant, and Tillandsia adpressa André var. orthiantha (Standl.) L. B. Sm. undergoes a change in status and is recombined as Racinaea adpressa (André) J. R. Grant ssp. orthiantha (Standl.) J. R. Grant. A key to all the taxa in the "assemblage" is provided.

Hönig, D. 1994. In memoriam: Clarence Kl. Horich. Kakteen Sukk. 49: 195.

Supplies some of the data missing from our earlier obituary. Clarence was born on in Lüdenscheid, Germany, on 11 May, 1930, and moved to Costa Rica in late 1957. He died on 1 March of this year, at the age of 63 (sorry, old boy). In German.

Jenny, R. 1993. Monograph of the genus Gongora Ruiz & Pavón (transl. M. Sommer & C. d'Heureuse). Koeltz Scientific Books, Champaign, IL, U. S. A./Königstein, Germany.

This (according to the dust jacket) "is the first complete survey of the genus Gongora [Orchidaceae] since it was first described in 1794." The operative word is "survey," as this is in no sense a rigorous taxonomic revision. A new infrageneric classification is introduced, comprising three subgenera and seven sections, all published here for the first time. The total number of species accepted in the genus is 53, of which nine are definitely recorded from Costa Rica (only G. amparoana Schltr. is apparently endemic); one new species and one new combination are here validated. Distribution is reported in a highly confusing fashion (e.g., "Central America, Mexico" seems to indicate that a species is only known from Mexico), and there are no maps. Furthermore, virtually no specimens are cited (nor is there an index to exsiccatae), even though the author alludes to "the great mass of material I have received over the past years from all over the world" (p. 119). Most of the specimens that are cited are types, other historical collections in major orchid herbaria (K-L, W-R), or personal collections of the author or his contacts. Indeed, many species are known (to Jenny) only from the type, or a handful of collections from the type locality. This approach may be admired as profoundly conservative, but adds little to our previous knowledge of the genus. Gongora is one of the most frequently collected of the showy orchid genera; a wealth of Costa Rican material has been generated in recent years by the Manual project, and the same must be true for other regions. Yet none of this recent material appears to have been consulted by Jenny. Jenny notes (p. 10) that "it is only rarely possible to classify a badly deformed Gongora flower on an herbarium sheet, especially if it belongs to a species distinguishable from closely related species only by the scent components." Yet we are unable to find a single instance in any of Jenny's keys in which "scent components" are even so much as mentioned, let alone deployed as the sole character for distinguishing species. If good, modern collections with decent notes cannot be identified, what of century-old types? Not all dried orchid flowers are "badly deformed," and orchid flowers can be boiled up like any others.

This is essentially a taxonomic treatment of a handful of living specimens (often of unknown or dubious provenance) scattered among various horticultural collections; the extent to which it is relevant to wild populations remains to be seen. On the positive side, the work is well illustrated with line drawings and both black-and-white and color photographs, the latter mainly of living specimens. The biggest surprise for us: Gongora quinquenervis Ruiz & Pav., a name that has been used in virtually every regional Mesoamerican floristic treatment, properly refers (fide Jenny) to a species restricted to mid-elevations on the eastern slope of the Peruvian Andes (and possibly also occurring in Ecuador).

Jones, C. E. & D. M. Kearns. 1994. New species of Cyclanthera (Cucurbitaceae) from Mexico and Central America. Novon 4: 373-380.

Cyclanthera cogniauxii C. E. Jones & Kearns is endemic to Costa Rica, where it occurs at elevations of 1,500-2,500 in the central part of the country from the San Ramón region to the northern flanks of Cerro de la Muerte. The new species is compared to C. steyermarkii Standl. and C. monticola Gentry. Features excellent line-drawings.

Lehmiller, D. J. 1994 ['1993']. The identity of Crinum americanum L. (Amaryllidaceae). Herbertia 49: 58-66.

Billed as a "focused discussion" of this long-standing controversy, this turgid and exquisitely esoteric analysis ultimately leaves us in the dark. Though profoundly researched, this account is poorly edited, cluttered with misused terms, and confuses nomenclature and taxonomy throughout: "Despite its common use, C. erubescens does not qualify as a legitimate taxon because it is a superfluous entity." Evidently the author considers the familiar name Crinum erubescens Aiton to be illegitimate, and we gather that C. americanum L. is the name we should be using for the common Costa Rican species. Frankly, we do not follow the argument, and are more than a tad skeptical of the author's interpretations of the Code, particularly with regard to his principal conclusion that "Commelin 15.t.15 must be the type specimen [sic] for C. americanum L." We are more than happy to defer this thorny problem to Alan Meerow, our eminently capable Amaryllidaceae contributor.

Maas, P. J. M., E. A. Mennega & L. Y. T. Westra. 1994. Studies in Annonaceae. XXI. Index to species and infrageneric taxa of neotropical Annonaceae. Candollea 49: 389-481.

Bibliographic citations, type information, and (where appropriate) indication of synonymy. Some new combinations (but none relevant to Costa Rica).

Maas-van de Kamer, H. & P. J. M. Maas. 1994. Triuridopsis, a new monotypic genus in Triuridaceae. Pl. Syst. Evol. 192: 257-262.

Features an updated key to all the world's genera of Triuridaceae, including (without comment) Lacandonia. The new genus is Peruvian.

Murakami, N. & B. A. Schaal. 1994. Chloroplast DNA variation and the phylogeny of Asplenium sect. Hymenasplenium (Aspleniaceae) in the New World tropics. J. Pl. Res. 107: 245-251.

A companion piece to Murakami & Moran's 1993 monograph of the section (Ann. Missouri Bot. Gard. 80: 1-38). The epiphytic habit evolved only once in this group, and A. delitescens (Maxon) L. D. Gómez is "shown to have a polyphyletic origin." The "Journal of Plant Research" was formerly known as "The Botanical Magazine, Tokyo."

Pupulin, F. & D. E. Mora-Retana. 1994. A revision of the Costa Rican species of Trichocentrum (Orchidaceae). Selbyana 15: 87-103.

Seven species are recognized from Costa Rica, with two described as new: T. dianthum Pupulin & D. E. Mora, from 980-2,000 m on the Pacific slope in the vicinity of the Valle de El General; and T. costaricense D. E. Mora & Pupulin, from 700-1,170 m in the vicinity of San Ramón and Ciudad Quesada, Alajuela Province. Four species are Costa Rican endemics, and two others are shared with Panama; only T. capistratum Linden & Rchb. f. is relatively widespread (Costa Rica to Colombia and Venezuela). Key, distribution map, descriptions, illustrations. Most of the specimens cited, including the types of both new species, are unicates at USJ collected by the authors (a common sort of situation in Orchidaceae). The captions of the composite plates do not indicate what is depicted. The name Trichocentrum dianthum is replaced on the distribution map by "T. biflorum," presumably an earlier, rejected version.

Symptomatic of the lamentable collection history and documentation status of Orchidaceae is the fact that we currently have just one name in Trichocentrum (T. pfavii Rchb. f.) on our data-base.

Rico, M. de L. 1994. Four new species of Zygia (Leguminosae: Mimosoideae). Kew Bull. 49: 547-554.

Zygia rubiginosa L. Rico & Q. Jiménez, the only novelty represented in Costa Rica, is known from just two specimens from the Golfo Dulce region. It is compared to Z. confusa L. Rico, the only other Central American species of sect. Macrophylla. Our reprint features a hand-written correction of the type collection number, cited in two places as Jiménez & Jaén 6050; it should read "Jiménez & Jaén 680." Our careful perusal of the Code fails to reveal whether this significant error invalidates publication of the name, at least until such time as the correction is effectively published (or did we just do that?).

Zygia is a segregate of Pithecellobium sensu lato, a genus richly deserving fragmentation (for once we side with the splitters!). A detailed composite plate accompanies the species description.

Rueda, R. 1994. Systematics and evolution of the genus Petrea (Verbenaceae). Ann. Missouri Bot. Gard. 81: 610-652.

The 11 extant species are exclusively neotropical (fossils are known from California, Colorado and the country formerly known as Czechoslovakia). Only the widespread P. volubilis L. is reported from Costa Rica. According to Rueda, "this species is highly variable" in a number of characters; we would add that a wide range of this variability can be observed within Costa Rica. Features a cladistic analysis, illustrations, maps, and extensive specimen citations; no novelties published here.

Senghas, K. 1994. Gomphichis merzii Sengh. spec. nov. Orchidee (Hamburg) Suppl. 2: 25-42.

The genus Gomphichis (Orchidaceae), with two species now known from Costa Rica, is compared with the closely related Cranichis. Gomphichis merzii Senghas, known only from the type collected at 2700 m on Cerro de la Muerte, differs from G. costaricensis (Schltr.) Ames, Hubb. & C. Schweinf. in the form and structure of the labellum. The new species (which is terrestrial) is dedicated to Dr. Stefan Merz, whose recently completed doctoral research at Heidelberg involved comparative ecological and pharmaceutical studies of epiphytes on the Cerro and the Osa Peninsula. Features several color photographs of varying quality, all of living plants in cultivation and in the wild; these will be important in interpreting the species, since the type is a unicate at HEID. Complete description and pictorial floral analysis. Fully bilingual (German/English).

Shinwari, Z. K., H. Kato, R. Terauchi & S. Kawano. 1994. Phylogenetic relationships among genera in the Liliaceae_Asparagoideae_Polygonatae s. l. inferred from rbcL gene sequence data. Pl. Syst. Evol. 192: 263-277.

The molecular data "are concordant to" the classification of Dahlgren et al., and "suggest that" the transfer of Smilacina to Maianthemum by LaFrankie was "perhaps reasonable."

Smith, J. F. 1994. Systematics of Columnea section Pentadenia and section Stygnanthe (Gesneriaceae). Syst. Bot. Monogr. 44: 1-89.

This revision treats 27 species, including nine in sect. Pentadenia and 18 in sect. Stygnanthe. Collectively, these two sections correspond to the genus Pentadenia sensu Wiehler. Three new combinations, but no new taxa described here. The only species in either section reported from Costa Rica is Columnea angustata (Wiehler) L. E. Skog (sect. Stygnanthe), from a few mid-elevation sites on the Atlantic slope; C. nervosa (Klotzsch ex Oerst.) Hanst. (sect. Pentadenia) is spottily distributed from southern Mexico to southern Ecuador, but has not been collected in Costa Rica (though it is known from Prov. Chiriquí, Panama).

Features a cladistic analysis based on combined morphological and cpDNA restriction site data. Key to all six sections of Columnea, combined key to species of the two sections treated, distribution maps, illustrations, index to exsiccatae.

Taylor, C. T. 1994. Revision of Hillia (Rubiaceae). Ann. Missouri Bot. Gard. 81: 571-609.

_____. 1994. Revision of Tetragonia (Aizoaceae) in South America. Syst. Bot. 19: 575-589.

Hillia is a neotropical genus comprising 24 species arranged in five subgenera, of which three are here described; subg. Ravnia, with five species (two in Costa Rica), was dealt with previously by Taylor (Selbyana 11: 26-34, 1989; Ann. Missouri Bot. Gard. 78: 521-523, 1991), and is not further treated here. Of the remaining 19 species, six occur in Costa Rica; H. macrophylla Standl., reported only from the Cordillera de Guanacaste, is disjunct from Colombia. Features cladistic analysis, illustrations and maps; no new taxa below the subgeneric level.

The 11 New World species of Tetragonia are restricted to the arid regions of Chile and Peru, with the exception of the cultivated and widely naturalized T. tetragonoides (Pall.) Kuntze ("New Zealand spinach"). The latter is well known in Costa Rica and accounts for virtually all of the greens sold as "espinaca" in the marketplace; however, we have not recorded it as adventive. Features illustrations and maps; no novelties.

Tucker, G. C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monogr. 43: 1-213.

The sequel to Tucker's 1983 treatment of the Costa Rican and Panamanian species, which comprised Vol. 2 of this series. Only marginally relevant to us, but we note some inevitable conflicts with C. D. Adams's (1994) Flora Mesoamericana (FM) treatment, and a few apparent additions to our data-base: Cyperus reflexus Vahl, omitted from FM, is mentioned here as "in Costa Rica" (no specimen cited). C. entrerianus Boeck., omitted from FM but presumably included in C. luzulae (L.) Retz., is reported from "northwestern Costa Rica" (no specimen cited). C. thyrsiflorus Jungh. is attributed to Costa Rica by virtue of the Costa Rican type of its alleged synonym Mariscus tribrachiatus Liebm.; the latter name is cited in synonymy under Cyperus hermaphroditus (Jacq.) Standl. in FM, which does not mention C. thyrsiflorus. C. regiomontanus Britton is attributed to Costa Rica by virtue of the Guanacaste type of its alleged synonym Cyperus pittieri Boeck.; both names are treated in FM as synonyms of C. pallens (Liebm.) Standl. & Steyerm., which Tucker includes under C. thyrsiflorus. The technical segregate genera Pycreus and Torulinium, recognized in FM, are sensibly (we think) included in Cyperus by Tucker; Kyllinga (revised by Tucker in Rhodora 86: 507-538, 1984) is maintained.

Young, A. M. 1994. The Chocolate Tree: A Natural History of Cacao. Smithsonian Institution Press, Washington & London.

Allen Young is an entomologist, based at the Milwaukee Public Museum, who has worked in Costa Rica (mainly in the Sarapiquí region) for more than 25 years. This is a semi-popular account emphasizing pollination biology, the main thrust of Young's research. The central theme of the book's final chapter is "that successful natural pollination of cacao is linked to the ecology of the tropical rain forest," a fact that is seen to "bode well *or the future of both economic development and biological conservation in the lowland tropics." Other chapters address the history and geography of cacao cultivation. Illustrated with black-and-white photos as well as eight excellent color plates.

Zhu, G. 1994. Lectotypification and epitypification of Dracontium gigas (Seemann) Engler (Araceae). Novon 4: 404-407.

This, the first publication heralding Zhu's highly anticipated revision of this difficult genus, fixes the application of the name of what is perhaps the most familiar of the four Costa Rican species (the one in the arboretum at La Selva). Constraints of the Code require lectotypification (as opposed to neotypification), even though none of the available original material (comprising three separate elements in a plate) suffices to identify the species involved. Zhu obviates this impasse by designating an epitype in the form of a good, fertile herbarium specimen, the taxonomic identity of which can be reasonably interpreted. This is certainly the first publication to propose epitypification, a new provision (Art. 9.7) of the most recent (Tokyo) Code.



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