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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XIV, Number 4, October 2007

News and Notes | Leaps and Bounds | Germane Literature | Season's Pick | Annotate your copy

Aiello, A., M. Correa & C. Galdames.  2007. Remarkable new plant record for Panama.  STRI News 2007 (August 31): 4.

The plant is indeed remarkable, none other than “our own” Osa pulchra (D. R. Simpson) Aiello (Rubiaceae), which revealed itself to startled botanists during a recent collecting expedition to a remote region in the lowlands of Prov. Bocas del Toro.  However, the record itself is not surprising, as we have known for some time that this sp. occurs in the Atlantic lowlands of southeasternmost Costa Rica, near Cahuita and Fila Carbón (Valverde & Zúñiga 1382, 1393; USJ), not far from the Panamanian locality.  What this does mean is that Costa Rica loses one of its few endemic plant genera.  Read the bad news, and feast your eyes on a stunning photo, at:


Alvarado-Cárdenas, L. O. & H. Ochoterena.  2007. A phylogenetic analysis of the CascabelaThevetia species complex (Plumerieae, Apocynaceae) based on morphology.  Ann. Missouri Bot. Gard. 94: 298–323.

This study confirms the monophyly of the complex named in the title, which comprises a total of 14 spp.  The two genera are also monophyletic, or at least can easily be made so with a single new combination (here validated).  The sister-group relationship of Cascabela and Thevetia complicates the basic issue of whether to accept one genus or two.  These authors opt to accept both, “because there are two important and easy-to-diagnose characters supporting each clade.”  They ignore the much more important criterion of nomenclature stability, which would perhaps favor the opposite decision:  with the exception of Flora de Nicaragua, no major regional (Mesoamerican) floristic work has accepted the name Cascabela.

Appendix 4 of this paper comprises a “taxonomic synopsis” of Cascabela and Thevetia, including keys to the genera and the spp. of each, formal generic and sp. descriptions, full synonymy and typology at all levels, and, for each sp., a distribution summary, discussion, and representative specimen citations.  According to this classification, Costa Rica has just one native sp. in each genus:  Cascabela ovata (Cav.) Lippold and Thevetia ahouai (L.) A. DC.; a third sp.,Cascabela thevetia (L.) Lippold [AKA Thevetia peruviana (Pers.) K. Schum.], is known from Costa Rica only in cultivation (although these authors give no such indication).

Anderson, C.  2007. Revision of Galphimia (Malpighiaceae).  Contr. Univ. Michigan Herb. 25: 1–82.

Galphimia is a genus of 26 spp., ranging from Texas to northern Nicaragua and (disjunctly) South America.  It therefore does not occur naturally in Costa Rica, although one sp. is commonly cultivated as an ornamental shrub and is briefly mentioned in the family discussion of the in-press Manual treatment of Malpighiaceae by William R. Anderson (MICH).  The name of that sp., Galphimia gracilis Bartl., remains unchanged, and there ends our immediate interest in this revision, which we include here mainly for reference purposes.  Readers with a need to investigate further will find a superb, fully realized treatment, much in the style of Systematic Botany Monographs, of which the author is the long-time editor.

Anderson, W. R.  2007. Lectotypification of names of Malpighiaceae—I.  Contr. Univ. Michigan Herb. 25: 83–93.

Dozens of names are lectotypified here, several of which (e.g., Bunchosia costaricensis Rose and Mascagnia vacciniifolia Nied.) apply to spp. occurring in Costa Rica.  As far as we can tell, no changes in usage result from these typifications, in keeping with the author’s characterization of the paper as “a first step toward the goal of stabilizing the nomenclature of the Malpighiaceae.”

-- & C. C. Davis.  2007. Generic adjustments in neotropical Malpighiaceae.  Contr. Univ. Michigan Herb. 25: 137–166.

Several “adjustments” are implemented in this paper, but just one concerns us:  the further dismemberment of Banisteriopsis.  Previously, these same authors removed one of the three subgenera of Banisteriopsis sensu Gates (Fl. Neotrop. Monogr. 30: 1–237.   1982) to the genus Diplopterys [see The Cutting Edge 13(4): 2–3, Oct. 2006], on the basis of their own unpublished molecular data.  That change was accommodated in the first author’s Manual Malpighiaceae treatment, currently in-press, although the impact for Costa Rica was minimal.  The latest “adjustments” have more extensive consequences for Costa Rican floristics, but have come to fruition too late for full implementation in the Manual (although a brief reference was inserted at the last moment).  Based on (apparently) the same body of molecular data (still unpublished), the authors here split off one of the two remaining subgenera of Banisteriopsis to a new genus, Bronwenia W. R. Anderson & C. Davis (honoring Banisteriopsis/Diplopterys monographer and former Anderson student Bronwen Gates).  Somewhat confusingly, Bronwenia corresponds to what was, according to Gates, the autonymic subgenus of Banisteriopsis; it turns out that Gates had overlooked an earlier typification, according to which her subgen. Hemiramma (Griseb.) B. Gates must become subgen. Banisteriopsis.  Actually, this is a fortunate development, as it means that the majority of the spp. involved remain in Banisteriopsis; in Costa Rica, these are Banisteriopsis caapi (Spruce ex Griseb.) C. V. Morton (cultivated only), B. elegans (Triana & Planch.) Sandwith, B. lyrata B. Gates, and B. muricata (Cav.) Cuatrec.  The remaining two Costa Rican representatives of Banisteriopsis sensu Gates, B. acapulcensis (Rose) Small and B. cornifolia (Kunth) C. B. Rob. var. maracaybensis (A. Juss.) W. R. Anderson, are transferred to Bronwenia (with eight other spp.), with all new combinations attributed to “W. R. Anderson & C. Davis.”  The authors express doubt as to the distinction between Bronwenia cornifolia and the Brazilian B. ferruginea (Cav.) W. R. Anderson & C. Davis, and suggest that “eventually they may have to be combined under” the latter name.  Includes a formal description and brief diagnosis of Bronwenia, a dichotomous key to spp. (adapted from Gates), full synonymy and typology for each sp., and a fine composite line drawing of B. acapulcensis.

Bogarín, D. & F. Pupulin.  2007. Las orquídeas del Parque Nacional Barra Honda, Guanacaste, Costa Rica.  Lankesteriana 7: 446–449.

The authors list (without explicitly citing vouchers) 26 spp. in 23 genera, rather surprising totals for one of the most arid sites in Costa Rica.  Seven spp. are terrestrial, including the adventive, African Oeceoclades maculata (Lindl.) Lindl. and two spp. (in Palmorchis and Sarcoglottis) undetermined beyond the level of genus.

Chung, K.-F.  2007.  Inclusion of the South Pacific alpine genus Oreomyrrhis (Apiaceae) in Chaerophyllum based on nuclear and chloroplast DNA sequences.  Syst. Bot. 32: 671–681.

The results of a previous paper helmed by this same author suggested that the north-temperate Chaerophyllum, a genus of ca. 30 spp., was paraphyletic with respect to the circum-Pacific Oreomyrrhis, with 26 spp. [see The Cutting Edge 13(2): 3, Apr. 2006].  Nomenclatural changes were postponed pending further investigation employing sequence data from chloroplast DNA (the original study having relied upon nuclear DNA data).  Further investigation has now been accomplished, and the initial hypotheses upheld:  Oreomyrrhis is monophyletic, but nested within Chaerophyllum, specifically, in a sister-group relationship with the two North American spp. of the latter genus.  The classificatory option of fragmenting Chaerophyllum to retain Oreomyrrhis is rejected, because a minimum of three additional genera would be required (leaving Chaerophyllum s. str. monospecific), based on “clades that lack apparent morphological synapomorphies.”  Thus Oreomyrrhis is formally sunk into Chaerophyllum, which thereby balloons to ca. 60 spp.  Recognition of Oreomyrrhis even at sectional rank is eschewed, as that would “again render sect. Chaerophyllum paraphyletic”; instead, the informal designation “Oreomyrrhis clade” is recommended.  All necessary new combinations and nomina nova are validated, in the name of the author, in a comprehensive enumeration of the spp. composing the “Oreomyrrhis clade” which also includes homotypic synonymy and typology.  The sole sp. occurring in Costa Rica is henceforth Chaerophyllum andicola (Kunth) K. F. Chung.

Cuevas Figueroa, X. M.  2005. A revision of the genus Amphipterygium (Julianiaceae).  Ibugana 13(1): 27–47.

Just one sp. of Amphipterygium reaches Costa Rica, where it is restricted to several islands off the Península de Santa Elena, in nothern Prov. Guanacaste, and scattered localities on the nearby mainland.  Though rare and little-known from a national perspective, this sp. (comprising small trees) may be a dominant element where it does occur, e.g., on several of the Islas Murcíelago.  We were content for many years with the apt name Amphipterygium molle (Hemsl.) Standl., first applied to Costa Rican material in Holdridge & Poveda’s (1975) classic Árboles de Costa Rica Vol. 1, until Carmen Ulloa Ulloa (MO) substituted A. adstringens (Schltdl.) Standl. in her Flora de Nicaragua (2001) treatment of Julianiaceae.  Still more recently, however, we have been perplexed to note that Anacardiaceae specialist John D. Mitchell (NY) has identified Costa Rica specimens as A. molle.  For the past several years, establishing the correct name for this plant has been a significant concern to botanists (ourselves included) and conservation workers in the Guanacaste region, where this is a marquee sp.  Finally, we have in our hands a generic revision (okay, so apparently it’s been out for awhile) to arbitrate this matter authoritatively.  And the decision is (envelope, please):  none of the above!  According to this Mexican worker, Amphipterygium comprises four spp., of which three (including A. adstringens and A. molle) are endemic to Mexico; the fourth sp., ranging from Oaxaca to Costa Rica, is branded Amphipterygium simplicifolium (Standl.) Cuevas-Figueroa comb. nov. (based on A. adstringens subsp. simplicifolium Standl.), which name we will now happily embrace.

This is a standard revision, with genus and sp. descriptions, a dichotomous, non-indented key to spp., a distribution map, and, for each sp., synonymy and typology, summaries of phenology and distribution, a list of vernacular names, a diagnostic statement, and comprehensive specimen citations.  Each sp. is illustrated with separate line drawings of male and female plants, plus at least two black-and-white photos depicting living and/or herbarium material.  The brief introduction deals mainly with taxonomic history, but a section at the end investigates phylogeny on the basis of nuclear DNA analysis (A. simplicifolium is sister to the other three spp.).

Incidental matters:  Don’t look for “Julianiaceae” in our forthcoming Manual volume, where that family would fall alphabetically; following the Angiosperm Phylogeny Group, and other reliable sources, we will include Amphipterygium in Anacardiaceae, slated for Manual Vol. 4 (our final volume, chronologically speaking); good thing, too, or we’d have used the wrong sp. name!  Ibugana is the bulletin of “IBUG,” i.e., the Instituto de Botánica de la Universidad de Guadalajara, and supersedes Bol. Inst. Bot. Univ. Guadalajara.  This article may be accessed in PDF format at the following source:


De-Nova, J. A., V. Sosa & V. W. Steinmann.  2007. A synopsis of Adelia (Euphorbiaceae s. s.).  Syst. Bot. 32: 583–595.

Adelia comprises nine spp. (according to this revision), ranging nearly throughout the Neotropics.  There is nothing new here for us, as only A. triloba (Müll. Arg.) Hemsl. is recorded from Costa Rica.  The last-mentioned sp. ranges southward to Ecuador, and is the only member of the genus occurring in this region.  Intriguingly, the generally more northern Adelia barbinervis Cham. & Schltdl. extends to quite near the Costa Rican border in south-central Nicaragua, and should be sought on the Llanuras de Los Guatusos and de San Carlos or along the Río San Juan.  In terms of its presentation, this is more of a revision than a synopsis, as it includes formal genus and spp. descriptions, along with a dichotomous, indented key to spp., full synonymy and typology, summaries of phenology and distribution, and representative specimen citations.  Illustrated with a composite line drawing of one sp. (neither of the aforementioned), and another showing morphological details of various spp. (but mainly A. barbinervis).  The introductory discussion is very brief.

Doust, A. N., A. M. Penly, S. W. L. Jacobs & E. A. Kellogg.  2007. Congruence, conflict, and polyploidization shown by nuclear and chloroplast markers in the monophyletic “bristle clade” (Paniceae, Panicoideae, Poaceae).  Syst. Bot. 32: 531–544.

The so-called “bristle clade,” already supported as monophyletic by previous studies, comprises several genera of grasses with inflorescences bearing “stiff sterile branches” (i.e., bristles), believed to be uniquely derived.  The important neotropical members of this group are Cenchrus, Ixophorus, Paspalidium, Pennisetum, Setaria, and Stenotaphrum.  The molecular phylogenies generated for this study “show that morphological similarity...generally does not indicate phylogenetic relationship” within this group—a sobering assessment indeed for conventional taxonomists!  Our comprehension of this paper is much facilitated by an explicitly written section headed “Implications for Classification” (other authors of such papers, take heed!).  Cenchrus is supported as monophyletic, but Pennisetum cannot be rejected as monophyletic; thus, the merger of these two genera can be neither supported nor rejected—in contrast to some previous molecular studies, in which Pennisetum has been portrayed as paraphyletic with respect to CenchrusPaspalidium plus Setaria does not comprise a monophyletic group, suggesting that those two genera should not be combined.  Indeed, Setaria itself is polyphyletic, according to the tree generated by the nuclear marker, with three well-supported major clades:  one including the generic type, S. viridis (L.) P. Beauv. (a rare adventive in Costa Rica); a morphologically very similar group including such spp. (in Costa Rica) as S. parviflora (Poir.) Kerguélen and S. sphacelata (Schumach.) Stapf & C. E. Hubb. ex M. B. Moss; and a group corresponding to Setaria sect. Ptychophyllum (A. Braun) Stapf, comprising morphologically distinctive plants with plicate leaves and open inflorescences [in Costa Rica, S. paniculifera (Steud.) E. Fourn. ex Hemsl. and S. poiretiana (Schult.) Kunth].  Nonetheless, the tree generated by the chloroplast marker “cannot reject monophyly of Setaria,” so the authors refrain from fragmenting the genus.  The morphologically distinctive sect. Ptychophyllum might justifiably be segregated by itself; however, the authors fear “that doing this might lead to a paraphyletic residue of Setaria species.”

Duno de Stefano, R.  2007.  Tratamiento taxonómico del género Dendrobangia Rusby (Cardiopteridaceae o Icacinaceae).  Candollea 62: 91–103.

We would love to have been able to cite this revision in our in-press Manual volume (where Dendrobangia is treated traditionally in Icacinaceae), but it hit the shelves just a few weeks too late.  Nonetheless, the changes for us are minimal, with Dendrobangia boliviana Rusby still the accepted name for the sole Costa Rican sp.  However, the relegation of the Brazilian Dendrobangia tenuis Ducke to synonymy under D. boliviana mandates a reduction in the generic sp.-total from three to two, and the citation of a collection of D. boliviana from Trinidad requires the addition of that country to the geographic range of both the sp. and genus.  Features lengthy genus and sp. descriptions, a key to spp., full synonymy and typology, distribution summaries, comprehensive specimen citations, and distribution maps.   Both sp. are depicted in photographic images of herbarium (type) material, and D. boliviana in a composite line drawing.

Endress, M. E. & B. F. Hansen.  2007. Pinochia, a new genus of Apocynaceae, Apocynoideae from the Greater Antilles, Mexico and Central America.  Edinburgh J. Bot. 64: 269–274.

Pursuant to a molecular study showing Forsteronia to be polyphyletic (see under Livshultz et al., this column), four spp. long appreciated as morphologically anomalous are segregated into a new genus, Pinochia M. E. Endress & B. F. Hansen.  Just one of these spp. occurs in Costa Rica, that being the recently described Forsteronia monteverdensis J. F. Morales [see The Cutting Edge 3(1): 10–11, Jan. 1996], now Pinochia monteverdensis (J. F. Morales) M. E. Endress & B. F. Hansen.  Features a detailed genus description (but no sp. descriptions), dichotomous (but non-indented) keys to spp. and (in one case) subspp., and full synonymy and typology.  Pinochia monteverdensis is (alone) depicted in a composite line drawing.  For the record, Pinochia differs from Forsteronia in its leaf-blades lacking colleters at the base, anthers with acuminate basal appendages, more numerous ovules, ovoid style-heads with a small annulus at the base, and seeds without longitudinal ribs.  Oh, and forget all about marionettes and exaggerated beaks; Pinochia is an anagram based on the surname of late Apocynaceae monographer Marcel Pichon (1921–1954).

Eriksson, R.  2007. A synopsis of Basellaceae.  Kew Bull. 62: 297–320.

This brief but useful summary recognizes four genera with a total of 19 spp., plus a few subspp.   Features synonymy and typology at all levels, a detailed family description, successively less detailed sp. and subspp. descriptions, a combined, dichotomous (though non-indented) key to genera, spp., and subspp., and, for each sp. and/or subsp., a paragraph on nomenclature, illustration references, a distribution summary, selected exsciccatae citations, and note(s).  The introductory part addresses taxonomic history, morphological details (with supplementary illustrations), reproduction, phylogeny, and distribution (including a map).  There is an index to names at the end.  Although it is not obvious from this source, Basellaceae is poorly represented in Costa Rica, with just three taxa recorded.  The ruderal and probably introduced Anredera cordifolia (Ten.) Steenis is fairly frequent in the Valle Central.   The mysterious Anredera ramosa (Moq.) Eliasson subsp. ramosa and A. vesicaria (Lam.) C. F. Gaertn. are both known from a few historical collections, but we’ve never encountered them.  It is also possible that the so-called “Malabar spinach,” Basella alba L., may be cultivated somewhere in Costa Rica on a minor scale.

Erkens, R. H. J., L. W. Chatrou, J. Koek-Noorman, J. W. Maas & P. J. M. Maas.  2007. Classification of a large and widespread genus of Neotropical trees, Guatteria (Annonaceae) and its three satellite genera Guatteriella, Guatteriopsis and Heteropetalum.  Taxon 56: 757–774.

Analysis of molecular data reveals that the three satellite genera mentioned in the title (none of which occurs in Costa Rica) are all nested within the large genus Guatteria, which is otherwise strongly supported as monophyletic.  It is suggested that all three satellites “should be merged with Guatteria,” but the indicated new combinations are not validated.

--, P. J. M. Maas & G. E. Schatz.  2007. Preliminary Flora Mesoamericana treatment of Guatteria.  Pp. 179–217 in, R. H. J. Erkens, From morphological nightmare to molecular conundrum.  Phylogenetic, evolutionary and taxonomic studies on Guatteria (Annonaceae).  Ph.D. thesis, Univ. Utrecht, Netherlands.  275 pp.

Because Guatteria is the largest and taxonomically most difficult genus of Annonaceae in Mesoamerica, this preliminary treatment provides valuable insights and promises to be extremely useful in the immediate future, even though changes are inevitable prior to its “official” publication.  Twenty-eight spp. of Guatteria are formally recognized from the Mesoamerican region, plus four “Insufficiently Known Species,” treated separately at the end of the (otherwise alphabetical) sequence.  Of these 32 spp., 17 are attributed to Costa Rica, including six country endemics:  Guatteria pudica N. Zamora & Maas, G. reinaldoi Erkens & Maas, G. rostrata Erkens & Maas, G. tenera R. E. Fr. (one of the “insufficiently known” spp.), G. verrucosa R. E. Fr., and “G. sp. 1” (one of three provisionally named spp.).  The last-mentioned entity is based on two collections, made on the same trip, from a remote (and presently inaccessible) site at ca. 1300 m elevation on the Atlantic slope of the Cordillera de Talamanca near the Panamanian border.  The otherwise Panamanian Guatteria alata Maas & Setten is reported from Costa Rica on the sole basis of a specimen collected by Manual co-PI Nelson Zamora (in the Barra del Colorado region) that was destroyed in a herbarium fire.  Features a brief genus description, dichotomous (and indented!) key to spp., adequately detailed sp. descriptions, brief distributional data in the standard Flora mesoamericana format, diagnostic discussions, black-and-white photos of herbarium specimens, and an index to exsiccatae.   Although this qualifies as an effective publication, no taxonomic novelties are validated here.  The bulk of this thesis comprises a collaborative, multidisciplinary assault on the phylogenetics and taxonomy of Guatteria; several of the other 10 chapters have been submitted for formal publication, and some have already seen the light of day [see foregoing entry, as well as The Cutting Edge 13(4): 3, Oct. 2006].

González, F., J. Betancur, O. Maurin, J. V. Freudenstein & M. W. Chase.  2007. Metteniusaceae, an early-diverging family in the lamiid clade.  Taxon 56: 795–800.

Although the genus Metteniusa has generally been included in Icacinaceae, it has been widely acknowledged as anomalous in that (or any other) family.  Until now, it has not been possible to bring molecular data to bear on this issue due to lack of suitable material, a problem here overcome through collaboration with Colombian biologists.  The resulting molecular analyses, involving three genes, suggest that Metteniusa occupies an isolated position near the monogeneric Oncothecaceae, endemic to New Caledonia.  This evidence is interpreted as supporting separate familial status for Metteniusaceae.  Unfortunately, all of this arrives a day late and a dollar short for us, as Icacinaceae (including Metteniusa!) are in-press for the Manual.

Goyder, D., A. Nicholas & S. Liede-Schumann.  2007. Phylogenetic relationships in subtribe Asclepiadinae (Apocynaceae: Asclepiadoideae).  Ann. Missouri Bot. Gard. 94: 423–434.

These molecular analyses suggest that the familiar genus name Asclepias must either be restricted to spp. of the New World (whence the confusingly named type sp., A. syriaca L.) or applied to the entire subtribe.  This conclusion obviously does not affect native spp. in Costa Rica, only Old World exotics, such as the cultivated (and escaped) Gomphocarpus physocarpus E. Mey. [= Asclepias physocarpa (E. Mey.) Schltr.].

Grose, S. O. & R. G. Olmstead.  2007. Evolution of a charismatic Neotropical clade:  molecular phylogeny of Tabebuia s. l., Crescentieae, and allied genera (Bignoniaceae).  Syst. Bot. 32: 650–659.

Sequence analysis of two chloroplast genetic regions “supports the monophyly of the Neotropical taxa [of Bignoniaceae] with palmately compound leaves.”  This group includes (to mention only those taxa occurring in Costa Rica) the neotropical genera Godmania (oligotypic) and Tabebuia (ca. 100 spp.), as well as tribe Crescentieae.  The last-mentioned taxon, also exclusively neotropical, comprises three smallish to medium-sized genera, Amphitecna, Crescentia, and Parmentiera, all represented in Costa Rica.  The results of this study suggest that unifoliolate leaves have evolved “independently multiple times” within the overall group, e.g., in Amphitecna and some spp. of Tabebuia.  More importantly, the entire tribe Crescentieae (along with two small Antillean genera) is nested within Tabebuia, which is thereby confirmed as paraphyletic.  The disposition of Godmania (and two other small genera not occurring in Costa Rica) relative to Tabebuia could not be resolved.

-- & ---. 2007. Taxonomic revisions in the polyphyletic genus Tabebuia s. l. (Bignoniaceae).  Syst. Bot. 32: 660–670.

In which the more definitive results of the foregoing paper are consecrated by the implementation of real taxonomic changes, based on the cladograms presented therein.  The paraphyly of Tabebuia, the basic problem to be addressed, might be resolved through either lumping or splitting.  The lumping option, not even alluded to here, would require the merger of Amphitecna, Crescentia, Parmentiera, Tabebuia, and at least a few smaller genera (with Crescentia by far the oldest name involved).  Instead, the authors elect to split Tabebuia, in accordance with three clades (two occurring naturally in Costa Rica) identified in their molecular analyses.  The name Tabebuia will remain attached to a the largest of these clades, that corresponding to Gentry’s (1992; Fl. Neotrop. Monogr. 25: 1–335) informal groups 2 and 6–10, characterized by “an indumentum of lepidote scales and a spathaceous, sometimes irregularly bi- or trilabiate calyx,” as well as white to red flowers.  Just two Costa Rican spp. belong to this group (and will thus remain in Tabebuia):  Tabebuia palustris Hemsl. (an anomalous, seldom-seen mangrove sp.) and T. rosea (Bertol.) DC.  A second clade, including Gentry’s informal groups 3–5, is characterized by “an indumentum of simple to stellate or dendroid hairs” (usually also with some lepidote scales) and a “campanulate to cupular and 5-dentate” calyx.  The flowers of this group, which entrains the generic name Handroanthus Mattos, are generally yellow.  The remaining Costa Rican spp. of traditional Tabebuia belong to this group:  T. chrysantha (Jacq.) G. Nicholson, T. guayacan (Seem.) Hemsl., T. impetiginosa (Mart. ex DC.) Standl. (with magenta flowers), and T. ochracea (Cham.) Standl.  The third clade, with just two spp., is “distinguished by its unique calyx...spathaceous in shape, and of the same color and texture as the corolla.”  No member of this clade occurs naturally in Costa Rica, but one sp., Tabebuia donnell-smithii Rose [syn. Cybistax donnell-smithii (Rose) Seibert], has been cultivated there as an ornamental.  The operative genus name for this last clade is Roseodendron Miranda.

In a separate taxonomic section, synonymy, typology, and formal (English) descriptions are provided for Handroanthus, Roseodendron, and Tabebuia s. str., and all accepted spp. and infraspecific taxa are enumerated for each genus; synonymy is indicated for taxa of Handroanthus and Roseodendron, and all necessary new combinations (occurring only in Handroanthus) are validated in the name of S. O. Grose.  A dichotomous, indented key separates all 12 genera comprising the “Tabebuia Alliance” (i.e., “the Neotropical taxa with palmately compound leaves” of the previous paper).

N.B.:  We are quite certain that Handroanthus chrysanthus (Jacq.) S. O. Grose subsp. pluvicolus (A. H. Gentry) S. O. Grose must be corrected to pluvicola, and we are suspicious of H. ochraceus (Cham.) Mattos subsp. neochrysanthus (A. H. Gentry) S. O. Grose; is Gentry’s neochrysantha—clearly referencing Tabebuia chrysantha—a noun in apposition?  Opinions are welcome!

Horn, S. P. & L. M. Kennedy.  2006. Pollen evidence of the prehistoric presence of cattail (Typha: Typhaceae) in Palo Verde National Park, Costa Rica.  Brenesia 66: 85–87.

Despite its reputation as a recent “invader” in the marshes of the lower Río Tempisque basin, Typha domingensis Pers. was already present there between 2850–2340 BC, according to pollen analysis of sediments so dated.  These results “confirm that the plant is native” in the area

Jobson, R. W. & M. Luckow.  2007. Phylogenetic study of the genus Piptadenia (Mimosoideae: Leguminosae) using plastid trnL-F and trnK/matK sequence data.   Syst. Bot. 32: 569–575.

To make a long story short, Piptadenia, most recently regarded as comprising some 27 spp., is resolved as polyphyletic and “is best divided into three genera.”  Piptadenia flava (Spreng. ex DC.) Benth., the sole Costa Rican representative, is one of 23 spp. in the clade putatively harboring the generic type (which itself was not sampled), and thus will retain its present binomial.  We need not concern ourselves with the residual segregates.

Kirkpatrick, R. E. B.  2007. Investigating the monophyly of Pellaea (Pteridaceae) in the context of a phylogenetic analysis of cheilanthoid ferns.  Syst. Bot. 32: 504–518.

DNA sequence analyses of several chloroplast genes show Pellaea to be polyphyletic.  On the surface, this would figure to be of no concern to us, as both Costa Rican spp. of Pellaea, P. ovata (Desv.) Weath. and P. ternifolia (Cav.) Link—indeed, all Mesomerican spp. of the genus—belong to the autonymic section.  However, Pellaea sect. Pellaea is itself polyphyletic, including the Old World genera Paraceterach and Paragymnopteris, as well as the New World genus Astrolepis.  Weighing the possible classificatory ramifications of her results, the author tentatively considers both the splitting and lumping alternatives.  The former, according to our analysis, would result in no name changes for any spp. occurring in Costa Rica; however, lumping the three aforementioned genera into Pellaea would necessitate a name-change for the only Costa Rican representative of Astrolepis, A. sinuata (Lag. ex Sw.) D. M. Benham & Windham.

Klitgaard, B. B.  2007. Three new species in Salvia subgenus Calosphace (Lamiaceae) from Mesoamerica.  Novon 17: 206–211.

Somehow we overlooked this article, in our own in-house journal, despite having reviewed this issue last quarter.  This was a dangerous oversight, as one of the new spp. entails critical implications for our upcoming Manual volume:  Salvia colonica Standl. & L. O. Williams ex Klitg. now applies to most of the material that has recently been identified (e.g., in Flora de Nicaragua and the in-press Manual treatment) as S. comayaguana Standl., here considered a Honduran endemic.  By rare good fortune, the substitution involved no changes in alphabetical order, so we were able to implement it at the (very!) last possible moment.

Livshultz, T., D. J. Middleton, M. E. Endress & J. K. Williams.  2007. Phylogeny of Apocynoideae and the APSA clade (Apocynaceae s. l.).  Ann. Missouri Bot. Gard. 94: 324–359.

The “APSA clade” comprises four subfamilies of Apocynaceae (including Asclepiadaceae), and this paper is mainly devoted to subfamilial and tribal classification, as interpreted from analyses of both molecular and morphological data.  Some results having potential impact at generic rank are:  Forsteronia and Prestonia are both resolved as polyphyletic, while both Allotoonia and Fernaldia are nested within Echites.  The Forsteronia situation has already been addressed (see under Endress & Hansen, this column).  That of Prestonia is more complicated, and additional sampling is indicated; the Costa Rican spp. may be relatively safe, as all three that were sampled (out of seven total) clustered in the same clade with the generic type (P. tomentosa R. Br.).  The authors consider that their results support the “inclusion of Allotoonia in Echites s. l. and the expansion of the circumscription of Echites to include Fernaldia”; the indicated changes could be easily implemented, as all three Costa Rican spp. of the recently segregated Allotoonia [see The Cutting Edge 12(1): 10–11, Jan. 2005] already have names in Echites, as does the more common of the two Costa Rican Fernaldia spp.

Löhne, C., T. Borsch & J. H. Wiersema.  2007. Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers.  Bot. J. Linn. Soc. 154: 141–163.

These analyses, involving nine different regions of the chloroplast genome and 24 accessions (plus five outgroup taxa), confirm the monophyly of sister-taxa Cabombaceae and (more or less) Nymphaeaceae.   However, the pantropical genus Nymphaea emerges as paraphyletic, with the Asian Euryale, the Australian Ondinea, and the South American Victoria nested therein.   These authors stop short of recommending formal changes in classification; however, one may speculate that, if the splitting option were to be imposed on Nymphaea based on the cladograms here presented, at least three separate genera would result.  Such a change would inevitably have consequences for the Costa Rican contingent of 12 Nymphaea spp., several of which belong to subgenera other than the autonymic one.

Martin, C. V., G. Cremers & R. Goldenberg.  2007. Lectotypification of Miconia, Octomeris, and Staphidium (Melastomataceae) names described by Naudin.  Novon 17: 362–368.

As far as we can tell, these are all perfunctory lectotypifications that do not affect usage.  The only names dealt with that appear relevant to Costa Rica are Miconia impetiolaris (Sw.) G. Don var. pandurifolia Naudin and M. obovalis Naudin [the latter a synonym of M. splendens (Sw.) Griseb.].

Miller, J. S.  2007. New Boraginales from tropical America 5:  new names and typifications for Neotropical species of Cordia and Varronia.  Novon 17: 372–375.

In our review of this author’s (with M. Gottschling) recent paper resurrecting Varronia (Boraginaceae s. l., or Cordiaceae) from Cordia, we noted [see The Cutting Edge 14(2): 8, Apr. 2007] that no combinations had been published for two spp. of the group occurring in Costa Rica.   Turns out we jumped the gun a bit:  those combinations must already have been in the works, as both—Varronia guanacastensis (Standl.) J. S. Mill. and V. linnaei (Stearn) J. S. Mill.—are here validated.  None of the other innovations in this paper appears relevant to Costa Rican floristics.

Moore, G.  2007. The handling of the proposal to conserve the name Acacia at the 17th International Botanical Congress—an attempt at minority rule.  Bothalia 37: 109–118.

This is another lucid and detailed recapitulation of the now notorious machinations in Vienna that culminated in the conservation of the name Acacia with a conserved, Australian type [see The Cutting Edge 13(4): 8, Oct. 2006].  This author suggests a challenge to the printed version of the Vienna Code when it goes up for ratification at the next International Botanical Congress (Melbourne, 2011), and offers several “Recommendations for future Nomenclature Sections.”  We suspect that Melbourne may be an unfavorable venue for the proposed challenge!

Morales, J. F.  2006. Panopsis acostana (Proteaceae), una nueva especie del Pacífico sur de Costa Rica.  Brenesia 66: 69–73.

The name Panopsis acostana J. F. Morales is validated for a sp., endemic to Costa Rica at 0–800 m elevation on the southern Pacific slope, that had been previously identified as P. mucronata Cuatrec. [see The Cutting Edge 8(4): 3–4, Oct. 2001].  The Costa Rican material differs from P. mucronata (based on a Colombian type) in its narrower petioles, usually much narrower leaf-blades with fewer secondary veins, and (especially) in its smaller tepals.  Despite its epithet (based on the Cantón de Acosta, in Prov. San José), the new sp. has been collected most frequently on the Península de Osa.  The author discusses certain sterile collections, from similar habitats in the Cordilleras de Guanacaste and de Tilarán, that might be assigned to Panopsis acostana, pending the availability of fertile material.  Illustrated with a photographic image of the holotype specimen.

Muñoz, M. & S. H. Kirby.  2007. An orchid inventory and conservation project at Bosque de Paz Biological Reserve, upper Río Toro Valley, Alajuela, Costa Rica.  Lankesteriana 7: 60–65.

When a preliminary orchid list was published for this same site, accounting for just 55 spp., we predicted that “at least five times as many” would eventually turn up [see The Cutting Edge 10(3): 6–7, Jul. 2003].  Well, they aren’t there yet, but are now able to enumerate (in quasi-vouchered fashion) a total of 163 orchid spp. in 47 genera.  Another four or five years should do the trick!  The most sp.-rich genera in the reserve are Epidendrum (with 24 spp.), Pleurothallis s. l. (23 spp.), and Maxillaria (22 spp.).

-- & J. Warner.  2007. Distribución de poblaciones silvestres y descripción del hábitat de Phragmipedium en Costa Rica.  Lankesteriana 7: 66–70.

The authors set out to provide distributional data for both Costa Rican representatives of the persecuted orchid genus Phragmipedium, based on herbarium and field studies.  However, they were unable to locate any populations of the taxon they refer to (following the Manual) as “Phragmipedium humboldtii (Warsz. ex Rchb. f.) J. T. Atwood & Dressler,” now correctly (if the dust has finally settled) P. warscewiczii (Rchb. f.) Christenson subsp. warscewiczii [see The Cutting Edge 13(3): 4–5, Jul. 2006].  Thus, what we end up with is a (mostly) vouchered, tabularized account of the known Costa Rican localities of P. longifolium (Warsz. & Rchb. f.) Rolfe, more detailed than the Manual summary, supplemented by a rather crude distribution map (and it would be foolhardy to provide a more accurate one!).

Ortiz, R. del C., E. A. Kellogg & H. van der Werff.  2007. Molecular phylogeny of the moonseed family (Menispermaceae):  implications for morphological diversification.  Amer. J. Bot. 94: 1425–1438.

This paper is mainly concerned with suprageneric classification, which is outside our interest.   At the level of genera, however, we glean a few intriguing details, e.g.:  Anomospermum is polyphyletic, while Odontocarya is paraphyletic with respect to the monospecific South American Borismene.  The possible classificatory repercussions of these situations are not addressed; however, based on our own superficial analysis, we would not expect name changes for either of the Costa Rican spp. (one each of Anomospermum and Odontocarya) involved.

Rainer, H.  2007. Monographic studies in the genus Annona L. (Annonaceae):   inclusion of the genus Rollinia A. St.-Hil.  Ann. Naturhist. Mus. Wien, B 108: 191–205.

Molecular studies “in prep.” by the author have “revealed” that Rollinia is “nested within Annona” (as is the South American Raimondia, dispatched in a previous paper), notwithstanding the fact that just two spp. of Rollinia were included in the analysis.  Actually, the notion that Annona may be paraphyletic with respect to Rollinia has been kicking around for a good long time, but was apparently not addressed (according to this author) in the most recent monograph of Rollinia (Maas et al., 1992; Fl. Neotrop. Monogr. 57: 1–188).  Rainer is up to the task, however, formally relegating Rollinia to synonymy and validating all the consequent new combinations and names.  The four affected Costa Rican spp. become:  Annona danforthii (Standl.) H. Rainer, A. mucosa Jacq., A. papilionella (Diels) H. Rainer (replacing Rollinia pittieri Saff., non Annona pittieri Donn. Sm.), and A. rensoniana (Standl.) H. Rainer (replacing Rollinia membranacea Triana & Planch., non Annona membranacea R. E. Fr.).  As an added bonus, full synonymy is provided for each accepted name, together with typological details.

Rojas Alvarado, A. F.  2006. Una nueva especie de helecho del género Danaea (Marattiales: Marattiaceae) endémica de Costa Rica.  Revista Biol. Trop. 54: 1057–1060.

Danaea tuomistoana A. Rojas, honoring Finnish specialist Hanna Tuomisto, is known only from the type, collected by the author near the summit of Cerro Rincón, the highest point (745 m) on the Península de Osa.  The new spp. is most similar to the sympatric D. crispa Endres & Rchb. f., differing in its “truncate blade” and “more incised pinna with cuneate base and flat margin.”  These two spp. and D. moritziana C. Presl are compared in a table, and the new sp. is depicted in a much reduced photo of the type.  Similarly illustrated is Danaea erecta Tuomisto & R. C. Moran, formerly believed restricted to Colombia and Ecuador but here reported from Costa Rica on the basis of the author’s collection from 1500–1800 elevation on the Pacific slope of the eastern Cordillera de Talamanca (Zona Protectora Las Tablas).  The distinctions between D. erecta and D. nodosa (L.) Sm. are discussed.

-- & M. Palacios Ríos.  2006. Una especie nueva de helecho del género Pteris (Pteridaceae) endémica de Costa Rica.  Revista Biol. Trop. 54: 1061–1066.

Dedicated to famed Costa Rica botanist Gerardo Herrera, Pteris herrerae A. Rojas & M. Palacios is known by just two collections, both hailing from ca. 750 m elevation on the Atlantic slope of the northern Cordillera de Talamanca.  The new sp. is depicted in an excellent line drawing, and compared in tabular form with three related spp., among which only P. paucinervata Fée is represented in Costa Rica.

Seaton, P.  2007. Conservation in Costa Rica.  Orchid Rev. 115: 214–217.

An interesting, informal interview with Diego Bogarín and Guillaume Gigot, Darwin Initiave Project Officers in Costa Rica (Jardín Botánico Lankester) and England (K), respectively.  The focus is on collaborative ventures involving orchids, with an aim (in Gigot’s words) “to monitor orchid diversity in Costa Rica and develop new conservation strategies.”  DNA barcoding is evidently one of these new strategies.  For more information on the Darwin Initiative (which embraces far more than just orchids and Costa Rica), navigate to:


Secco, R. de S.  2004. Alchorneae (Euphorbiaceae) (Alchornea, Aparisthmium e Conceveiba).  Fl. Neotrop. Monogr. 93: 1–194.

How these things sneak past us, we’ll never know.  The neotropical members of tribe Alchorneae (Hurus.) Hutch. total 37 spp., of which 23 are in Alchornea, 13 in Conceveiba, and one in Aparisthmium.  Four spp. in the group are attributed to Costa Rica:  Alchornea costaricensis Pax & K. Hoffm., A. glandulosa Poepp., A. latifolia Sw., and Conceveiba pleiostemona Donn. Sm.  The new combination Alchornea glandulosa subsp. iricurana (Casar.) Secco, for a South American taxon, consigns Mesoamerican material to the autonymic subsp.  That is about the only new information here for us; however, we have some new information for the author:  both Alchornea triplinervia (Spreng.) Müll. Arg. (or is it A. guatemalensis Lundell?) and Aparisthmium cordatum (Juss.) Baill. have now been collected in Costa Rica.  Features synonymy and typology at all levels, formal descriptions of the tribe, genera, and all lower taxa, dichotomous keys to genera, spp., and infraspecific taxa, comprehensive lists of exsiccatae, distribution maps, an account of dubious spp., an index to exsiccatae, and many illustrations (both photos and line drawings).  The extensive introductory section addresses taxonomic history, intergeneric relationships, morphology and anatomy, dispersal, pollination, sexual systems, chromosomes, chemistry, uses, conservation, and distribution.  In Portuguese (with English abstract).  By the way, since the publication of this volume, a new revision of Conceveiba has appeared, by a different author [see The Cutting Edge 13(2): 8–9, Apr. 2006].

Simões, A. O., T. Livshultz, E. Conti & M. E. Endress.  2007. Phylogeny and systematics of the Rauvolfioideae (Apocynaceae) based on molecular and morphological evidence.  Ann. Missouri Bot. Gard. 94: 268–297.

While the focus of this paper is on subfamilial and tribal classification, a few results have repercussions (at least potentially) at generic rank.  Tonduzia, recently (on the basis of morphological-cladistic analyses) treated as a section of the large, otherwise Old World Alstonia [see The Cutting Edge 5(2): 7, Apr. 1998], does not even belong to the same tribe and is thus resegregated (the sole Costa Rican sp. involved would thus become Tonduzia pittieri Donn. Sm.).  Also, Tabernaemontana is apparently paraphyletic, with respect to the sole sp. of Stemmadenia sampled [S. litoralis (Kunth) L. Allorge], perhaps foreshadowing the eventual scuttling of the latter genus (which is not pursued here).

Tripp, E. A.  2007. Evolutionary relationships within the species-rich genus Ruellia (Acanthaceae).  Syst. Bot. 32: 628–649.

This is a meaty paper, but much of its subject matter lies outside our restricted field of interest.  Ruellia is a nearly cosmopolitan genus with ca. 300 spp., including at least 18 in Costa Rica.  Analyses of molecular data from both nuclear and chloroplast genetic regions for 196 specimens reveals that three oligotypic genera are nested within Ruellia, which is weakly supported as monophyletic with these genera included.  The only one of these nested genera of interest to us is Blechum, of which two spp. occur in Costa Rica:  B. costaricense Oerst. and the weedy B. pyramidatum (Lam.) Urb.  The author proposes to merge all three offending genera in Ruellia, and provides the necessary new combinations for two of them, but not, for whatever reason, for Blechum.  Instead, the formal transfer of Blechum to Ruellia is deferred to two other papers, one “in press” and the other “in prep.”  We believe the name Ruellia blechum L. is available for B. pyramidatum, but must await a new combination based on B. costaricense.

Tzvelev, N. N.  2006. The synopsis of the genus Glyceria (Poaceae).   Bot. Zhurn. (Moscow & Leningrad) 91: 255–276.

We quote from the English abstract:  “A complete though brief review of the genus Glyceria...is given in the paper.  The 55 accepted species are arranged according to a new system of the genus.  6 new species, 2 new sections and 2 new varieties are described; a few new combinations of different ranks are made.”  All of this has little impact on Costa Rica, with but a single sp. of Glyceria, the rare European adventive called G. plicata (Fr.) Fr. in the Manual (and all previous important regional accounts).  Here, the name G. plicata appears in synonymy under Glyceria notata Chevall., which is evidently the name we must now adopt for our material.  Includes a dichotomous (though unindented) key to spp., along with (we think) brief distributional summaries for each sp., but apparently no desciptions (all of this is in Russian, so we are groping).

Wallnöfer, B.  2007. A revision of neotropical Diospyros (Ebenaceae):  part 1.  Ann. Naturhist. Mus. Wien, B 108: 207–247.

This paper inaugurates “a series of separate publications” in which “all the currently known neotropical species of Diospyros [100–130 in all] will be dealt with. ”  If this contribution is any indication, the series will be a long one, since just three spp. (one comprising three subspp.) are here considered.  The basis for the association of these three spp. is unclear to us; it may be that the author is proceeding alphabetically, since all three epithets begin with “c,” although it is difficult to imagine that there are none beginning with “a” or “b” (moreover, the three “c” names are themselves not ordered alphabetically).  Having said all of that, only one of these first three spp. occurs in Costa Rica, where it has been known most recently as Diospyros costaricensis Provance & A. C. Sanders, considered endemic to the Cordilleras de Guanacaste and de Tilarán.  Our readers may recall [see The Cutting Edge 13(4): 7–8, Oct. 2006] that the authors of the last-mentioned name revised Mesoamerican material that had been determined as D. conzattii Standl. and D. riojae Gómez Pompa, and ended up retaining those two spp., while describing three others (including D. costaricensis) as new.  The present author undoes all of that and more, returning D. costaricensis and the other recent new spp. to the synonymy of D. conzattii, together with D. riojae!  This all boils down to status quo for us, since D. conzattii had long been applied to the Costa Rican plants, and is the name used in the Manual draft of Ebenaceae by José González (LSCR); we do, however, lose a short-lived endemic.  Features distribution maps and photographic illustrations of herbarium material; there is no key.

Webster, G. L.  2007. Taxonomic and nomenclatural changes in American Euphorbiaceae sensu lato.  Contr. Univ. Michigan Herb. 25: 235–239.

This posthumous contribution from long-time Euphorbiaceae mastermind Grady L. Webster (DAV) includes but one item of potential interest to us:  the new combination Bia cordata (Baill.) G. L. Webster (based on Zuckertia cordata Baill.).  This name would apply to the sp. presently known as Tragia bailloniana Müll. Arg. (listed in synonymy), should future authors accept Bia Klotzsch as a genus distinct from Tragia; according to Webster, Bia “appears distinct in its larger stamen number (8–40) and distinctive bifurcate inflorescences with staminate and pistillate flowers on different axes.”  He also cites a supportive molecular study [see The Cutting Edge 12(4): 13, Oct. 2005].  Incidentally, the epithet cordata is preoccupied in Tragia by T. cordata Michx. (1803).

Xiao, L.-Q. & H. Zhu. 2007. Paraphyly and phylogenetic relationships in Lasianthus (Rubiaceae) inferred from chloroplast rps16 data.  Bot. Stud. (Taipei) 48: 227–232.

According to this study, Lasianthus, “a large pantropical genus...comprising more than 180 spp.” (but with just three in the New World), is paraphyletic.  However, the potential consequences for us are unclear, inasmuch as L. panamensis (Dwyer) Robbr., the only sp. reaching Costa Rica, was not included in the analysis; moreover, the infrageneric classification of Lasianthus could not be resolved.


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