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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XIII, Number 3, July 2006

News and Notes | Recent Treatments | Germane Literature | Season's Pick | Annotate your copy

Achille, F., T. J. Motley, P. P. Lowry II & J. Jérémie.  2006. Polyphyly in Guettarda L. (Rubiaceae, Guettardeae) based on nrDNA ITS sequence data.  Ann. Missouri Bot. Gard. 93: 103-121.

The 21 spp. of Guettarda involved in this study wound up in four separate clades, with the majority of neotropical spp. in the same clade as the generic type (the paleotropical G. speciosa L.).  This would presumably secure membership in Guettarda for most of the 10 or so spp. occurring in Costa Rica; however, since only three of these were included in the study, we can be certain only of G. macrosperma Donn. Sm.  This “Guettarda sensu stricto” clade is “characterized by dichasial cymes (sometimes reduced to solitary flowers) and a caducous calyx with a circular abscission zone.”  At least three other Costa Rican congeners could well end up in other genera, based on this evidence.  Guettarda crispiflora Vahl and (by association) the closely related G. tournefortiopsis (Rusby) Standl. occur in a separate clade, sister to the oligotypic Hawaiian genus Bobea.  Were these Guettarda spp. to be accorded generic status (separate from Bobea), Tournefortiopsis Rusby would be the operative name.  Also remote from the Guettarda s. str. clade is G. acreana K. Krause, sister to the type sp. of Stenostomum, a genus comprising most of the neotropical spp. formerly referred to Antirhea (now restricted to the Old World).  The authors conclude that “it might thus be appropriate to include...G. acreana within Stenostomum.”  Their results also confirm the distinctness of Antirhea s. str. from Stenostomum (as well as from Pittoniotis).

Andersson, L., M. Kocsis & R. Eriksson.  2006. Relationships of the genus Azorella (Apiaceae) and other hydrocotyloids inferred from sequence variation in three plastid markers. Taxon 55: 270-280.

The genus Azorella, as presently conceived, is paraphyletic; however, no resolution is ventured, in lieu of "denser sampling and more informative data." We cannot even speculate as to the fate of the sole Costa Rican representative, A. biloba (Schltdl.) Wedd., because the generic type (A. filamentosa Lam.) was not included in the study.  Consistent with previous results, these analyses portray Hydrocotyle as "closely related to core Araliaceae."

Andersson, S. 2006. On the phylogeny of the genus Calceolaria (Calceolariaceae) as inferred from ITS and plastid matK sequences.  Taxon 55: 125-137.

Tribe Calceolarieae, consisting of Calceolaria and two minor satellite genera (neither occurring in Costa Rica), has traditionally been classed in Scrophulariaceae, but was recently elevated to family rank on the basis of molecular studies [see The Cutting Edge 8(2): 12–13, Apr. 2001].  The present analyses provide strong support for Calceolarieae as monophyletic (and for one of the satellite genera as nested within Calceolaria), and moderate support for a sister-group relationship with Gesneriaceae.  If the last-mentioned hypothesis proves to hold water, we’d vote to include Calceolarieae in Gesneriaceae; what the hey, Calceolaria looks enough like a gesneriad to us, and who needs another little stray family floating around?

Christenhusz, M. J. M. & H. Tuomisto.  2005. Some notes on the taxonomy, biogeography and ecology of Danaea (Marattiaceae).  Fern Gaz. 17: 217-222.

Danaea was monographed very recently, with 17 spp. accepted for the entire genus [see The Cutting Edge 12(2): 14, Apr. 2005].  According to the present authors, who are revising Danaea yet again, that monograph featured “a large degree of synonymization...with which we do not agree.”  Indeed, the second author has already published a paper discriminating 18 Danaea spp. in Ecuador alone, and two other papers proposing eight additional new spp. from various regions are cited as “in press.”  Having said all of that, the purpose of the present paper is not at all obvious to us.  We do learn that the spp. known in Costa Rica as Danaea nodosa (L.) Sm. will probably wind up being called D. media Liebm., and that D. alata Sm. (attributed to Costa Rica in the aforementioned monograph) is in fact “confined to the Lesser Antilles.”  For anything more substantive than that, we’ll have to await the formal revision.

Christenson, E. A.  2006. Brevilongium, a new orchid genus from the Neotropics.  Austral. Orchid. Rev. 71(1): 32-33.

Here the author presents the original English manuscript of a paper already published in French (and reviewed in this column of our last issue).

-- . 2006. Das frühere Phragmipedium caudatum aus Panama/The former Phragmipedium from Panama.  J. Orchideenfr. 13: 141-143.

This story now rivals or even trumps the celebrated case of Sphagneticola trilobata (L.) Pruski [see The Cutting Edge 4(1): 7, Jan. 1997].  The rare, epiphytic orchid long known as Phragmipedium caudatum (Lindl.) Rolfe or P. warszewiczianum (Rchb. f.) Garay was rechristened in 1999 as “Phragmipedium humboldtii (Warsz. ex Rchb. f.) J. T. Atwood & Dressler” (the name used in the Manual), on the grounds that the first two names had been misapplied.  In the same paper, the Mexican Phragmipedium exstaminodium Castaño, Hágsater & J. Aguirre was subordinated to P. humboldtii at subspecific rank [see The Cutting Edge 6(3): 3, Jul. 1999].  More recently, the name P. humboldtii was revealed as invalid, and replaced by the newly coined Phragmipedium popowii Braem, Ohlund & Quéné [see The Cutting Edge 12(1): 4–5, Jan. 2005].  There we thought the story had ended, but we missed an important subsequent paper by Manual Orchidaceae coordinator Robert L. Dressler (Orchid Digest 69: 86–91.  2005), acknowledging his error with respect to P. humboldtii.  However, he continued to regard P. exstaminodium as distinct only at the subspecific level from the taxon known (for the previous year or so) as P. popowii; thus the latter name became a synonym of P. exstaminodium (which has priority at sp. rank) under Dressler’s classification.  But rather than validate a new combination based on Phragmipedium popowii for the southern (Panama and perhaps Costa Rica) subsp. of P. exstaminodium, Dressler created a new taxon name, P. exstaminodium subsp. warszewiczii Dressler.  Cased closed?  Not on your life!  Now Florida orchid maven Eric Christenson has unearthed a much older name, overlooked for more than a century, that exerts priority over all contenders.  This is Selenipedium warscewiczii Rchb. f. (1873), upon which the author bases the combination Phragmipedium warscewiczii (Rchb. f.) Christenson (and yes, Virginia, substantival and adjectival epithets based on the same personal name are permissible within a single genus).  Christenson does agree with Dressler’s notion of a subspecific relationship between the Mexican and Panamanian taxa already alluded to, prompting another new combination, Phragmipedium warscewiczii subsp. exstaminodium (Castaño, Hágsater & J. Aguirre) Christenson.  This latter action suggests that Selenipedium warscewiczii was based on material corresponding to the Panamanian subsp. (as seems probable from the epithet alone), although Christenson, rather curiously, avoids any mention of types.  Indeed, he provides no evidence at all for his equation of S. warscewiczii with P. exstaminodium sensu Dressler, other than to state that the former name is “based on the earlier Cypripedium caudatum var. roseum Hort.,” itself an ambiguously typified entity (in all fairness, though, we should note that Dressler himself effectively cited the latter name as a synonym of P. exstaminodium).

This is, taxonomically as well as nomenclaturally, one of the most volatile and confusing situations we have yet encountered.  If we accept Christenson’s interpretation, the correct name for the taxon called Phragmipedium humboldtii subsp. humboldtii in the Manual is currently P. warscewiczii (Rchb. f.) Christenson subsp. warscewiczii (though you may want to hold off on annotating your copies).  But even if the story really does end here, we are left with a nomenclatural rubble of mind-boggling perplexity, including three separate, heterotypic entities based on the same personal name (variously spelled) in the same genus, all of which have been used at one time or another for the same taxon (we refer to Phragmipedium warscewiczii, P. warszewiczianum, and P. exstaminodium subsp. warszewiczii), not to mention both Phragmipedium exstaminodium subsp. warszewiczii and P. warscewiczii subsp. exstaminodium (which apply to different taxa).  By the way, although Christenson uses the spelling “warszewiczii,” we consulted the protologue and established that Selenipedium warscewiczii is correct (hence Phragmipedium warscewiczii, etc.).

Clark, J. L., P. S. Herendeen, L. E. Skog & E. A. Zimmer.  2006. Phylogenetic relationships and generic boundaries in the Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data.  Taxon 55: 313-336.

These are the raw data on which the first author’s reestablishment of Glossoloma, restricting Alloplectus to just seven spp. [see The Cutting Edge13(2): 3–4, Apr. 2006], was predicated.  Episcieae is the largest tribe of Gesneriaceae, with 22 genera, all but one of which (the monospecific Lampadaria) were represented in this study.  Most of the sampled genera are supported as monophyletic, in most cases strongly (though Drymonia received only weak support).  Especially significant is the strong support for Columnea as a single monophyletic genus, as opposed to the five segregate genera distinguished by the late Hans WiehlerAlsobia and Episcia are upheld as separate genera, while Oerstedina and Rufodorsia are strongly supported as sister taxa (though only one sp. each was studied).  Notably, the evidence for Glossoloma as distinct from Alloplectus—one of the primary interests of the authors—is equivocal.  One of the data-sets weakly supports a sister-group relationship of these taxa, suggesting that “they could be treated as congeners following traditional classification.”  Nonetheless, the authors “argue that they should be treated as distinct genera because their recognition is practical since they are morphologically distinct with well defined morphological synapomorphies.”  We would counter with our maxim that, unless cladistic data provide strong evidence to the contrary, the nomenclatural status quo should prevail.

Dalton, R.  2006. Cashing in on the rich coast.  Nature 442: 567-569.

INBio gets some much-needed attention in a big-time journal, and an even more sorely needed shot in the arm in the form of $3.5 million in NIH funding over the next four years. This would appear to bode well for a worthy institution that had recently fallen on hard times, as detailed herein. At least from our reading of this article, the future looks bright, and we sincerely hope that it pans out that way!

Dathe, S. & H. Dietrich.  2006. Comparative molecular and morphological studies in selected Maxillariinae orchids.   Willdenowia 36 (Special Issue): 89-102.

Analyses of nuclear ribosomal DNA sequences support Orchidaceae subtribe Maxillariinae as monophyletic.  However, the pivotal genus Maxillaria is itself “clearly paraphyletic”; nested within it are Cryptocentrum, Mormolyca, and Trigonidium, all generally accepted as distinct genera, as well as numerous occasional or former segregates, e.g., Camaridium, Chrysocycnis, Heterotaxis, Neo-urbania, and OrnithidiumCamaridium, Heterotaxis, and Ornithidum are themselves polyphyletic, contrary to previous studies that spurred the recent reestablishment of Heterotaxis [see The Cutting Edge 13(1): 10, Jan. 2006].  The authors conclude that “further multidisciplinary studies are required to decide if and how Maxillaria s. l. can be divided in smaller genera.”  Well, obviously it can be!  The other question we would ask is not how, but why?

Drábková, L., J. Kirschner & Č. Vlček.  2006. Phylogenetic relationships within Luzula DC. and Juncus L. (Juncaceae):   a comparison of phylogenetic signals of trnL-trnF intergenic spacer, trnL intron and rbcL plastome sequence data.  Cladistics 22: 132-143.

This study corroborates a previous one by the same lab [see The Cutting Edge 11(2): 6, Apr. 2004] in portraying Luzula as monophyletic, but Juncus as non-monophyletic.  The problem for Juncus is that the oligotypic, Southern Hemisphere genera Distichia, Oxychloe, Patosia, and Rostkovia are all nested within it, in a sister-group relationship with a clade containing the generic type of Juncus.  Taking these cladograms at face value, the only two Costa Rican representatives of Juncus included in this study (J. bufonius L. and J. effusus L.) would find themselves in a different genus under any reclassification scheme that involved splitting.

Fior, S., P. O. Karis, G. Casazza, L. Minuto & F. Sala.  2006. Molecular phylogeny of the Caryophyllaceae (Caryophyllales) inferred from chloroplast matK and nuclear rDNA ITS sequences.  Amer. J. Bot. 93: 399-411.

We gather that Arenaria is quite seriously polyphyletic; however, we cannot assess the potential impact for us, as no spp. occurring in Costa Rica were included in the study.

González, J.  2005. Plantas comunes de la Reserva Biológica Hitoy Cerere/Costa Rica/Common plants of the Hitoy Cerere Biological Reserve.  Edit. INBio, Santo Domingo de Heredia, Costa Rica.  245 pp.

This most recent of INBio field guides presents superb color photos and informed discussions for 71 plant spp. occurring at the Reserva Biológica Hitoy Cerere, an area of nearly 10,000 ha, on the Caribbean slope of the Cordillera de Talamanca ranging from ca. 100–1025 m elevation.  The spp. accounts are grouped according to eight categories, based on habit and basic leaf features.  Each sp. photo in the main section is paired with text providing a brief description, diagnosis, summaries of distribution and phenology, and “notes.”  Featured families are described in a separate section toward the back, followed by miscellaneous photos of other spp. mentioned in the volume (unaccompanied by text), a glossary, and an index.  In his introduction, the author describes topography and vegetation of the reserve, listing many common spp.  Perplexingly, numerous spp. alleged as conspicuously abundant in the introduction are not included in the main section of the work, e.g., Acacia ruddiae D. H. Janzen and Zygia longifolia (Humb. & Bonpl. ex Willd.) Britton & Rose (Acacia/Mimosoideae), Fleischmannia sideritides (Benth.) R. M. King & H. Rob. (Asteraceae), and Gynerium sagittatum (Aubl.) P. Beauv. and Pennisetum purpureum Schumach. (Poaceae).  Also omitted are a few local specialties, viz., Herpetacanthus stenophyllus Gómez-Laur. & Grayum (Acanthaceae) and Nyssa talamancana Hammel & N. Zamora (Cornaceae).  On the other hand, we do get excellent color photos (possibly the first ever published) of several rare or little-known spp., e.g., Clarisia mexicana (Liebm.) Lanj. (Moraceae), Cleidion castaneifolium Müll. Arg. (Euphorbiaceae), and Lecointea amazonica Ducke (Fabaceae/Faboideae).  For those fortunate enough to visit Hitoy Cerere, this volume will be an indispensable companion.  Fully bilingual (Spanish/English).

Hágsater, E.  2006. The genus Epidendrum.  Part 5.  Species new & old in Epidendrum.  Icon. Orchid. 8: pl. 801-900.

Another century of Epidendrum spp. is paraded forth, but this time only 32 are new; the rest are “old species which are now botanically illustrated with their precise determination based on the most recent information available.”  Eleven of the new spp. are from Costa Rica:  Epidendrum acroamparoanum Hágsater & L. Sánchez, E. atypicum Hágsater & E. Santiago, E. chalcochromum Hágsater, E. dolichochlamys Hágsater & E. Santiago, E. haberi Hágsater & L. Sánchez, E. microrigidiflorum Hágsater, E. pachytepalum Hágsater & E. Santiago, E. philowercklei Hágsater & E. Santiago, E. platystomoides Hágsater & L. Sánchez, E. salpichlamys Hágsater & E. Santiago, and E. vulgoamparoanum Hágsater & L. Sánchez (the last applying to the sp. previously known as E. amparoanum Schltr., here revealed as a synonym of E. barbeyanum Kraenzl.).  These novelties are all Costa Rican endemics with the exception of E. microrigidiflorum, E. pachytepalum, E. philowercklei, and E. vulgoamparoanum, all shared with Panama.  Curiously, most are from páramo or high elevations, habitats that had not previously yielded many Epidendrum spp.  Among the “old” Epidendrum spp. are most of those that were treated in the Manual in the genus Oerstedella, since abandoned by Hágsater [see The Cutting Edge 12(3): 5, Jul. 2005].  The sp. entries are alphabetically arranged, and each comprises a full page, front and back; one side features a composite line drawing, the other typology and synonymy, a formal, bilingual (Spanish/English) description, specimen citations, a distribution summary and diagnostic paragraph, information on conservation status (usually indeterminable) and etymology, and a range map.  In many cases, the specimen citations are shunted to an appendix (oddly, in the front of the volume).  Adding Pupulin’s three new spp. (see this column), our running total of new orchid spp. described from Costa Rica since the inception of this newsletter jumps to 240.  Considering also the merger of Oerstedella, the Costa Rican sp.-total for Epidendrum now hovers in the vicinity of 185 (vs. 158 in the era of the Manual Orchidaceae treatment).

Hammel, B.  2005. Plantas ornamentales nativas de/Costa Rica/ Native ornamental plants.  Ed. 3.  Edit. INBio, Santo Domingo de Heredia, Costa Rica.  269 pp.

Already in its third edition, the best-selling title in INBio’s field-guide series has a fresh, new look.  Compared with the first edition [see The Cutting Edge 6(4): 6, Oct. 1999], the format is larger (sleeker and not so chunky), and the relative positions of the English and Spanish texts have been changed.  Inevitably, some of the photos have been upgraded.  An appendix provides illustrations of 41 additional spp., including some that have never or rarely been cultivated, as a means “to demonstrate [the] rich potential of Costa Rican native plants.”  A key component of any Costa Rican plant library!

Hofreiter, A.  2006. Bomarea edulis (Tussac) Herb., a nearly forgotten pre-Columbian cultivated plant and its closest relatives (Alstroemeriaceae).   Feddes Repert. 117: 85-95.

Bomarea edulis was cultivated in pre-Columbian times for its tubers (up to 5 cm in diameter), which “are eaten like roasted potatoes and also a white cream is produced from them.”  These are said to be still consumed in parts of Mexico and Guatemala, as are those of B. acutifolia (Link & Otto) Herb.  This synoptic treatment of B. edulis and four similar and closely related spp. features a key to separate the four subgenera of Bomarea, along with the five spp. here treated (all members of the autonymic subgenus, which harbors all the spp. with edible tubers).  Of the five chosen spp., only B. edulis and B. obovata Herb. occur in Costa Rica (for some reason, B. acutifolia is left out).  Each sp. entry includes full synonymy and typology and exsiccatae citations, but only B. edulis merits a detailed morphological description, distribution summary, and discussion.  Illustrated with photos from herbarium specimens and a range map for B. edulis.

Jenny, R.  2003–2004. The genus Stanhopea.  Caesiana 21-22: 1-291.

Although the author refers to this as a “revision,” it does not quite meet our criteria, as there is no key to spp. and no new taxa.  What it is is a rich compendium of information on all 56 spp. of the neotropical genus Stanhopea (Orchidaceae), as well as one subsp. and six hybrids; for good measure, both spp. of the segregate genus Embreea are thrown in.  Each taxon entry includes synonymy, a detailed description, specimen citations, an often extensive discussion, and illustrations (mostly color photos from life).   The introductory part pays lip service to phylogeny (by reproducing borrowed cladograms), and also addresses generic characteristics, taxonomic history, pollination, culture, and hybridization and breeding.  This is presented in two consecutively paginated parts (with the spp. ordered alphabetically), and may have been published originally as two separate issues of Caesiana; however, the MO copy is bound as a single volume.

Jordaan, M. & A. E. van Wyk.  2006. Sectional classification of Gymnosporia (Celastraceae), with notes on the nomenclatural and taxonomic history of the genus.  Taxon 55: 515-525.

Gymnosporia is once again characterized as “an Old World genus” by these South African workers [see The Cutting Edge 6(4): 7, Oct. 1999], but this time they do acknowledge the existence of “three non-thorny species from the New World” (including the Costa Rican G. haberiana Hammel), the generic affinity of which “needs to be critically examined..., as these taxa may constitute a new genus.”  The authors admit that both the Old and New World spp. are “dioecious trees and share the same dichasial cymose inflorescence and capsule types.”  However, in addition to lacking thorns, the New World spp. differ in having flowers “tending to be 4-merous” and in being “large rain-forest trees, up to 30 m high.”  By contrast, the Old World Gymnosporia spp. have flowers that are “usually 5-merous” and “usually grow in arid regions, in savanna and open thicket habitats.”

Kappelle, M.  2006. Structure and composition of Costa Rican montane oak forests.  Pp. 127–139 in, M. Kappelle (ed.), Ecology and conservation of neotropical montane oak forests.  Ecol. Stud. Analysis Synth. 185: 1-483.

This contribution documents the geographical and elevational distribution of Quercus spp. (Fagaceae) in Costa Rica, as well as the phytogeography, structure, and floristic composition of Costa Rican oak forests.  Surprisingly (to us), nearly 75% of the vascular plant genera in these forests have a tropical distribution, the greatest affinity being with the Colombian Andes.  About 1300 vascular plant spp. have been recorded from Costa Rican oak forests, including about 750 spp. of dicots, 250 spp. of monocots, and over 300 spp. of pteridophytes.  Nearly 500 spp. are woody.  The largest angiosperm families are Asteraceae (60+ spp.), Rubiaceae (50+ spp.), Piperaceae (40+ spp.), Lauraceae and Melastomataceae (each with 35+ spp), and Ericaceae and Solanaceae (each with 30+ spp.).  Considering all vascular plants, the most diverse genera are Elaphoglossum (30+ spp.), Peperomia (25+ spp.), and Maxillaria and Miconia (each with 20+ spp.).  We focus on this chapter as representative of this rich and multi-faceted volume, which addresses a fascinating range of topics relevant to neotropical oak forests.  More than half of its 34 chapters deal largely or exclusively with Costa Rica.

Kathriarachchi, H., R. Samuel, P. Hoffmann, J. Mlinarec, K. J. Wurdack, H. Ralimanana, T. F. Stuessy & M. W. Chase.  2006. Phylogenetics of tribe Phyllantheae (Phyllanthaceae; Euphorbiaceae sensu lato) based on nrITS and plastid matK DNA sequence data.  Amer. J. Bot. 93: 637-655.

Phyllanthus “in its traditional circumsciption” is paraphyletic, with several smaller genera including Breynia and Sauropus (both cultivated in Costa Rica) nested therein.  The authors advocate lumping these smaller genera to fashion a new Phyllanthus s. l. with “an estimated 1269 species, making it one of the ‘giant’ genera.”  Assuming their cladograms to be substantially correct, the pursuit of any option involving splitting would segregate the Phyllanthus spp. occurring in Costa Rica into at least four smaller genera.  Phyllanthus acuminatus Vahl, P. anisolobus Müll. Arg, P. caroliniensis Walter, P. graveolens Kunth, P. niruri L., and P. salviifolius Kunth might all remain in Phyllanthus s. str.; however, P. acidus (L.) Skeels (cultivated in Costa Rica), P. amarus Schumach. & Thonn., and P. urinaria L. would have to be assigned to three different genera.

Kriebel, R.  2006. A new species and notes on Drymonia (Gesneriaceae) from Costa Rica.  Novon 16: 65-86.

The new sp., Drymonia rubripilosa Kriebel, is endemic to Costa Rica, where it occurs at 50–300 m elevation on the Atlantic slope of the Cordillera Central and northern Cordillera de Talamanca, and on the Llanura de San Carlos.  It was first found and distinguished at the Estación Biológica La Selva, where it is sympatric with the very similar D. multiflora (Oerst. ex Hanst.) Wiehler.  As implied by its epithet, Drymonia rubripilosa differs from D. multiflora (which typically occurs at higher elevations) in its reddish (vs. whitish) pubescence, but also by its generally larger leaf-blades, shorter pedicels, and larger calyx lobes.  The new sp. was already included as provisionally named in the author’s Manual draft treatment of Gesneriaceae [see The Cutting Edge 12(3): 1–2, Jul. 2005].  Illustrated with an excellent composite line drawing by INBio’s Claudia Aragón.

Lammers, T. G.  2006. Validation of names at subspecific rank in North American Campanulaceae.  Novon 16: 69-73.

Inevitably, some of these novelties impact us in an oblique manner.  The validation of two Mexican subspp. (elevated from varietal rank) in Lobelia irasuensis Planch. & Oerst. automatically establishes the autonymic subsp. for use in southern Central America.  Similarly, the subordination of Triodanis biflora (Ruiz & Pav.) Greene to T. perfoliata (L.) Nieuwl. as a subsp. creates the corresponding autonym, which would presumably apply to the Costa Rican populations of T. perfoliata.

Maas, P. J. M. & L. Y. T. Westra.  2005. Neotropical plant families.  3 ed.  Gantner Verlag, Ruggell [Liechtenstein].  358 pp.

This handy reference provides brief descriptions and other critical details (number of genera and spp., distribution, uses, literature, etc.) for every family of vascular plants known to occur in the Neotropics.  Many are illustrated with drawings, but there is no key of any sort.  The big innovation, in the latest edition of this vademecum, is that all angiosperm family circumscriptions are now according to the Angiosperm Phylogeny Group (APG) system (the previous edition having followed Cronquist).  Families are grouped by order and higher groups (these, for angiosperms, also following APG); as far as we can tell, alphabetical sequencing is not observed at any level.  Includes a glossary and indices to vernacular and scientific names.

Maas-van de Kamer, H.  2006. (1721) Proposal to conserve Canna tuerckheimii, nom. cons. (Cannaceae) against an additional name, C. sylvestris.  Taxon 55: 531.

The author has already successfully proposed that Canna tuerckheimii Kraenzl. (used as an accepted name in the Manual) be conserved against three earlier names [see The Cutting Edge 12(1): 10, Jan. 2005].  Here she tacks on one more, C. sylvestris Roscoe, at least as obscure as the names already rejected.

Nicolson, D. H.  2006. (1720) Proposal to conserve the name Caladium (Araceae) with a conserved type.  Taxon 55: 529-530.

A thorough and erudite bibliographic investigation has established that the earliest effective lectotypification of Caladium Vent. applied that name in the modern sense of Colocasia [indeed, the name Caladium is derived from keladi, a Malay word for Colocasia esculenta (L.) Schott].  This proposal seeks to uphold the prevailing and very long-standing usage of two familiar genus names by designating Caladium bicolor (L.) Vent. as the conserved type of Caladium.  In the exceedingly unlikely event this proposal should fail, then (if we follow the proposal correctly) the name Caladium would replace Colocasia for an Old World genus that includes the cultivated taro (now Colocasia esculenta), and would itself be replaced by Phyllotaenium André (designating a New World genus that includes the familiar ornamental currently called Caladium bicolor).

Ossenbach, C.  2006. Orchids in Costa Rica.  Part I:  The era of Rudolf Schlechter.  Orchids (West Palm Beach) 75: 38-47.

This new series runs parallel to a similar one, dealing with Central American orchid history, recently launched by the same author (see this column in our last issue).  Despite its title, the first installment is mostly devoted to what we could call the Pittier era, when most of the Costa Rican field work that would fuel Schlechter’s orchid descriptions was realized.  In addition to Pittier himself, Biolley, Tonduz, Wercklé, the Brade brothers, and Brenes are all discussed in some detail.  And finally we learn something about Guillermo Acosta (1878–1955), a wealthy businessman!  Sidebars address the significance of type specimens and the cultivation of Ponthieva brenesii Schltr.  Features many excellent (mostly color) illustrations, including a black-and-white photo of Doña Amparo de Zeledón that is different (and more flattering) than the one used in the Manual.

-- . 2006. Orchids in Costa Rica.  Part II:  Charles H. Lankester and Oakes Ames.  Orchids (West Palm Beach) 75: 120-127.

The focus here is squarely on Lankester, an Englishman long resident in Costa Rica, whose collections were a principal basis of the many Costa Rican orchid spp. described by Harvard botanist Ames.  It turns out that Ames was in direct competition with “the Germans” (i.e., Schlechter, working with the collections of Tonduz, Brenes, etc.), and recruited Lankester as a means “to keep abreast” of their efforts.  The collaboration lasted 20 years, and accomplished its goal (as an appended list of new orchid spp. collected by Lankester will attest).  In a story all too familiar to botanists, Lankester died (in 1969) “having spent his last years in a difficult economic condition. ”  But his was a long and rich life, and anyone interested in the details should begin with this article.  The cultivation of myrmecophilous orchids is addressed in a sidebar.  Among the many illustrations are several rare photos from Lankester’s era.

-- . 2006. Orchids in Costa Rica.  Part III:  Rafael Lucas Rodríguez.   Orchids (West Palm Beach) 75: 198-205.

This is an objective consideration of Dr. Rafael Lucas Rodríguez, the most revered and influential Costa Rica biologist of the mid-20th century.  Rodríguez earned his Ph.D. at the University of California (Berkeley) under the late Lincoln Constance, who later characterized Don Rafa as a conflicted figure, “‘torn between a career as a professional biologist...or as an artist.’”  Although he had no aptitude for horticulture, was out of his element in the field, and described just one new plant sp., Rodríguez played critical founding roles in the establishment of the University of Costa Rica Department of Botany, the Revista de Biología Tropical, the Organization for Tropical Studies, the Costa Rican Orchid Society, and the Lankester Botanical Garden.  His dissertation was a masterful study of Myrrhidendron (Apiaceae), but Rodríguez will best be remembered by botanists for his beautiful and scientifically accurate watercolors of Costa Rican orchids, over 1000 in all, most of which have never been published.  Rodríguez’s foray into orchid illustration was instigated by Charles H. Lankester (see previous entry).  Copiously illustrated.

-- . 2006. Orchids in Costa Rica.  Part IV:  the Charles H. Lankester Botanical Garden.  Orchids (West Palm Beach) 75: 284-291.

This contribution (which we gather to be the last in this series) charts the course of the Lankester Botanical Garden (LBG), near Cartago, from a small, private collection (purchased ca. 1973 by the University of Costa Rica for $50,000) to “the most important center for orchid research in Central America and the Caribbean.”  This remarkable transformation has been overseen by just two directors, the late Dora Emilia Mora de Retana (1979–2000) and Jorge Warner (2000–present).  Mora de Retana produced the illustrated series Icones Plantarum Tropicarum (in collaboration with John T. Atwood) and a checklist of Costa Rican orchids (with the late Joaquín B. García Castro), and organized the first orchidology course at the University of Costa Rica (with Robert L. Dressler).   Under Warner’s watch, the garden achieved the status of “experimental station” and launched the highly successful taxonomic journal Lankesteriana, edited by staff-members Franco Pupulin and (later) Carlos O. Morales.  Sidebars by Warner and Diego Bogarín discuss (respectively) orchid conservation in Mesoamerica and orchid collections at LBG.  Features numerous photos, including several of historical interest.

Persson, C. & E. Ljungstrand.  2006. (1716) Proposal to conserve Duroia, nom. cons. (Rubiaceae, Gardenieae) against an additional name, Coupoui.  Taxon 55: 227-238.

The overlooked Coupoui Aubl. (1775) threatens Duroia L. f. (1782), an established name for a neotropical genus of some 20–25 spp., at least one of which occurs in Costa Rica.

Pirie, M. D., S. Kankainen & P. J. M. Maas.  2005. Revision and phylogeny of Cremastosperma (Annonaceae).  Pp. 87–188 in, M. D. Pirie, Cremastosperma (and other evolutionary digressions): molecular phylogenetic, biogeographic, and taxonomic studies in Neotropical Annonaceae.  Ph.D. dissertation, Univ. Utrecht, Netherlands.  256 pp.

This is one of five chapters (and the only one of interest to us) in what we take to be an effectively published thesis (although one of the other chapters has already been published elsewhere, and two have been submitted).  The revision of Cremastosperma is state-of-the-art, with moleculary phylogeny, both dichotomous and synoptical keys, and plenty of maps and illustrations.  None of the 31 accepted spp. is newly validated herein, and two are given only provisional names, “Cremastosperma spec. A” and “spec. B.”  The former designation applies to the only sp. occurring in Costa Rica, which means that we don’t get no satisfaction!  Although we have known of this apparently endemic sp. for more than 20 years, no one has managed to collect flowering material, deemed necessary by the authors (though they acknowledge that, “on the basis of the fruits, Cremastosperma spec. A appears distinct”).  We hope to get to the bottom of this before the Manual Annonaceae treatment goes to press.

Prince, L. M. & W. J. Kress.  2006. Phylogenetic relationships and classification in Marantaceae:  insights from plastid DNA sequence data.  Taxon 55: 281-296.

Calathea, the largest genus in Marantaceae, is paraphyletic, with spp. occurring in two distinct lineages.  Considering only those sampled spp. represented in Costa Rica, Calathea crotalifera S. Watson occupies the smaller clade, while the larger clade includes C. foliosa Rowlee ex Woodson, C. gymnocarpa H. Kenn., C. micans (L. Mathieu) Körn., C. vinosa H. Kenn., and C. warscewiczii (L. Mathieu) Planch. & Linden.  The authors do not suggest possible synapomorphies for these clades and, pending sampling of the type sp. (Calathea discolor G. Mey., = C. lutea Schult.), cannot say which clade should retain the name CalatheaIschnosiphon and Pleiostachya emerge as sister taxa, but the resolution of this study was insufficient to determine whether they should be merged.

Pruski, J. F. & G. Sancho.  2006. Conyza sumatrensis var. leiotheca (Compositae: Astereae), a new combination for a common Neotropical weed.  Novon 16: 96-101.

Despite its name, Conyza sumatrensis (Retz.) E. Walker (Asteraceae) is “seemingly native to the Neotropics,” where it has been much confused with the closely related C. bonariensis (L.) Cronquist and C. canadensis (L.) Cronquist, more familiar names both.  These authors adopt a relatively inclusive concept of C. sumatrensis, at the expense of the latter two spp., involving two vars.:  the autonymic var., now a pantropical weed, and C. s. var. leiotheca (S. F. Blake) Pruski & G. Sancho (comb. nov.), restricted to the Neotropics.  The latter var. resembles and has been confused with Conyza canadensis, whereas the same can be said about the autonymic var. with respect to C. bonariensis.  We know that both vars. of Conyza sumatrensis occur in Costa Rica, because specimens are cited.  We strongly suspect that C. bonariensis and C. canadensis do as well, based on the authors’ statement that “each species now occurs pantropically”; however, we cannot be certain in the absence of voucher citations.  A dichotomous key is presented for the four taxa discussed in this paragraph (all of which have sometimes been included in Erigeron).  A fine composite line drawing (by the second author) depicts capitular and floral features of C. sumatrensis var. leiotheca.

Pupulin, F. [“and collaborators”].  2005. Vanishing beauty:  native Costa Rican orchids.  Vol. 1.  Acianthera–Kegeliella.  Edit. Univ. Costa Rica, San José, Costa Rica.  421 pp.

Having already established himself as a superlative artist, Jardín Botánico Lankester orchidologist Franco Pupulin abundantly demonstrates his equally accomplished photographic skills in this first volume of his heralded magnum opus on Costa Rican orchids [see The Cutting Edge 11(4): 1–2, Oct. 2004].  The introduction consists of an absorbing, 20-page historical resumé of Costa Rican orchidology, co-authored by Pupulin and colleague Carlos Ossenbach, who has published much of this separately (see this column).  The genus entries follow, in alphabetical order, and the volume concludes with a bibliography and index to scientific names.  Each genus entry is anchored by a textual account and features at least one color photo (several, in most cases).  The photos, mostly by Pupulin, are all exquisitely rendered and detailed.  The accounts, written by a wide variety of well chosen and internationally renowned experts, vary in their length, content, and relevance to Costa Rica, depending on the genus and the familiarity of the author(s) with the Costa Rican flora.  The volume embraces all the most recent new generic circumscriptions suggested by molecular research, e.g., Chelyorchis, Cohniella, Cuitlauzina, Cyrtochiloides, Epilyna, Guarianthe, Heterotaxis, and the Pleurothallis segregates advocated by Pridgeon & Chase [see The Cutting Edge 9(1): 9–10, Jan. 2002].  Three new spp. are published, all based on plants flowering in cultivation:  Chondroscaphe yamilethiae Pupulin (as “ yamilethae”), from 1200–1300 m elevation on the Pacific slope of the Cordillera de Talamanca; Dichaea filiarum Pupulin, from 800 m on the Atlantic slope of the Cordillera Central (Monumento Nacional Guayabo); and Galeandra arundinis Garay & G. A. Romero, from the San Carlos region.  All three novelties are illustrated with a composite line drawing, as well as at least one color photo.  A new form is also validated (Comparettia falcata Poepp. & Endl. f. alba Pupulin), as well as a new combination pertaining to a taxon not recorded from Costa Rica.  Some names are used that are still apparently unpublished (e.g., in Dichaea).  At least one sp., Encyclia amanda (Ames) Dressler, is vouchered for the first time (as far as we know) for Costa Rica.  Additionally, a few spp. are newly (at least from our perspective) attributed to Costa Rica, e.g., Bletia edwardsii Ames and Heterotaxis sessilis (Sw.) F. Barros, albeit without voucher citations.

Although this work does not include keys to spp. or even checklists, it will serve as an invaluable adjunct to the identification of Costa Rican orchids (at least to genus) by virtue of its many excellent photos.  Are only major criticism would be that, in some cases, the photos are too many.  For example, Cattleya dowiana Bateman is depicted in eight virtually interchangeable (to our undiscerning eyes) color photos, occupying seven pages; space and considerable expense could have been saved by selecting just one of these.   Also, a few photos portray spp. not known from Costa Rica (e.g., Acineta sella-turcica Rchb. f.), a fact that is not made evident in every case.

Rakotondrainibe, F. & T. Deroin.  2006. Comparative morphology and rhizome anatomy of two new species of Zygophlebia (Grammitidaceae) from Madagascar and notes on the generic circumscription of Zygophlebia and Ceradenia.  Taxon 55: 145-152.

The title gives it away:  the discovery of two new Madagascan spp. of Grammitidaceae blurs the distinction between Ceradenia and Zygophlebia (both of which occur in Costa Rica), suggesting that a redefinition of the latter genus is needed.  This would appear to be one more strike against the reigning generic classification of Grammitidaceae [see The Cutting Edge 11(4): 12, Oct. 2004], and further reason to return to the traditional Grammitis s. l.

Robson, N. K. B.  2006. Studies in the genus Hypericum L. (Clusiaceae).   Section 9. Hypericum sensu lato (part 3):  subsection 1. Hypericum series 2. Senanensia, subsection 2. Erecta and section 9b. Graveolentia.   Syst. Biodivers. 4: 19-98.

This is the ninth installment in the author’s protracted monograph of the major genus Hypericum, and completes his coverage of sect. Hypericum s. l.  None of the spp. treated here reaches Costa Rica; indeed, as far as we are aware, all the Costa Rican Hypericum spp. have already been dealt with in this series.  We cite this paper mainly because it features “an introduction to the genus and a summary of the aims and methods of the project.”

Rojas Alvarado, A. F.  2006. Notas sobre Enterosora Baker (Filicales: Grammitidaceae) en Costa Rica.  Lankesteriana 6: 9-13.

Two entities previously regarded as of hybrid origin are here accepted (albeit somewhat tentatively) as good spp., on the grounds that they are fertile and more abundant locally than their purported parents.  The name Enterosora bishopii A. Rojas sp. nov. is applied to material that has previously gone by the apparently unpublished name “Enterosora ×symmicta L. E. Bishop,” supposed to be a hybrid between E. campbellii Baker subsp. spongiosa (Maxon) L. E. Bishop and E. trifurcata (L.) L. E. Bishop.  Specimens previously considered as hybrids between Enterosora percrassa (Baker) L. E. Bishop and E. trifurcata may now be determined as Enterosora enterosoroides (Christ) A. Rojas comb. nov. (based on Polypodium enterosoroides Christ).  Both erstwhile hybrids are widespread in the Costa Rican cordilleras, principally at elevations of 1200–2000 m, and extend into westernmost Panama (though only Costa Rican specimens of E. enterosoroides are cited).   Both are illustrated with images from herbarium specimens.

Smith, G. F., A. E. van Wyk, M. Luckow & B. Schrire.  2006. Conserving Acacia Mill. with a conserved type.  What happened in Vienna?  Taxon 55: 223-225.

Well, we’ve been following this story, so why stop here?  It was predictable that the rest of the world would be outraged by the official decision to conserve the name Acacia with an Australian type.  These authors (two of which are based in South Africa) provide some of the lurid details of the political machinations that went on behind the scenes in Vienna, culminating in the egregious ruling [see The Cutting Edge 12(4): 1, Oct. 2005].  It turns out, despite all kinds of nefarious subterfuges, the majority of members in the Nomenclatural Section (54.9%) actually voted against the conservation proposal!  The authors sagely pronounce that “exceptions should not be made to the Code unless the case is overwhelming and in this instance it certainly was not”; they suggest that “a fairer outcome would surely have been to let simple priority and normal typification rule in this case.”  We say, impeach the current rulers and install these blokes!  But the authors sign off with an ominous coda:  “Bear in mind that the ICBN does not prescribe what classification system to use, hence the end-users of plant names now have a choice as to whether they want to use the name Acacia in a strict or a wide sense.”  Will reckless and ill-advised decisions by the ruling nomenclatural junta inspire regional boycotts of robust new classifications?  Stay tuned!

Sousa S., M.  2005. Especies nuevas de Lonchocarpus (Millettieae: Leguminosae) para Costa Rica y Panamá; lectotipificación de la sect. Densiflori Benth./New species of Lonchocarpus (Millettieae: Leguminosae) from Costa Rica and Panama; lectotypification of sect. Densiflori Benth.  Revista Mex. Biodivers. 76: 119-127.

Four new Lonchocarpus spp. are described here, of which three occur in Costa Rica.  Two are endemic to Costa Rica and further restricted, as implied by their epithets, to the Monteverde region of the Cordillera de Tilarán.  Curiously, both Lonchocarpus haberi M. Sousa and L. monteviridis M. Sousa have only been collected on the Pacific slope, the former at somewhat lower elevations (600–1200 m) than the latter (1400–1600 m).  Lonchocarpus haberi (the only new sp. described here that does not belong to sect. Densiflori) is most closely related to L. chiangii M. Sousa and L. oliganthus F. J. Herm., while L. monteviridis is nearest to L. luteomaculatus Pittier.  Lonchocarpus latisiliquus M. Sousa sp. nov. occurs in Panama as well as Costa Rica; in the latter country it is known exclusively from the Estación Biológica La Selva and environs, where it has previously been called L. heptaphyllus (Poir.) DC. or L. pentaphyllus (Poir.) Kunth ex DC.  It is indeed most closely related to the latter two spp., which may or may not themselves occur in Costa Rica.  Includes a dichotomous, indented key to the nine spp. of sect. Densiflori recorded from Costa Rica and Panama.  All four new spp. are illustrated with composite line drawings.  Only the title and abstract are bilingual (the rest is in Spanish).

Specht, C. D.  2006. Systematics and evolution of the tropical monocot family Costaceae (Zingiberales):  a multiple dataset approach.  Syst. Bot. 31: 89-106.

This is one of the cladistic studies upon which the new classification of Costaceae discussed under the following entry was predicated.  Since we are only interested in the classification...read on!

-- & D. W. Stevenson.  2006. A new phylogeny-based generic classification of Costaceae (Zingiberales).  Taxon 55: 153-163.

Cladistic analyses (see foregoing entry) of molecular data from both chloroplast and nuclear genomes, in combination with morphological data, have established that the major, pantropical genus Costus is polyphyletic.  On the other hand, the three smaller genera of Costaceae (including Dimerocostus, represented in Costa Rica) are upheld as monophyletic.  This paper, by two of the original investigators, formally implements a new generic classification of Costaceae based on these cladograms (one of which is reproduced).  The authors’ solution is to divide Costus s. l. into four smaller genera, three of which are described as new.  Amazingly, this drastic reconfiguration has minimal consequences for the Costa Rican flora, as all the indigenous spp. treated under the name Costus in the Manual remain in Costus s. str.; the three new genera are either paleotropical (Cheilocostus, Paracostus) or restricted to South America (Chamaecostus).  We only need to digest a single nomenclatural change:  the Asian Costus speciosus (J. Koenig) Sm., naturalized in Costa Rica, becomes Cheilocostus speciosus (J. Koenig) C. Specht.  Dimerocostus remains intact.  Other solutions were obviously possible (according to the cladogram), but this one seems as good as any.  Includes full synonymy and typology (for the family, genera, and all lower names dealt with), family and genus descriptions, a key to genera, and extensive discussions.

Taylor, C. M.  2006. Rubiacearum americanarum magna hama pars XVIII:  new species of Psychotria subg. Psychotria from Central America and western South America.  Novon 16: 142-154.

This time around the big bucket runneth over with new South American spp., nine in all, just one of which barely reaches Costa Rica.  Psychotria convergens C. M. Taylor is centered in Colombia and Ecuador, with a disjunct population at about 2350–2600 m elevation on the Atlantic slope of the Cordillera de Talamanca on both sides of the Costa Rica/Panama border (mainly in the area known as “Valle de Silencio”).  In our area, the new sp. has been confused with Psychotria sylvivaga Standl., from which it differs in its coarser pubescence and longer corollas, internally pubescent in a medial ring (vs. above the middle).  Illustrated with an excellent composite line drawing.

Weigend, M.  2006. Validating subfamily, genus and species names in Loasaceae (Cornales).  Taxon 55: 463-468.

Horror of horrors!  Due to a new ruling implemented at the International Botanical Congress in Vienna last year, many of the author’s new names in Loasaceae that had been widely accepted as effectively published in this 1997 Ph.D. dissertation (Ludwig-Maximilians-Universität, München) are now lost to us.  This ruling requires that, retroactively to 1 January 1953, “an independent non-serial work stated to be a thesis submitted to a university or other institute of education for the purpose of obtaining a degree [is] not effectively published unless it include[s] an explicit statement...or other internal evidence that it was regarded as an effective publication by its author or publisher” (quoting Weigend here, not the Code).  Thus, not only are the names of new genera, spp., etc., proposed in Weigend’s thesis now invalid, but “all species which have since been described in or transferred to the new genera appearing in the thesis require new publication.”  The present paper begins the task of re-validating all the author’s fallen names, addressing only those proposed in the thesis itself.  Among these, the genus names Nasa and Chichicaste and the sp. name Chichicaste grandis (Standl.) Weigend are germane to Costa Rican floristics.  The author promises that “the other names [those proposed after his thesis] will be published elsewhere in the near future.”  We hope this is true, since Loasaceae are due to appear in our next Manual volume!  Two sp. names and one subsp. name (all in Nasa) remain to be validated for Costa Rica.  We expect that this new ruling may impact other families as well.

Zhang, L.-B. & M. P. Simmons.  2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes.  Syst. Bot. 31: 122-137.

There is some vindication here for Manual co-PI Barry Hammel and associates, as the two genera of Lepidobotryaceae (including Ruptiliocarpon) “were unambiguously supported as a monophyletic group” in these analyses.   However, they are also “unambiguously supported as the sister group of the Celastraceae/Parnassiaceae clade,” and thus “properly assigned to Celastrales” (Barry’s team had posited a relationship with Euphorbiaceae or Sapindales).  Also of interest is the status of Perrottetia, traditionally classed in Celastraceae but long recognized as anomalous within that family.  Here, it “was resolved as the sister group of the monotypic Tapiscia (Tapisciaceae),” and therefore “should be transferred out of Celastraceae. ”  By our reckoning, the closest relative of Perrottetia in Costa Rica would thus be Huertea, traditionally a member of Staphyleaceae, but assigned to Tapisciaceae by the Angiosperm Phylogeny Group [see The Cutting Edge 10(3): 5–6, Jul. 2003].


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