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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume VIII, Number 2, April 2001

News and Notes | Recent Treatments | Leaps and Bounds | Germane Literature | Season's Pick

Balick, M. J., M. H. Nee & D. E. Atha. 2000. Checklist of the vascular plants of Belize with common names and uses. Mem. New York Bot. Gard. 85: 1–246.

A rare and glorious occasion it is when we have two country-level floristic accounts for the Mesoamerican region to report in the same issue (see also under Stevens et al., below)! This checklist tallies 3408 spp. for Belize, in 209 families and 1219 genera. The largest families are Fabaceae s. l. (295 spp.), Orchidaceae (279 spp.), Poaceae (248 spp.), Asteraceae (153 spp.), and Cyperaceae (146 spp.), while the largest genera are Miconia and Rhynchospora (45 spp. each), followed by Psychotria (39 spp.), Paspalum (34 spp.), and Panicum (33 spp.). The supremacy of Fabaceae over Orchidaceae, the reverse of the situation in Costa Rica, is not surprising in this relatively dry country. What is surprising is that Belize boasts 41 endemic spp., as well as one endemic genus, the monotypic Schippia (Arecaceae). And who would have guessed the family with the most endemic Belizean spp. (six) to be Eriocaulaceae? The checklist itself is vouchered, with annotations indicating synonymy, literature references, uses, common names, and habit. Unfortunately, the families are ordered "phylogenetically" (although there is a comprehensive index to scientific names in the back). The introductory part includes a section on the history of botanical work in the country, emphasizing the role of prominent native collector Percy H. Gentle, to whom the volume is dedicated.

Brummitt, R. K. 2000. Report of the Committee for Spermatophyta: 50. Taxon 49: 799–808.

Three decisions affect names for taxa occurring in Costa Rica. Retypification of Poa pratensis L. (Poaceae) is recommended unanimously, because the current typification threatens the most widely accepted application of this well-known name. A proposal to conserve Zizyphus (Rhamnaceae) with that spelling, and as feminine, is not recommended; thus the more widely used Ziziphus will prevail, with feminine gender retained (conservation of the latter having been deemed unnecessary). Conservation of Scirpus miliaceus L. (Cyperaceae) with a new type is wisely not recommended, upholding (at least for the time being) the Flora mesoamericana usage of the name (followed in Jorge Gómez-Laurito's Manual treatment). Noting that "the world is now split into two camps" over the usage of this name, which has thus become "completely ambiguous," the Committee "sees the need to resolve the problem, but does not believe the present proposal is the best solution." In effect, acceptance of this proposal would have simply passed the buck from one camp to the other, rendering Flora mesoamericana and a host of other publications retroactively wrong. But in an odd twist, the Committee votes to invite a second proposal to reject the name altogether, thus extending (rather than terminating) nomenclatural instability, and begging a situation (acceptance of the second proposal) in which every flora in the world will become retroactively wrong! This seems like a step backward. Why not simply stand by the first lectotypification, "which should still stand today" (in the Committee's own words)? That is the function of types: to anchor the application of names. Now we are at sea.

Camelbeke, K., B. De Wilde, D. A. Simpson & P. Goetghebeur. 2000. Typifications in the genus Scleria P. J. Bergius (Cyperaceae). Kew Bull. 55: 993–996.

Scleria purdiei C. B. Clarke is the only name addressed here that applies to a sp. occurring in Costa Rica. The erroneous citation of a holotype in Flora mesoamericana Vol. 6 (1993) is noted, and a prior lectotypification is accepted. This apparently entails no consequences for the usage of the name.

Carnevali Fernández-Concha, G. & W. Fritz. 2000. The Maxillaria porrecta Lindley complex (Orchidaceae) in northeastern South America, including a new species. Selbyana 21: 144–150.

Although not strictly germane to our area, this article does include notes and Costa Rican specimen citations for three spp. represented in Central America. Of particular interest is the treatment of Maxillaria brunnea Linden & Rchb. f., a name currently in use for a sp. occurring in Costa Rica, as a synonym of M. porrecta Lindl. This change is also endorsed by Manual Maxillaria contributor John T. Atwood (SEL; in litt.). The new sp. (illustrated) is South American.

Davis, A. P., D. Bridson, C. Jarvis & R. Govaerts. 2001. The typification and characterization of the genus Psychotria L. (Rubiaceae). Bot. J. Linn. Soc. 135: 35–42.

The generic type, Psychotria asiatica L., was based on a mixture of New and Old World elements belonging to different spp. The authors can find no reason to reject the first (1964) valid lectotypification of the name, on the Old World (Asian) element. However, the true identity of this element has never been known. Here, the authors succeed in "unambiguously" linking the name Psychotria asiatica with wild populations from southeast China, previously identified as P. rubra (Lour.) Poir. This unequivocal typification of Psychotria on a particular sp. is obviously critical to the application of the name, especially considering recent overtures to fragment the traditional generic concept. That the type sp. is Asian could have unfortunate consequences for at least some New World members of this gigantic (perhaps 2000 spp.) and ecologically significant genus. Incidentally, the authors also establish that, contrary to previous assertions [see The Cutting Edge 6(4): 7–8, Oct. 1999], Psychotrophum P. Browne cannot be the correct name for the clade comprising Palicourea and Psychotria subgen. Heteropsychotria Steyerm.

Doyle, J. J., J. A. Chappill, C. D. Bailey & T. Kajita. 2000. Towards a comprehensive phylogeny of legumes: evidence from rbcL sequences and non-molecular data. In, P. S. Herendeen & A. Bruneau (editors), Advances in Legume Systematics 9: 1–20. The Royal Botanic Gardens, Kew, UK.

Some emerging taxonomic notions regarding Fabaceae are briefly discussed, largely on the basis of rbcL parsimony analysis of over 300 sequences, representing 77 outgroup genera and 194 genera of Fabaceae. The family is resolved as monophyletic, but with only "moderate" support. Subfamilies Mimosoideae and Papilionoideae (Faboideae) are supported (strongly and weakly, respectively) as monophyletic, but nested in a paraphyletic Caesalpinioideae "that includes the basal elements of the family." Monophyly of Cassia s. l. is not supported, lending credence to the controversial splinter genera Cassia s. str., Chamaecrista, and Senna.

González, F. & D. W. Stevenson. 2000. Perianth development and systematics of Aristolochia. Flora 195: 370–391.

The nature of the perianth in Aristolochia and some allied genera of Aristolochiaceae has apparently been a bone of contention, although we were not aware of the controversy. This careful study confirms that it corresponds, in all cases (42 spp. of Aristolochiaceae were investigated), to a trimerous calyx. We consulted Aristolochia descriptions in about a dozen standard floras and other references dating back to 1925, all but two of which guessed correctly. Only the Flora of Guatemala (Standley & Steyermark, 1946) and Bailey's Manual of cultivated plants (1949) opted conservatively for the appropriately noncommital "perianth."

Harley, R. M. & G. Paucar. 2000. List of species of tropical American Clinopodium (Labiatae), with new combinations. Kew Bull. 55: 917–927.

The cosmopolitan assemblage included by some authors in a broad concept of Satureja (Lamiaceae) has been eroding in recent years. The best molecular evidence, as interpreted by Cantino and Wagstaff [see the Cutting Edge 5(2): 3, Apr. 1998], restricts Satureja (along with Micromeria) to the Old World. New World taxa formerly included in Satureja would all be assigned to the genus Clinopodium, with the exception of the monospecific, South American Obtegomeria. In connection with Harley's forthcoming contribution to Klaus Kubitzki's Families and genera of vascular plants, the present paper provides the correct names in Clinopodium and Obtegomeria, with synonymy and typology, for all Central and South American spp. of the Satureja complex. This classification is accepted with some reservation; the authors clearly do not consider the matter settled, particularly in regard to a largely Andean group of long-flowered spp. often segregated as Gardoquia. Eleven new combinations are validated here, none affecting Costa Rica. For the record, the correct names in Clinopodium for the three "Satureja" spp. attributed to Costa Rica are: Clinopodium brownei (Sw.) Kuntze [Satureja brownei (Sw.) Briq.]; Clinopodium foliolosum (Benth.) Govaerts [Gardoquia foliolosum Benth.; G. discolor Kunth; Satureja discolor (Kunth) Briq., non Clinopodium discolor (Diels) C. Y. Wu & S. J. Hsuan ex H. W. Li]; and Clinopodium vimineum (L.) Kuntze (Satureja viminea L.). The last of these is a cultivated ornamental.

Hiepko, P. 2000. Opiliaceae. Fl. Neotrop. Monogr. 82: 1–55.

This obscure little family of the order Santalales receives the deluxe FNM treatment, including an extensive introductory section that fills nearly half the volume. Agonandra, endemic to the Neotropics and with 10 spp., is the only New World representative of Opiliaceae, a family with just 10 genera altogether. There are no changes for Costa Rica, with Agonandra macrocarpa L. O. Williams (ranging from Yucatán to the Guanacaste region and south to near Tivives) our only sp. Includes family, genus, and sp. descriptions, a key to spp., mostly representative specimen citations, line-drawings, distribution maps, and an index to exsiccatae. The introductory part, with three sections contributed by other authors, features SEM's of trichomes, flowers, and pollen grains.

Hughes, C. 1998. Monograph of Leucaena (Leguminosae-Mimosoideae). Syst. Bot. Monogr. 55: 1–244.

Pardon our belated coverage of this volume which, for some reason, did not hit our bookshelves until just now. No urgency, however, as there are no significant changes for Costa Rica's small contingent of Leucaena, a neotropical genus of 22 spp. strongly centered in Mexico and northern Central America. Our spp. are just two, Leucaena leucocephala (Lam.) de Wit and L. multicapitula Schery, with only the latter certainly indigenous. Leucaena leucocephala, long better-known by the misapplied L. glauca (L.) Benth., is represented in Costa Rica by subsp. glabrata (Rose) Zárate, a taxon of considerable economic importance (for ornament and edible pods and seeds) but unknown in the wild. This is an extravagant production, even by SBM standards, but with just one taxonomic novelty (a hybrid). We do learn something that slightly fortifies our primitive understanding of mimosoid genera: Leucaena is the only genus in the subfamily with hairy anthers, and though this feature is not universal in Leucaena, it is manifest in both Costa Rican spp.!

Jawad, J. T., D. S. Seigler & J. E. Ebinger. 2001 ['2000']. A systematic treatment of Acacia coulteri (Fabaceae, Mimosoideae) and similar species in the New World. Ann. Missouri Bot. Gard. 87: 528–548.

The informal "Acacia coulteri sp. group" comprises 13 spp. of Acacia ser. Vulgares Benth. that are characterized by a similar growth habit (erect shrubs or small trees), persistent stipules, and spicate inflorescences. The last-mentioned feature has provoked confusion with Piptadenia and some spp. of Lysiloma, from which a careful distinction is here made. Taxonomy of this group has been tangled up to now, with "up to half the specimens" in most herbaria misidentified. Fortunately, for our purposes, the group is practically restricted to Mexico, with only Acacia centralis (Britton & Rose) Lundell occurring in Costa Rica. Not surprisingly, the Costa Rican range of A. centralis is limited to the seasonally dry, northwestern corner of the country, where it ascends to ca. 1000 m. Includes a key to spp., full descriptions, representative specimen citations, and drawings of petiolar glands of several spp. Species circumscriptions are confirmed by principal components analysis. One new combination is validated.

Judziewicz, E. J., L. G. Clark, X. Londoño & M. J. Stern. 1999. American bamboos. Smithsonian Inst. Press, Washington, DC & London.

How do these things sneak by us? And what a valuable, authoritative, and beautifully illustrated volume it is! Covered here are all 41 genera of bamboos (Poaceae subfam. Bambusoideae) native to the New World, including the herbaceous bamboos (tribe Olyreae; 20 gen.) as well as the woody ones (tribe Bambuseae; 21 gen.). Thrown in, for good measure, are three monogeneric tribes of "basal grasses" (comprising Anomochloa, Pharus, and Streptochaeta) that were formerly classed among the bamboos (Streptogyna, now believed allied to Oryza and kin, is omitted). The taxonomic part of this work kicks off with a section that characterizes subfam. Bambusoideae and its two tribes, and discusses phylogeny, classification, and evolution in the group. An illustrated key to the five tribes treated precedes the formal generic entries. A description, discussion, and illustrated key to genera is provided for each of the two polygeneric tribes, under which the genera are considered alphabetically. Each genus entry includes a color distribution map, a description, an annotated list of spp., a discussion, and at least one line-drawing or photographic illustration. Generous introductory chapters delve into such topics as morphology, anatomy, growth and development, cytology, ecology and distribution, reproductive biology, ethnobotany and economic botany, cultivation and propagation, and pathology. Appendices compiling information on generic distribution by country, avian bamboo specialists, common names, and bamboo spp. in cultivation are followed by a glossary and separate indices to scientific names and subjects. This is a masterful piece of work by four individuals with as many lifetimes of first-hand experience.

Krings, A. 2000. Floristics and ecology of Mesoamerican montane climber communities: Monteverde, Costa Rica. Selbyana 21: 156–164.

The author, having already plied his trade at the Jardín Botánico Wilson [see The Cutting Edge 7(1): 7–8, Jan. 2000], now samples dicot woody climbers (at least 2.5 cm dbh) in the Monteverde Cloud Forest Reserve. Species composition and other parameters were analyzed in four 0.1 ha belt transects, which yielded a total of 19 spp. of dicot woody climbers. The most sp.-rich family was Asteraceae (hey, we're talking just three spp. here!), followed by Sapindaceae and Vitaceae (with two spp. each); similar results were obtained in four other Mesomerican transects, three in Mexico and one in Nicaragua. In terms of basal area, Fabaceae, Hydrangeaceae, and Piperaceae were most important in the Monteverde transects. Elsewhere, Celastraceae also figured prominently in this regard.

Lellinger, D. B. 2000. On the lectotypification of Danaea elliptica. Amer. Fern J. 90: 100–103.

The traditional application of the name Danaea elliptica Sm. (Marattiaceae) has been threatened by lectotypifications rendering it synonymous with D. nodosa (L.) Sm. Here, a case is made to overturn previous lectotypifications and designate a new lectotype that will allow the name to be used in its conventional sense. We wonder if this can be done unilaterally, without a formal proposal published in Taxon and voted upon by the Committee for Pteridophyta. We also wonder whether lectotypification is even an option in this case; the original author explicitly cited just one element, an illustration, which (at least according to some interpretations of the Code) must therefore be accepted as the holotype. In typical fashion, the Code (Art. 9.1 and Note 1) is ambiguous to a fault on this critical issue. Perhaps conservation of D. elliptica with a different type (Art. 14.9) would be a more appropriate strategy for maintaining current usage.

Liede, S. & A. Täuber. 2000. Sarcostemma R. Br. (Apocynaceae—Asclepiadoideae)—a controversial generic circumscription reconsidered: evidence from trnL-F spacers. Pl. Syst. Evol. 225: 133–140.

The results of this molecular study support morphological evidence previously adduced by the first author [see The Cutting Edge 3(4): 7, Oct. 1996] in concluding that Sarcostemma s. l. is polyphyletic and should be subdivided. Under this view, all Mesoamerican spp. in the group would be classed in the genus Funastrum.

Lombardi, J. A. 2000. Vitaceae—gêneros Ampelocissus, Ampelopsis e Cissus. Fl. Neotrop. Monogr. 80: 1–251.

This contribution covers three of the four genera of Vitaceae represented in the Neotropics, including Cissus, by far the largest. It seems strange indeed that the author did not make a clean sweep of things and include Vitis, which he allows has "few" neotropical spp. All three of the genera treated occur in both hemispheres, with a minority of their spp. in the New World. Accepted are four neotropical spp. of Ampelocissus, one of Ampelopsis, and 75 of Cissus. Ampelopsis does not reach Costa Rica, and Ampelocissus is represented only by A. javalensis (Seem.) W. D. Stevens & A. Pool, restricted to the Talamanca lowlands. The last-mentioned sp. would be a Costa Rican endemic, were it not for the lost (and dubiously conspecific) Nicaraguan holotype of Vitis javalensis Seem. [see The Cutting Edge 6(4): 8–9, Oct. 1999]. There are no endemics among the 13 spp. of Cissus attributed to Costa Rica. Features keys to spp. (but not to genera), thorough genus and sp. (but not family) descriptions, extensive (though often representative) specimen citations, useful composite line-drawings of all spp., distribution maps, a section on dubious and excluded names, and an index to exsiccatae. The taxa are conveniently arranged alphabetically. The introductory section is quite brief, by the standards of this series. None of the three new spp. or five newly designated lectotypes affects us; however, we are impacted by a few new synonymies, such as Cissus pseudosicyoides Croat under C. fuliginea Kunth. Entirely in Portuguese (including even the locality data from specimen labels originally in English, French, or Spanish).

Luer, C. A. 2000. Icones pleurothallidinarum XXI. Systematics of Masdevallia. Part Two. Monogr. Syst. Bot. Missouri Bot. Gard. 82: 265–518+.

The first part of this series contained introductory material relevant to the genus Masdevallia (Orchidaceae) as a whole, including a key to the 10 subgenera and their sections, plus a revision of Masdevallia subgen. Polyantha Luer [see The Cutting Edge 7(2): 10, Apr. 2000]. The present contribution treats 100 spp., 67 in four (of eight) sections of subgen. Masdevallia, the rest in subgen. Pygmaeia Luer (all four sections). Of the four sections of subgen. Masdevallia covered here, only sect. Reichenbachianae Woolward (with 13 spp. total) is represented in Costa Rica, by the following 9 spp. (all endemic): Masdevallia calura Rchb. f., M. chasei Luer, M. demissa Rchb. f., M. fulvescens Rolfe, M. marginella Rchb. f., M. reichenbachiana Endres ex Rchb. f., M. rolfeana Kraenzl., M. schroederiana Sander ex H. J. Veitch, and M. walteri Luer. Two other entities, of unknown provenance, are perhaps attributable to Costa Rica: Masdevallia epallax Königer (here spelled "enallax"), a putative hybrid, and M. lankesteriana Luer, both known only from cultivation. None of the four Costa Rican representatives of subgen. Pygmaeia is endemic, all ranging southward at least to Ecuador. These are: Masdevallia erinacea Rchb. f. and M. pygmaea Kraenzl., of sect. Pygmaeia, and M. schizopetala Kraenzl. and M. zahlbruckneri Kraenzl., of sect. Zahlbrucknerae (Luer) Luer, comb. nov. This last, plus two new spp. not occurring in Costa Rica, are the only novelties. Features descriptions of the subgenera, keys to sections and spp., and an index to scientific names.

McDade, L. A., T. F. Daniel, S. E. Masta & K. M. Riley. 2001 ['2000']. Phylogenetic relationships within the tribe Justicieae (Acanthaceae): evidence from molecular sequences, morphology, and cytology. Ann. Missouri Bot. Gard. 87: 435–458.

Combined analysis of nr ITS and cp trnL-trnF sequence data reveals (among other things) that the large (ca. 700 spp.), pantropical genus Justicia is not monophyletic. The Old World members form a grade, and the New World members are monophyletic only if several other genera are synonymized. The ca. 400 spp. of New World "justicioids" (including, e.g., Chaetochlamys, Megaskepasma, and Poikilacanthus) apparently have a sister-group relationship to subtribe Diclipterinae (Dicliptera, Hypoestes), although the authors were unable to identify any non-molecular synapomorphies to confirm the relationship. N.B.: Justicia hyssopifolia L., the generic type, is apparently an Old World sp.; therefore, if the conclusions of this study prove substantially correct, the well-known name Justicia could not be used for any New World plants!

Miller, J. T. & R. J. Bayer. 2000. Molecular phylogenetics of Acacia (Fabaceae: Mimosoideae) based on the chloroplast trnK/matK and nuclear Histone H3-D DNA sequences. In, P. S. Herendeen & A. Bruneau (editors), Advances in Legume Systematics 9: 181–200. The Royal Botanic Gardens, Kew, UK.

Cladistic analyses of sequence data from both chloroplast and nuclear DNA regions were congruent in clearly establishing the genus Acacia as polyphyletic, with three distinct lineages. However, the relationships among the major clades remain to be clearly elucidated (this study involved just 43 spp.). Any breakup of the vast (ca. 1200 spp.) genus Acacia could entail undesirable nomenclatural consequences for the Mesoamerican spp. [as discussed in The Cutting Edge 7(3): 8, Jul. 2000].

Morales, C. O. 1999. Sobre el holotipo de Kefersteinia retanae (Orchidaceae). Brenesia 52: 75–76.

The name Kefersteinia retanae G. Gerlach (Brenesia 41-42: 99–106. 1994) was invalidly published, because its author failed to provide any concrete reference to the type, beyond locality information, as required by Art. 37, Note 1, of the current Code [see The Cutting Edge 3(4): 5–6, Oct. 1996]. The problem is here remedied with information furnished (in litt.) by the original author. According to Art. 45.1, "when the various conditions for valid publication are not simultaneously fulfilled, the date is that on which the last is fulfilled." Thus, the priority of K. retanae must date from 1999, not 1994. Because the requisite "concrete reference" is quoted from Gerlach, we would treat this as an "in," rather than an "ex" citation, yielding: Kefersteinia retanae G. Gerlach, in C. O. Morales, Brenesia 52: 75. 1999.

Morales, J. F. 2001. Orquídeas, cactus y bromelias del bosque seco/Costa Rica/Orchids, cacti and bromeliads of the dry forest. Inst. Nac. de Biodiversidad, Sto. Domingo de Heredia, Costa Rica. 162 pp.

This, the latest in INBio's slick field-guide series, is one of the best of the lot. The subject matter is somewhat limited, there being relatively few spp. of orchids, cacti, and bromeliads in the drier portions of Costa Rica. But because of this, Chico's little guide is reasonably comprehensive. Moreover, these dry areas are heavily visited, and the plants are easy to see. Although this guide was written for the northwestern corner of the country (Áreas de Conservación Guanacaste and Tempisque), it ought to serve admirably for dry-forest habitats all the way to the Panamanian border. As throughout this series, the illustrations here play a starring role. The fine water-color paintings of bromeliads by North American artist Anita W. Cooper, previously showcased in Chico's bromeliad guide [The Cutting Edge 6(3): 8–9, Jul. 1999], are here supplemented by the exquisite work of INBio's own Claudia Aragón. Within each family, the genera and spp. are arranged alphabetically. The fully bilingual text features a diagnostic description for each sp., along with remarks on phenology, habitat, and distribution, and some miscellaneous, narrative commentary. This last is generally the most interesting part, as Chico imparts his hard-earned wisdom. The introductory sections characterize each of the three plant families, as well as the dry forest habitat type and related areas; there is also a no-nonsense discussion of illegal trade. A glossary, bibliography, and index to scientific names complete the volume. One minor quibble: it might have been a good idea to include Agavaceae (only a few spp., and superficially similar to some Bromeliaceae).

Olmstead, R. G., C. W. dePamphilis, A. D. Wolfe, N. D. Young, W. J. Elisons & P. A. Reeves. 2001. Disintegration of the Scrophulariaceae. Amer. J. Bot. 88: 348–361.

They keep beating this horse [see, e.g., The Cutting Edge 7(1): 12, Jan. 2000], but it shows no signs of dying anytime soon. Here, sequence analysis of three plastid genes in 39 genera, representing both Scrophulariaceae s. l. and other families of Lamiales, shows that "the traditional Scrophulariaceae are an unnatural assemblage of plants distributed throughout the phylogenetic tree of Lamiales." At least five distinct lineages were resolved, and the authors ask that we accept these at family rank. According to the recommended classification, Scrophulariaceae s. str., with "moderately strong" support, would include (to mention only genera occurring in Costa Rica) Alonsoa and Buddleja (but not other genera sometimes referred to Buddlejaceae, such as Peltanthera and Polypremum). A second family, here called Veronicaceae (but correctly Plantaginaceae), has "weak" (some might say nil) support. Among those genera assigned to Veronicaceae are Angelonia, Bacopa, Digitalis, Limosella, Tetranema, and Veronica, as well as Callitriche and Plantago. Orobanchaceae correspond to a "well supported" clade comprising almost exclusively holo- or hemiparasitic plants, not only Orobanchaceae s. str. (Conopholis, Orobanche, etc.), but also many genera traditionally assigned to Scrophulariaceae, e.g., Alectra, Buchnera, Castilleja, and Lamourouxia. Calceolaria and two oligotypic austral genera form "one of the most distinct and isolated of all clades in Lamiales," with a bootstrap value of 100%. The name Calceolariaceae (G. Don) Raf. ex Olmstead is validated for this clade. The fifth family, Stilbaceae, is strictly African.

This paper exemplifies a dilemma that molecular studies (and cladistic analyses in general) often pose for plant taxonomy. The authors propose that botanists accept their division of Scrophulariaceae s. l. into five smaller families, going even so far as to publish a new family name. They state (p. 354) that "...the primary conclusion from this...study is that Scrophulariaceae, as traditionally circumscribed, are not monophyletic." At the same time, they admit (p. 353) that "...the strength of the evidence is modest and we do not wish to overstate that conclusion." These studies were conceived because the authors were bothered by the "absence of uniquely defining traits" in Scrophulariaceae s. l.; yet, they are unable to identify any morphological or anatomical synapomorphies for the new groups, and do not seem bothered by that. In addition, the authors provide no "comprehensive morphological descriptions" for their segregate families. Why would the authors or editors of any flora be moved to adopt a modestly supported, yet radical, new classification scheme, in which neither the content nor the circumscription of the major taxa have been rigorously elucidated? Who will prepare keys and formal, parallel descriptions for these taxa, when even their proponents will not dare to do so? And what should be done with the ubiquitous loose ends (like Peltanthera, Polypremum, and Mimulus)? The implementation, in floras and other practical works, of many of the taxonomic changes envisioned by molecular workers and so often discussed in these pages is, we fear, a long way off. We are frustrated by this, especially because we appreciate the necessity for monophyletic taxa, and would like to be on the proverbial cutting edge with this sort of thing.

For better or worse, floras are probably the primary vehicle by which plant-taxonomic innovations are disseminated to the larger community of botanically inclined workers, including horticulturists, conservationists, ecologists, and even most plant taxonomists (those not specialized on a particular group). If it is important for molecular workers to have their conclusions embraced by this community, they must take the necessary time to iron out the kinks, tie up all the loose ends, and rigorously formalize their classifications. Otherwise, they are just spinning their wheels and waving their arms.

Rocha, J. F. & L. de J. Neves. 1999. Anatomia foliar de Hibiscus tiliaceus L. e Hibiscus pernambucensis Arruda (Malvaceae). Rodriguésia 51: 113–131.

Various differences in leaf anatomy uphold the notion that the two littoral spp. mentioned in the title (which are sympatric in Brazil) are indeed distinct. This suits us, because otherwise we would might have to revert to Hibiscus tiliaceus for the only entity occurring in Costa Rica, which presently answers to H. pernambucensis.

Simmons, M. P., C. C. Clevinger, V. Savolainen, R. H. Archer, S. Mathews & J. J. Doyle. 2001. Phylogeny of the Celastraceae inferred from phytochrome B gene sequence and morphology. Amer. J. Bot. 88: 313–325.

Simultaneous analysis of morphological and molecular data supports an earlier conclusion [see, e.g., The Cutting Edge 7(1): 11–12, Jan. 2000], based solely on morphology, that Hippocrateaceae is monophyletic, but nested within a paraphyletic Celastraceae. However, support for the monophyly of the former taxon is poor. The problematic genus Goupia (occurring in Mesoamerica, but not yet found in Costa Rica) is resolved as unrelated to Celastraceae (Goupiaceae was assigned to the order Malpighiales by the Angiosperm Phylogeny Group). Perrottetia, another aberrant genus sometimes classed in a separate family, can be accommodated in Celastraceae after all, though it has (together with Mortonia) a sister-group relationship to the rest of the family. Maytenus s. l. (including Gymnosporia) is resolved as polyphyletic, with three disparate groups, supporting the recognition of Gymnosporia at genus rank (but no New World spp. of Gymnosporia were sampled). This study involved 51 spp. from 38 genera of Celastraceae s. l.

Stevens, W. D., C. Ulloa Ulloa, A. Pool & O. M. Montiel (editores). 2001. Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot. Gard. 85: 1–2666.

Twenty-three years in the making, here it is all at once, the work of 177 contributors, seamlessly edited and complete in three hefty volumes! Seldom if ever has a rigorous flora for a tropical country of comparable diversity been produced so quickly. Only a total team effort (to which we have borne witness) could have accomplished such a task. This is truly an amazing and invaluable contribution to Mesoamerican floristics, and one that we will both profit from and be hard-pressed to match. The gauntlet has indeed been thrown, and the bar set high. Treated here in the generous format of standard floras are 5796 spp, in 1699 genera and 225 families. However, 442 of these spp. are not definitely known from the country, and included hypothetically. Apart from these, all the spp. are vouchered (a significant departure from most temperate floras). According to the analysis of project leader W. D. Stevens, the five most diverse families in Nicaragua are Orchidaceae (601 spp.), Poaceae (388 spp.), Fabaceae s. str. (311 spp.), Asteraceae (262 spp.), and Rubiaceae (229 spp.). Fabaceae s. l. (as according to our Manual) would have 525 spp. The most sp.-rich genera are Psychotria (69 spp.), Miconia (66 spp.), Piper (63 spp.), Epidendrum (60 spp.), and Cyperus (58 spp.). The geographical affinities of the overall flora are varied, but with a slight northern bias. Thirty-three taxa (32 spp. and one var.) are listed as apparently extinct in Nicaragua, and 761 spp. are known from the country by a single collection. Endemic "taxa" number 79, about 68% of which are restricted to the mountainous north-central region.
The page design of this flora is handsome and easy on the eyes. Each volume has its own indices to common and scientific names. The first volume features an introductory part, with sections on the history of the project, geography and history of the country, vegetation, phytogeography, floristic analysis, and history of botanical exploration. The Nicaraguan flora more or less resembles what the Manual will probably look like, mainly because we deliberately adopted substantially the same style. Still, there are several notable differences. First, as mentioned above, their flora appears on the stage already fully formed, whereas ours will come out in four parts published on different dates. Partly as a consequence of that, the angiosperm families are presented in strict alphabetical sequence in the Nicaraguan flora, with monocots and dicots intermixed. The Manual will observe the traditional monocot/dicot dichotomy, though more out of necessity than conviction. Whereas the Nicaraguan flora follows the Cronquist system throughout, the Manual does so only for dicots; for monocots, we opted for the Dahlgren system (a decision we now regret). Because the Costa Rican flora is so much better known, the Manual will formally treat very few, if any, hypothetical spp., i.e., those not vouchered by at least one specimen collected in the country. Also, the Nicaraguan flora provides more lavish treatments of infraspecific taxa than planned for the Manual. On the other side of the ledger, the Nicaraguan flora lacks family keys and illustrations, both of which the Manual will include, as well as a glossary, which we hope to include. But talk is cheap, and now we have our work cut out for us. We will be thrilled to even approximate the standard of excellence set by this landmark publication.

Szlachetko, D. L., R. González Tamayo & P. Rutkowski. 2000. Zhukowskia, a new orchid genus from Mesoamerica. Adansonia sér. 3, 22: 235–238.

Yet another new orchid genus is established, on the basis of technical, morphological details, and without the benefit of cladistic analysis. Zhukowskia harbors "at least three species," characterized as intermediate between Pelexia and Sarcoglottis. No consideration is given to the possibility of combining the last two genera. Just one sp. occurring in Costa Rica is affected: Sarcoglottis smithii (Rchb. f.) Schltr., which would become Zhukowskia smithii (Rchb. f.) Szlach., R. González & Rutk. A partial description of the new genus is provided and three new combinations are validated (but this is not a taxonomic revision).

Thorne, R. F. 2000. The classification and geography of the flowering plants: dicotyledons of the class Angiospermae (subclasses Magnoliidae, Ranunculidae, Caryophyllidae, Dilleniidae, Rosidae, Asteridae, and Lamiidae). Bot. Rev. (Lancaster) 66: 441–647.

As Bobby Zimmerman once said, the times they are a-changin'. Still carrying the torch for the profoundly influential Cronquist/Dahlgren/Tahktajan/Thorne generation of phylogenists, the author integrates a wealth of data from recent studies, including molecular and cladistic, in updating his last (1992) classification of dicots. The introduction references this recent literature (cited in a voluminous bibliography of over 2000 entries), and summarizes the changes in the author's system. The classification is presented in synoptic form, from subclass down to subfamily (tribes in Asteraceae), with annotations indicating the estimated number of genera and spp. in each taxon, the indigenous distribution of each family and subfamily, and the author's degree of confidence in his circumscriptions of higher taxa (family and above). Many of the family-level changes (the ones in which we are most interested) reflect conclusions already reported in these pages. Innovations in the Thorne system at the family level (and below) include: segregation of Siparunaceae from Monimiaceae; inclusion of Mahonia (Berberidaceae) in Berberis; absorption of Bombacaceae and much of Tiliaceae and Sterculiaceae into Malvaceae; recognition of Muntingiaceae as a distinct family; separation of Celtidaceae from Ulmaceae; removal of Drypetes from Euphorbiaceae as Putranjivaceae; insertion of Hippocrateaceae into Celastraceae (indeed, into an existing subfamily thereof); demotion of Lacistemataceae to subfamily status within Flacourtiaceae, and segregation of Kiggelariaceae from the latter family; removal from Simaroubaceae of Alvaradoa and Picramnia (to Picramniaceae) and Recchia (to Surianaceae); inclusion of Swartzia in subfam. Faboideae (Fabaceae); recognition of Brunelliaceae as distinct from Cunoniaceae, of Memecylaceae as distinct from Melastomataceae, of Phyllonomaceae as distinct from Escalloniaceae, of Nyssaceae as distinct from Cornaceae, of Hydroleaceae as distinct from Hydrophyllaceae, of Plocospermataceae and Strychnaceae as distinct from Loganiaceae, and of Petreaceae as distinct from Verbenaceae; removal of Hydrocotyle and related genera from Apiaceae to Araliaceae, and of Potalia from Loganiaceae to Gentianaceae; and finally, fragmentation of Scrophulariaceae s. l. into several families, including Scrophulariaceae s. str., Veronicaceae, Orobanchaceae, and Schlegeliaceae (with Plantaginaceae and Callitrichaceae maintained as distinct, and Buddlejaceae subsumed within Scrophulariaceae s. str.). Features an index to scientific names, and various useful tables.

Wagstaff, S. J., K. Martinson & U. Swenson. 2000. Divergence estimates of Tetrachondra hamiltonii and T. patagonica (Tetrachondraceae) and their implications for austral biogeography. New Zealand J. Bot. 38: 587–596.

Parsimony analysis of rbcL sequences supports previous studies in suggesting that the monospecific, herbaceous Polypremum (occurring in Costa Rica), bounced among Buddlejaceae, Loganiaceae, and Rubiaceae, is in fact most closely related to the enigmatic, subaquatic, ditypic, Gondwanan Tetrachondra. The latter genus has itself been shuttled among several families, including Boraginaceae, Crassulaceae, Lamiaceae, and Scrophulariaceae, but lately has come home to roost in its own family. That Polypremum and Tetrachondra are evidently sister genera, forming a distinct lineage within Lamiales, justifies the inclusion of the former genus in Tetrachondraceae, as already proposed by Oxelman et al. (Syst. Bot. 24: 164–182. 1999). But although the two genera are quite similar morphologically and phytochemically, they differ in important respects, especially fruit type (many-seeded capsules in Polypremum; one-seeded mericarps in Tetrachondra).

Windisch, P. G. 2000. On the itineraries of Alfred and Alexander Curt Brade in Costa Rica. Amer. Fern J. 90: 108–109.

German-born botanist Alexander Curt Brade, who ultimately achieved glory in Brazil, made an important early collection series in Costa Rica, where his brother Alfred maintained a horticultural enterprise. This brief contribution presents a detailed itinerary of the Brade brothers' explorations, beginning with Curt's arrival at Limón in February, 1908, and ending with his departure from the same port in August, 1910. The Brades made a specialty of orchids and ferns.

ERRATUM: In our last issue, we reported ("Germane Literature," under Ebinger et al.) that the sp. formerly known in Costa Rica as Acacia farnesiana (L.) Willd. (Fabaceae: Mimosoideae) is henceforth to be called A. guanacastensis (H. D. Clarke, Seigler & Ebinger) Ebinger, Seigler & H. D. Clarke. This is incorrect. The real Acacia farnesiana does indeed occur in Costa Rica, and grows side-by-side with A. guanacastensis in the northwestern part of the country.



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