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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XXIII, Number 2, April 2016

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

The Angiosperm Phylogeny Group.  2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants:  APG IV.  Bot. J. Linn. Soc. 181: 1–20.

More than six years have passed since the publication of APG III [see under "The Angiosperm Phylogeny Group," this column, in The Cutting Edge 17(1), Jan. 2010], and much has transpired during that time.  This latest installment considers studies published since APG III, as well as the results of an online survey regarding family delimitations [see under "Christenhusz," this column, in The Cutting Edge 22(4), Oct. 2015]; however, it does not necessarily accept the recommendations from those sources.  The following innovations have significant consequences for Costa Rican floristics:  pursuant to a Nomenclature Committee decision [see under "Applequist," this column, in The Cutting Edge 22(2), Apr. 2015], Asphodelaceae supplants Xanthorrhoeaceae as the correct name for a family that includes several genera cultivated in Costa Rica, and discussed under the names Asphodelaceae and Hemerocallidaceae in Manual Vol. 2 (2003: 624–625); the formerly monogeneric Metteniusaceae is expanded to include several additional genera [see under "Stull," this column, in The Cutting Edge 23(1), Jan. 2016]; Hydnoraceae is now inlcuded in Aristolochiaceae [see under "Massoni," this column, in The Cutting Edge 21(2), Apr. 2014]; Peraceae is (finally!) segregated from Euphorbiaceae, and both Microteaceae and Petiveriaceae (which name has priority over Rivinaceae) from Phytolaccaceae (Microteaceae being monogeneric, Petiveriaceae with Petiveria, Rivina, Seguieria, and Trichostigma, in Costa Rica); Forchhammeria (formerly Capparaceae and, briefly, Stixidaceae, misspelled as "Stixaceae" in Manual Vol. 8) is moved to Resedaceae (wherein it was treated in Flora mesoamericana), as a consequence of the online survey; Nyssaceae has been revealed as "widely separated from Cornaceae" (wherein it was treated in Manual Vol. 5), and consequently reaccepted as distinct (their merger having been an innovation of APG III!); and Mazaceae (with only Mazus in Costa Rica) is accepted as distinct from Phrymaceae, contrary to the aforementioned online survey (which would have subsumed both families in Orobanchaceae).  Changes have also been made at suprafamilial levels, but these are of little interest to us.

Various other proposed changes have been resisted (at least for the time being) by APG:  Burmanniaceae remains apart from Dioscoreaceae, contrary to the results of the online survey; a suggestion to maintain Aceraceae and Hippocastanaceae as distinct from Sapindaceae by means of peeling off an additional (monogeneric) family [see under "Buerki," this column, in The Cutting Edge 16(4), Oct. 2010] is rejected, on the grounds that "no new phylogenetic information has become available" and "Sapindaceae s.l. are easily diagnosed morphologically"; Olacaceae s. l., while acknowledged as non-monophyletic, is maintained in place of numerous smaller families (Coulaceae, Erythropalaceae, etc.) for which "strong support...is lacking"; Agdestidaceae is retained within Phytolaccaceae in lieu of more data, though it "may require recognition as a segregate family" (see also under "Hernández-Ledesma," this column); Peltanthera (to be treated under Buddlejaceae in Manual Vol. 4) remains in no man's land, as an unplaced genus in Lamiales, although it may "eventually...need to be recognized in its own family" (see under "Molinari-Novoa," this column) or (as favored by the online survey) "included [together with Calceolariaceae] in an expanded Gesneriaceae"; and Adoxaceae is retained as the name for a family which, at this point in time, must correctly be called Viburnaceae [see under "Applequist," this column, in The Cutting Edge 21(2), Apr. 2014], "in the hope that the General Committee will not approve [the latest decision] in its report to the next botanical congress..." (we share their hope, as Adoxaceae is slated to appear in Manual Vol. 4, and the Manual volume that would have included Viburnaceae has already been published!).

As in the case of APG III, this contribution is not self-contained, so users will have to look elsewhere (APG II, or Peter Stevens's Angiosperm Phylogeny Website) to learn the generic content of most families.

Assis, F. C., T. E. Almeida, S. J. Russell, H. Schneider & A. Salino.  2016. Molecular phylogeny and recircumscription of the fern genus Pecluma (Polypodiaceae–Polypodiopsida).  Phytotaxa 247: 235–246.

The sundering of Polypodium sensu Flora mesoamericana Vol. 1 (1995) has been proceeding for some time now, involving the segregation of Serpocaulon [see The Cutting Edge 14(1): 10–11, Jan. 2007] and the reformulation of Pleopeltis [see the first entry under "Smith," this column, in The Cutting Edge 21(2), Apr. 2014], and largely in accordance with the plan envisioned in our review of the Serpocaulon paper.  However, the present paper marks a deviation from that trajectory, which suggested that the "Grupo de Polypodium dulce" of Flora mesoamericana would "apparently wind up...in Pecluma."  Subsequent papers, that we did not see or did not read carefully enough, have established that the "Grupo de Polypodium dulce" is in fact polyphyletic, with some spp. nested in Pecluma and others in Polypodium s. str.  Four spp. in the former situation are transferred to Pecluma in this paper, but just one of these definitely occurs in Costa Rica:  Polypodium dulce Poir. itself, henceforth Pecluma dulcis (Poir.) F. C. Assis & Salino (corrected by us from the authors' "Pecluma dulce").  One of the four new members of Pecluma, the former Polypodium rhachipterygium Liebm., had been accommodated in the "Grupo de Polypodium plesiosorum" of Flora mesoamericana, and thus would have been expected to remain in Polypodium s. str.  The sp. total for Pecluma now stands at 44.  Unfortunately, the analyses undertaken by these authors were not comprehensive, omitting 13 spp. of the "Grupo de Polypodium dulce" that "should currently be considered incertae sedis until additional molecular study is carried out."  This circumstance will prevent us from deviating one iota from the Flora mesoamericana circumsription of Polypodium, pending a final resolution.  Among the 13 spp. presently in limbo are at least four that occur in Costa Rica:  Polypodium echinolepis Fée, P. pinnatissimum R. C. Moran, P. plectolepidioides Rosenst., and P. ursipes Moritz ex C. Chr.  These could either remain in Polypodium s. str., or be transferred to Pecluma.  In any case, the morphological distinctions for Pecluma (and most of the other "genera" segregated from Polypodium s. l.) are nit-picking, to say the least, and we look forward to the next lumping cycle in this complex.

Baranow, P. & D. Szlachetko.  2016. The taxonomic revision of Sobralia Ruiz & Pav. (Orchidaceae) in the Guyanas (Guyana, Suriname, French Guiana).  Pl. Syst. Evol. 302: 333–355.

We detected just one potential ramification for Costa Rican floristics in this contribution:  Sobralia suaveolens Rchb. f., synonymized under S. bletiae Rchb. f. in the Manual Orchidaceae treatment by Robert L. Dressler (JBL), is recognized as distinct by these authors, based on the usual labellar details.  That said, we gain no insights as to which of these two ostensible spp. is to be attributed to Costa Rica; the geographic distribution of Sobralia bletiae s. str. is not provided and, although the Manual treatment is cited, Costa Rica is not among the countries enumerated in the "Distribution area" entry for S. suaveolens (which includes both Nicaragua and Panama!).  The type of S. bletiae hails from western Panama, so anything is possible (both "spp." may well occur in Tiquicia).

Blanco, M. A., J. E. Jiménez & P. Juárez.  2016. Mormodes salazarii (Orchidaceae, Catasetinae), a new species with greenish-white flowers from Costa Rica.  Phytotaxa 245: 161–168.

First things first:  congratulations to Esteban Jiménez (USJ) on his debut as a published taxonomic authority!  And, if we are not mistaken, Pedro Juárez (JVR) deserves the same accolades.  As for the plant itself:  Mormodes salazarii M. A. Blanco, J. E. Jiménez & P. Juárez equates with Mormodes sp. B of the Manual Orchidaceae treatment (Vol. 3), its epithet honoring Gerardo A. Salazar (MEXU), the first author of the Manual Mormodes account.  The new sp. is compared most closely with Mormodes fractiflexa Rchb. f., next to which it was keyed in the Manual.  The type specimen was collected by first author Mario Blanco (USJ) at 664 m elevation on the Pacific slope of the Cordillera de Tilarán (in the Monteverde region), extending the geographic range of the sp. as given in the Manual; otherwise, it remains endemic to Costa Rica (see under "Jiménez," this column, for further insights on this story).  Mormodes sp. A, we think, still has not received a formal name.  Includes a distribution map, a fine composite line drawing, and color photos of living material.

We will seize this opportunity to update our fabulous running count (of new orchid spp. described from Costa Rica since 1994) to 363.

Braem, G. J.  2016. The Phragmipedium longifolium complex.  Orchid Digest 80: 36–43.

It is a pleasure to be offered a respite from all the contentious diatribe surrounding Phragmipedium warscewiczii (Rchb. f.) Christenson [see, e.g., under "Braem," this column, in The Cutting Edge 21(3), Jul. 2014] with a consideration of P. longifolium (Warsz. & Rchb. f.) Rolfe (Orchidaceae), the only other member of its genus in Costa Rica.  But we are never satisfied:  P. longifolium turns out to be comparatively boring, and although the author provides detailed historical information, a lengthy description, and the usual plethora of color illustrations, his only real innovation is the validation of three combinations at the rank of forma.  None of these would appear to pertain directly to Costa Rica, though one extends the geographic range of the sp. (quite disjunctly) to south-central Brazil.

Caires, C. S. & C. E. B. Proença.  2015. Typification of two Neotropical names of Loranthus Jacq. (Loranthaceae).  Candollea 70: 197–199.

We rather resent having to waste time on these typification papers, and are surprised that the editors of real journals do not feel the same way.  However, we feel obliged to at least mention them when a name is involved that pertains to a sp. occurring in Costa Rica.  In the present case, that name is Loranthus cucullaris Lam., the basionym of Psittacanthus cucullaris (Lam.) G. Don; well, okay, the authorities for P. cucullaris were cited erroneously as "(Lam.) Blume" in Manual Vol. 6 (2007), so we do learn something new here (annotate your copy!).  Otherwise, this is barren and tedious fare (though certainly the easiest way to get published in a peer-reviewed taxonomic journal).

Clark, J. L., M. Nee, L. Bohs & S. Knapp.  2015. A revision of Solanum section Aculeigerum (the Solanum wendlandii group, Solanaceae).  Syst. Bot. 40: 1102–1136.

This seems to be the season for revisions of taxa with but a single sp. in Costa Rica (see, e.g., under "Cotton" and "Gutiérrez," this column), and this time there are no big surprises (see, especially, under "Sartori"):  although the eight spp. of the titular taxon are widespread collectively in the Neotropics, only Solanum wendlandii Hook. f. itself occurs in Costa Rica.  Though seldom encountered in the wild, S. wendlandii is quite well-kwown in Tiquicia as a cultivated ornamental.  Readers may wish to annotate their Manual copy with some additional information that is presented here, but none of it is earth-shattering.  Features synonymy and typology, technical descriptions of Solanum sect. Aculeigerum Seithe and each of its spp., a dichotomous and indented key to spp., distribution and phenology summaries, distribution maps, discussions (sometimes lengthy), representative specimen citations, and a section on "Names not validly published."  The brief introductory part conflates taxonomic history with taxonomy and phylogeny, and considers important aspects of morphology.  All spp. are illustrated with black-and-white photos of herbarium specimens, and Solanum wendlandii also with a color lithograph of "the plant from which the neotype specimen was probably taken."  Selected spp. are depicted (in the introductory pages) in two plates of color photos from life; one of these, showing the fruits of S. wendlandii, is revelatory (the measurements given in the Manual description should be doubled!).

Cotton, E., F. Borchsenius & H. Balslev.  2014. A revision of Axinaea (Melastomataceae).  Sci. Danica B 4: 1–120.

We missed this the first time around, and finally came to it by a circuitous route.  While it is a highly satisfying piece of work in all respects (the page formatting is exemplary), this revision (the first of Axinaea in nearly 80 years) offers nothing radically new for Costa Rican floristics:  Axinaea costaricensis Cogn., the only member of its genus included in the Manual Melastomataceae treatment (Vol. 6), retains that status, and is, indeed, the only Central American representative of this genus of 41 spp., otherwise restricted to Andean South America.  However, A. costaricensis no longer qualifies as a Central American endemic, as was indicated in the Manual:  it also occurs in Venezuela (according to these authors), which "represents the largest disjunction in distribution found among any species of Axinaea."  In general, the genus is characterized by a high level of endemism (28 of its spp. being endemic to one country).  Features synonymy, typology, and technical descriptions at all ranks, a dichotomous and indented key to spp., distribution summaries and maps, citations of specimens examined, generally brief discussions, a section on "Doubtful names and excluded taxa," and indices to exsiccatae and scientific names.  Topics addressed in the well-illustrated introductory part include taxonomic history, morphology and anatomy, karyology, relationships, and distribution.  Eight new (South American) spp. are described, all of which (along with selected other spp.) are illustrated with composite line drawings and/or color photographic plates (one of the latter being provided for A. costaricensis).

Dórea, M. C., R. P. Oliveira & C. A. D. Welker.  2016. Lectotypifications in Orthoclada (Poaceae, Panicoideae, Zeugiteae).  Phytotaxa 246: 159–162.

Lectotypes are designated for Aira laxa Rich. [the basionym of Orthoclada laxa (Rich.) P. Beauv., well known in Costa Rica] and one of its synonyms.  The applications of the names are not affected.

Gahagen, B., M. Terbush & H. Ballard, Jr.  2015. A taxonomic synthesis of the Tovomita weddelliana (Clusiaceae) species complex.  Syst. Bot. 40: 968–988.

Phenetic analyses suggested to these authors that "there are multiple morphologically distinct species currently subsumed under the name Tovomita weddelliana," prompting "the resurrection of synonyms and description of new taxa."  So resolved, the titular complex comprises at least 16 spp., 11 denoted by resurrected synonyms and five here described as new (one name, the Colombian Tovomita glossophylla Cuatrec., could not be evaluated).  All the Costa Rican material referrable to this complex has long been subsumed under the name Tovomita weddelliana Planch. & Triana as, e.g., in co-PI Barry Hammel's Manual Clusiaceae treatment (Vol. 5), which cited Clusia oblanceolata Rusby and C. pithecobia Standl. & L. O. Williams as synonyms.  Both of the latter names are here "resurrected" (though with C. oblanceolata banished from the genus Tovomita), whereas T. weddelliana is now restricted to a small area of Bolivia.  Clusia pithecobia (type from the Golfo Dulce region) is recast as Tovomita pithecobia (Standl. & L. O. Williams) Gahagen (comb. nov.) and deployed for populations from the Pacific-slope portion of the range of T. weddelliana (in the sense of the Manual).  All the Atlantic-slope populations in Costa Rica are referred to one of the new spp., Tovomita divesora Gahagen, although (somewhat discouragingly) three collections from the Pacific slope are included among the paratypes; so, is it the biogeography that makes no sense, or the authors' methodology?  And from a slightly different perspective:  phenetic analysis may (or may not!) be an effective means of documenting local morphological differentiation; but does local differentiation necessarily imply speciation?  We are strongly persuaded that it does not.  Whatever the case, this new taxonomy, should it hold up to scrutiny, does reward us with two new Costa Rican endemics!  Features a dichotomous and indented key to spp., distribution maps, and very crude line drawings for several spp. (including T. divesora, but not T. pithecobia).  According to the key, T. pithecobia differs from T. divesora by its fruits with "the stigmatic region constricted or raised ca. 1 mm from the surface" (vs. sessile).

Gutiérrez, D. G. & L. Katinas.  2015. Systematics of Liabum Adanson (Asteraceae, Liabeae).  Syst. Bot. Monogr. 97: 1–121.

Liabum, in the modern sense (i.e., shorn of such genera as Erato, Munnozia, Oligactis, and Sinclairia, which it once included), is a neotropical assemblage of 22 spp. (according to this revision), best represented in Andean South America.  Just one sp. has been ascribed to Costa Rica, and that situation remains unchanged.  However, we must now accustom ourselves to a new name, as the familiar Liabum bourgeaui Hieron. (type from southern Mexico) is synonymized by these workers under L. asclepiadeum Sch. Bip. (based on a Venezuelan collection).  Their decision was based on the observation that winged petioles, "the only useful character to distinguish" the two entities, "can be present in both species."  Liabum bourgeaui had been considered a Mesoamerican endemic, but its rebranding as L. asclepiadeum extends the geographic range to Colombia and Venezuela.  It bears mentioning, however, that the authors' taxonomic conclusions appear based overwhelmingly on herbarium work, with very limited field experience, so time will tell how well they hold up.  In terms of its presentation, this is an exemplary piece of work, upholding the very high standards of this time-honored series, now edited by David M. Johnson (OWU).  In addition to all the usual features of taxonomic monographs, indices to exsiccatae and scientific names are much appreciated, as are the distribution maps and excellent composite line drawings provided for each sp.  Two new spp. are described (from Ecuador and Peru), and numerous lectotypes, neotypes, and epitypes are newly designated (including lectotypes for both Liabum bourgeaui and L. asclepiadeum).

Hernández-Ledesma, P., W. G. Berendsohn, T. Borsch, S. von Mering, H. Akhani, S. Arias, I. Castañeda-Noa, U. Eggli, R. Eriksson, H. Flores-Olvera, S. Fuentes-Bazán, G. Kadereit, C. Klak, N. Korotkova, R. Nyffeler, G. Ocampo, H. Ochoterena, B. Oxelman, R. K. Rabeler, A. Sanchez, B. O. Schlumpberger & P. Uotila.  2015. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales.  Willdenowia 45: 281–383.

The consensus classification presented by this group accepts 39 families and 749 genera for this order, comprising approximately 12,500 spp.  The list of accepted families inevitably deviates from the Manual (given our alphabetical constraint), but there are just two mild surprises vis-à-vis APG (see the first entry in this column):  the acceptance of Agdestidaceae (with only Agdestis) as distinct from Phytolaccaceae, and of Chenopodiaceae as distinct from Amaranthaceae (will this ever be straightened out?).  We are less interested in the accepted genera (and the cactus crowd will never be bound by these decisions; see, e.g., under "Lodé," this column).  The checklist that occupies most of this paper supplies bibliographic references, type spp., and synonymy, as well as frequent discussions.  The extensive (22 pages!) bibliography will certainly prove extremely useful to our work on these families.  It's too bad that the authors did not see fit to include an index to genus names (or at least synonyms), and their Web site (http://caryophyllales.org/) does not provide a satisfactory resolution, in our opinion.

Hurtado-Hernández, P. M. & J. A. González-Ramírez.  2013. Lista anotada de las plantas del Parque Nacional Tortuguero.  Revista Biol. Trop. 61(Supl. 1): 121–130.

The authors' list (which, despite the claim made in the title, is not annotated in any way) totes 732 spp. of vascular plants in 413 genera, based on their own collections in the study region and (largely, we suspect) the ATTA and TROPICOS databases.  Even though the site has been well-collected and the sources are impeccable, we are surprised by the low sp. total—especially considering that the nearby (and much smaller) Estación Biológica La Selva boasts at least twice as many!  It is easy to imagine various explanations for this discrepancy, and a glance at the list suggests one possibility:  that collectors working in Tortuguero have not paid much attention to certain families, in particular, Orchidaceae (with just six spp. accounted for).  So, as the authors realize, their list is not the final word, and "sin duda otros estudios revelarán nuevas especies" in the area.  At the same time, many of the plant spp. characteristic of the Costa Rican Atlantic coastal plain range very widely, as illustrated by the occurrence at Tortuguero of just two spp. that are endemic to Costa Rica—Paullinia austin-smithii Standl. (Sapindaceae) and Pseuderanthemum pittieri Leonard (Acanthaceae).  La Selva is backed up against the Cordillera Central, home to many locally endemic spp. that must enrich the flora to a considerable degree.

Jiménez, J. E., P. Juárez & A. Díaz.  2016. Checklist of the vascular flora of Reserva Biológica San Luis, Costa Rica.  Check List 12(2): 1859(1–23).

The titular reserve, occupying 251 ha on the Pacific slope of the Cordillera de Tilarán (below Monteverde) at 540–855 m elevation, was created in 1995 by the Tropical Science Center.  It is in an area of wet forest (bosque húmedo).  The authors' inventory, involving 15 excursions (45 days total) during 2011–2014, generated 1086 herbarium specimens, which were augmented by 48 others collected previously on the site.  These revealed 716 spp. of native vascular plants, representing 477 genera and 130 families.  Fabaceae (67 spp.) was the most diverse family, and Piper (15 spp.) the most diverse genus.  Befitting the journal, the bulk of this paper consists of a vouchered checklist, annotated to indicate life form (shrub, tree, etc.) and habit (terrestrial, epiphytic, etc.), and whether each sp. also occurs in the El Rodeo region of the Valle Central (with a similar vegetation type).  The introductory part, featuring numerous color photographs as well as tables and graphs, expounds upon the vegetation analyses.  One new sp. was discovered by the authors, viz., Mormodes salazarii M. A. Blanco, J. E. Jiménez & P. Juárez (Orchidaceae; see under "Blanco," this column).

Judkevich, M. D., R. M. Salas & H. Keller.  2016. Randia brevituba (Rubiaceae), a new species from the Southern Cone of America and comments on Randia armata.  Syst. Bot. 41: 238–244.

As indicated in the discussion for the widespread Randia armata (Sw.) DC. in the Manual treatment of Randia by Charlotte M. Taylor (MO) and colleagues, the circumscription and distribution of said sp. have been contentious issues.  According to the interpretations of the present authors, the real R. armata does not occur in the South American "Southern Cone."  While the latter phrase is nowhere defined precisely in this paper, we surmise that readers inclined to accept the taxonomy of these authors will want to delete both "Par." and "Arg." from the Manual distrbution summary for R. armata.

Lodé, J.  2015. Taxonomy of the Cactaceae.  2 vols. Joël Lodé.

Here we have nothing less than "the new classification of Cacti mainly based on molecular data and explained" (to quote the subtitle of this work).  With cactophiles stumbling over themselves in an effort to mark out their taxonomic territory, this represents at least the third massive contribution along these lines in the past 15 years, following on the heels of Edward Anderson's The cactus family [see The Cutting Edge 8(4): 2–3, Oct. 2001] and David Hunt's New cactus lexicon (2006).  Hunt's group, in particular, has sought to position itself as the ultimate arbiter (APG-style) in matters of cactus classification (and we would probably be doing well to accept them as such).  But Joël Lodé, an energetic, well-connected, and evidently well-heeled amateur hip to the ongoing revolution in molecular systematics, is having none of that.  Repeatedly bashing Hunt for his perceived overlumping at the rank of genus, Lodé seeks to set the record straight in this vanity-press publication by recognizing a total of 177 genera for Cactaceae (vs. 127 that were accepted by Hunt, and only 125 by Anderson).  In so doing, he heeds the results of molecular studies (when it suits him), but also seems guided by an odd notion of priority requiring that every genus name of a certain antiquity be deployed (if only for a single sp.).  Two other motivations (that are also recurring themes in this work) for Lodé's splitting are the fear that, with fewer genera, Cactaceae will be perceived as less diverse than he fancies, and the trauma of frequently having to change the labels of potted plants.  Never mind that amateur cactophiles themselves (including Lodé) have been major players in the name-changing game, seizing every opportunity to recombine forms and varieties as subspp., subspp. as spp., sections and subgenera as genera, and vice-versa back down the line, in their own hobbyist journals (of which there are dozens internationally).  Most of the changes proposed or debuted in these volumes, and that impact Costa Rican floristics, have already been reported upon in these pages.  There is just one minor revelation, viz., the sequestering of the sp. known lately to us as Stenocereus aragonii (F. A. C. Weber) Buxb. in the monospecific genus Marshallocereus [the combination M. aragonii (F. A. C. Weber) Backeb. being already available].  This, we suspect, reflects a reluctance on Lodé's part to sweep Marshallocereus and Stenocereus (and probably other genera) into Carnegiea, a sacred name and concept to cactophiles—or, perhaps, the mistaken notion of Arias et al. [see The Cutting Edge 10(4): 4, Oct. 2003], which we ourselves blindly accepted, that Carnegiea and company might be swept into Pachycereus [Carnegiea itself is actually the oldest of the contending genus names in those authors' "Pachycereus group"!].  Let's get the rest of our criticisms out of the way, before moving along to the good points (and there are some) of this publication.  First, the English edition of this work (we understand there are editions in other languages) would have benefited by more (or better) textual editing.  The author's style is overly animated and polemic, and the text (particularly in the introductory part) tends to be poorly organized, disjointed, and repetitive.  His English is adequate, but far from perfect, and should have been refined (or better refined) by an editor.  And (here is where the "vanity" part comes into play) the "Illustrations" sections, which occupy approximately half of each volume, should have been eliminated.  These volumes are extremely unwieldy (one review pegs their combined weight at 15½ lbs.!) and, minus these extra illustrations, could have been condensed into a single volume at a lower price.  Although they are nice to look at (and will probably help sell copies), the supplemental illustrations (mostly color photos of plants) are unnecessary (the text is already well-illustrated) and break up the flow, undermining what we believe to be the author's main agenda.

So, what do we actually like about this work?  That the author has one foot in the amateur cactophile community and the other in the heady realm of molecular systematics positions him uniquely to bridge that gap.  And he (together with someone named Fabrice Cendrin) makes a valiant, and at times successful, effort to do just that in the introductory portion of Vol. 1, addressing such topics as classification, speciation, interfertility, phylogeny, genetics and molecular systematics, "Why plants change names?," the origin of cacti, and pollinators and dispersers, all in language tailored for the amateur cactophile.  Indeed, a few pearls of wisdom are to be found among these words.  The main portion of each volume (exclusive of the supplemental photos!) is an alphabetical compendium of cactus genera, including even those genus names that are not accepted (with appropriate cross-referencing).  The entries for the accepted genera generally comprise at least three pages (one being a full-page color photo), with an explanation of etymology, a brief description, "Comments" (often a loose diatribe on various alternative classifications), and habitat and distribution summaries, as well as miscellaneous additional photos, a distibution map and a graph charting altitudinal range, and a list of currently accepted spp.  Most useful to our work will be the author's firm command of the far-flung hobbyist literature on Cactaceae, which we are at great pains to track; many recent articles dealing with (or delving into) relevant taxonomic issues are cited here, often in journals that our library does not even receive.

Now, imagine the outcry from this crowd if Cactaceae had been sunk into Portulacaceae (or vice-versa), as (arguably) ought to have been done [see, e.g., The Cutting Edge 14(2): 8–9, Apr. 2007].

Loría-Naranjo, M., J. Samper-Villarreal & J. Cortés.  2014. Structural complexity and species composition of Potrero Grande and Santa Elena mangrove forests in Santa Rosa National Park, North Pacific of Costa Rica.  Revista Biol. Trop. 62: 33–41.

"Structural complexity" is where the emphasis lies, in this appraisal of two of the northernmost mangrove communities in Costa Rica.  There is not much here regarding "species composition," beyond lists of the five or six (depending on the site) mangrove tree spp. themselves.  The authors' discovery of Pelliciera rhizophorae Planch. & Triana (Tetrameristaceae) in the Santa Elena mangroves nudges the northern distributional limit for that sp. in Costa Rica to the north coast of the Península de Santa Elena (it was already known from the Potrero Grande site, on the south coast of the same peninsula).  The authors recommend the creation of a "marine protected area" in the Bahía de Santa Elena.  This paper appears in a special issue entitled "Investigaciones científicas en la zona marino-costera del Pacífico Norte de Costa Rica," edited by Jorge Cortés and Álvaro Morales-Ramírez.

Marhold, K., M. Šlenker, H. Kudoh & J. Zozomová-Lihová.  2016. Cardamine occulta, the correct species name for invasive Asian plants previously classified as C. flexuosa, and its occurrence in Europe.  PhytoKeys 62: 57–72.

Recent evidence, both molecular and morphological, has revealed that European and east Asian populations that have been identified as Cardamine flexuosa With. "belong to two different taxa."  The name C. flexuosa correctly applies to the taxon native to Europe, while C. occulta Hornem. is adduced by these authors (who lecto- and epitypify it) for the taxon native to east Asia.  The authors are not altogether forthcoming as regards the morphological distinctions between these two alleged spp. (a paper is said to be "in prep.") nor their geographical distributions, which is exasperating for us, because one (or both?) is widely adventive in the New World, including Costa Rica.  We suspect that ours may be the east Asian C. occulta, based mainly on the following statement:  "...unlike European C. flexuosa, C. occulta represents an invasive species that is quickly spreading from its area of origin in Eastern Asia to other continents."  And while we cannot find Costa Rica mentioned anywhere in the article, C. occulta is attributed (in addition to east Asia) to Canada, the United States, Mexico, Cuba, the Canary Islands, and Australia, and has even become (since 1993) widespread in Europe itself!  We look forward to a more rigorous explication of this situation.

Meza-Torres, E. I.  2015. Notas nomenclaturales en Ophioglossum (Ophioglossaceae).  Bol. Soc. Argent. Bot. 50: 627–630.

The name Ophioglossum ellipticum Hook. & Grev., generally accepted as a synonym of O. nudicaule L. f. (very scarce in Costa Rica), is lectotypified on a specimen suggested to have been the basis for an illustration previously considered to be type (based on our comparison of the two relevant images, we are not persuaded!).  Usage would appear to be unaffected.  Moving right along...

Molinari-Novoa, E. A.  2016. Two new lamiid families for the Americas.  Weberbauerella 1(7): 1–4.

So now we have two new family names, both potentially applicable to genera occurring in Costa Rica, validated by an individual who evidently spends much of his time surfing the Internet to peruse the careful work of others in search of just this sort of opportunity, and who now has his own vanity-press, online journal to facilitate the whole nefarious enterprise.  Enough said.  Namaceae Molinari, segregated from Hydrophyllaceae, would include four genera, one of which (Wigandia) is represented in Costa Rica.  Peltantheraceae Molinari accommodates the monospecific Peltanthera, widespread in the Neotropics (including Costa Rica), which has been bounced among Buddlejaceae, Loganiaceae, and Solanaceae, and could eventually find a home in Gesneriaceae (see under "The Angiosperm Phylogeny Group," at the head of this column).  Oddly, the subfamily names Namoideae Molinari and Peltantheroideae Molinari are also validated here (can a family have just one subfamily?).  All the new taxa are minimally diagnosed.

Mora, M. M. & J. L. Clark.  2016. Molecular phylogeny of the Neotropical genus Paradrymonia (Gesneriaceae), reexamination of generic concepts and the resurrection of Trichodrymonia and Centrosolenia.  Syst. Bot. 41: 82–104.

Paradrymonia, with 40 recognized spp. (according to these authors), "is resolved as polyphyletic and includes taxa in three clades" and, as a consequence, is split into three distinct genera, under the names mentioned in the title.  Paradrymonia s. str. (include the generic type sp.) is reduced to just eight spp., sharing (inter alia) anisophyllous leaves with decurrent bases, linear to lanceolate calyx lobes, crenate to fimbriate corolla lobes, and bearded anthers.  So defined, Paradrymonia is left with but a single sp. in Costa Rica (vs. eight, according to Manual Vol. 5); fortunately, it is our most common sp., P. decurrens (C. V. Morton) Wiehler, but unfortunately, that familiar name must now yield to the prior P. ciliosa (Mart.) Wiehler.  The remaining seven spp. treated under Paradrymonia in the Manual (together with P. alba Wiehler and P. ommata L. E. Skog, mentioned in the genus discussion) are all transferred in straightforward fashion to the "resurrected" Trichodrymonia Oerst., suddenly bloated to some 40 spp. having in common leaves clustered in an apical rosette, sulcate petioles, salverform to trumpet-shaped or hypocyrtoid corollas, and glabrous anthers (among other things).  Although it has been used in the past for some Costa Rican spp. now included in Paradrymonia s. str. or Trichodrymonia, the name Centrosolenia, as here "resurrected" on the basis of the generic type sp., now applies to 15 spp. endemic to the Guiana Shield region, and is of no concern to us.  The results of this study "also show that Nautilocalyx is clearly paraphyletic, but the recognition of additional new genera is not recommended until more comprehensive taxon sampling is included, as well as additional loci."  Nonetheless, four of its erstwhile spp. [including N. panamensis (Seem.) Seem., mentioned in the Manual genus discussion] share a clade with spp. of Chrysothemis (based on molecular and/or morphological evidence) and are transferred to the latter genus, nearly doubling its size.  We are unable to speculate on the potential fate of the two Nautilocalyx spp. in Costa Rica [N. dressleri Wiehler having recently been transferred to Drymonia; see under "Clark," this column, in The Cutting Edge 19(1), Jan. 2012], as they were not included in the analyses.  We wonder whether all of these genera might better have been combined into one (Centrosolenia being the oldest name, though Nautilocalyx is conserved against it).  Features a (highly artificial) key to the genera of the "Paradrymonia alliance," revised generic descriptions, an index to accepted names, and color photos of living specimens.  All the new combinations in this paper are credited to "M. M. Mora & J. L. Clark."

Perrie, L. R., L. D. Shepherd & P. J. Brownsey.  2015. An expanded phylogeny of the Dennstaedtiaceae ferns:  Oenotrichia falls within a non-monophyletic Dennstaedtia, and Saccoloma is polyphyletic.  Austral. Syst. Bot. 28: 256–264.

The title says just about everything that can be said about this paper.  Dennstaedtia appears diphyletic, with the oligospecific, Australasian Oenotrichia nested in one of its two clades.  Saccoloma would also appear diphyletic, although, rather problematically, two samples identified as S. elegans Kaulf. (the type sp.!) were resolved in different clades.  This is trumpeted as "the first phylogenetic study to include all of the genera recognised in the Dennstaedtiaceae...and the first to include multiple species for all of the genera that are not monotypic."  Be that as it may, it is so stunningly preliminary and ill-conceived (the type spp. of several important genus names, including Dennstaedtia itself, having been omitted) that almost nothing may be concluded with regard to the circumscriptions or correct names of potential segregate genera (though the authors spin their wheels trying).

Rueda, R., O. Jarquín, B. Munguía, A. Reyes & I. Coronado.  2013. Inventario preliminar de plantas asociadas a los pantanos dominados por palmas del Caribe de Costa Rica y Nicaragua.  Revista Biol. Trop. 61(Supl. 1): 101–120.

Palm swamps, or "yolillales," generally dominated in the region specified in the title by Raphia taedigera (Mart.) Mart. or (more locally) Astrocaryum alatum H. F. Loomis or Manicaria saccifera Gaertn., are notoriously low in plant-sp. diversity, often verging on monocultures.  What is not specified in the title is that this inventory is limited to a relatively small portion of the Caribbean coastal plain of Nicaragua and Costa Rica, specifically, at the southern end of Lago de Nicaragua in the former country, and in the Refugio de Fauna Silvestre Barra del Colorado and Parque Nacional Tortuguero, in northeastern Costa Rica (a total of eight palm swamps in five locations); furthermore, it involves only trees.  The authors' master list accounts for 60 tree spp. (three of which could not be identified) in the their study sites, among which are 13 spp. culled from an unpublished Ph.D. dissertation by another worker.  Even so, we are surprised by the total, though it clearly includes many spp. found growing in close proximity to the "yolillales," rather than in their very midst.  Had the inventory been extended southward, to the Panamanian border, many other interesting spp. could have been added on similar grounds, e.g., Anaxagorea panamensis Standl. and Unonopsis storkii Standl. & L. O. Williams (Annonaceae) and Casearia standleyana Sleumer (Salicaceae).  No vouchers are cited in this paper, but specimens are said to have been collected and deposited in Nicaraguan herbaria.  Twenty-three of the more common tree spp. are briefly described (with phenological and distributional data), and most of these are illustrated with color photos from life.  The supplement that includes this paper (as well as that of "Hurtado-Hernández," this column) is entitled "Yolillales:  los humedales olvidados" (edited by Mahmood Sasa).

Sartori, A. L. B., G. P. Lewis, V. F. Mansano & A. M. G. A. Tozzi.  2015. A revision of the genus Myroxylon (Leguminosae: Papilionoideae).  Kew Bull. 70(4): 48(1–12).

In Manual Vol. 5 (2010), co-PI Nelson Zamora characterized Myroxylon as comprising two spp., with only M. balsamum (L.) Harms represented in Costa Rica.  The status of both the genus and M. balsamum remain substantially unchanged, although these authors decline to recognize infraspecific taxa for the latter [Nelson had accepted Mesoamerican material as belonging to M. balsamum var. pereirae (Royle) Harms].  Also, the range of M. balsamum (and hence that of the genus) should be extended to include Cuba.  We had assumed that would be the end of the matter, but in a shocking development, the authors cite a Costa Rican voucher for the other sp. of Myroxylon, M. peruiferum L. f.  This has been considered exclusively South American and, indeed, TROPICOS contains just one specimen record for Mesoamerica (a Mexican specimen, determined by the first author of this paper).  Nonetheless, specimens of M. peruiferum are cited in this revision from throughout Mesoamerica, including El Salvador, Honduras, and Nicaragua.  The Costa Rican voucher, Cascante 1324, is a sterile collection from 900 m elevation on the Pacific versant of the Valle Central (Zona Protectora El Rodeo).  A second specimen (also sterile) from the same site in TROPICOS (as M. balsamum) could not be located in the herbarium at MO.  Vegetatively, Myroxylon peruiferum is distinguished from M. balsamum by "its leaflet with a shiny adaxial surface, the leaflet blade with conspicuous pellucid dots and streaks" (vs. leaflets dull adaxially in M. balsamum, with the pellucid dots and streaks "inconspicuous...unless held up to the light").  Comparing various specimens (including Cascante 1324) determined as one sp. or the other by the authors, we judge these particular characters to be of little or no practical utility (at least to the unschooled observer) in the herbarium, and prefer to await fertile Costa Rican collections before accepting M. peruiferum as a member of the flora.  Features synonymy and typology, detailed descriptions, distribution and phenology summaries, representative specimen citations, "notes," and a section on "Excluded taxa."  The introductory part, concerned mainly with taxonomic history, ethnobotany, and phytochemistry, also includes a composite distribution map.  Both spp. are depicted in very fine composite line drawings.

Treiber, E. L., A. L. Gaglioti, S. Romaniuc-Neto, S. Madriñán & G. D. Weiblen.  2016. Phylogeny of the Cecropieae (Urticaceae) and the evolution of an ant-plant mutualism.  Syst. Bot. 41: 56–66.

The most significant result of this study, from our viewpoint, is the demonstration by morphological and (especially) molecular analyses that the familiar, largely myrmecophytic neotropical genus Cecropia is paraphyletic, with respect to the non-myrmecophytic and much smaller African Musanga, embedded in the former with strong support.  It would appear that monophyly could be restored through either of two strategies:  the synonymization of Musanga (the later name) under Cecropia, or the transfer of a single South American Cecropia sp. to Musanga.  While the authors betray a clear inclination toward the former option, they take no action at the present time, pending "more complete sampling" (in particular, one of the two Musanga spp. has not been sequenced).  Cecropia and Musanga are portrayed as "ecologically similar" and "with highly similar vegetative and reproductive characteristics..."

Wahlert, G. A., H. E. Ballard, Jr. & J. Paula-Souza.  2015. Ixchelia, a new genus of Violaceae from Mexico and Mesoamerica.  Brittonia 67: 273–283.

The opening salvo from this lab [see under "Wahlert," this column, in The Cutting Edge 21(2), Apr. 2014] promised to break up the demonstrably polyphyletic Hybanthus in such a way that only H. yucatanensis Millsp. (among the Costa Rican representatives) would remain in that genus, with our other 10 spp. to be apportioned among four segregate genera.  The present contribution, which follows the unveiling of Pombalia [see under "Paula-Souza," this column, in The Cutting Edge 22(1), Jan. 2015], brings us to the halfway point (in terms of genera).  The new genus name Ixchelia H. E. Ballard & Wahlert (derived from the name of a Mayan deity) is coined, with two new combinations at sp. rank validated thereunder.  One of the latter, Ixchelia mexicana (Ging.) H. E. Ballard & Wahlert (based on Hybanthus mexicanus Ging.),  pertains to a sp. occurring in Costa Rica.  The second sp. (a transfer from Rinorea) is distributed disjunctly in southern Mexico and central Panama.  A table is deployed for comparison of the new genus with its closest relatives, Isodendrion and Pombalia.  This paper amounts to a fully realized revision of Ixchelia, with synonymy and typology, generous descriptions, a key to spp., distribution and phenology summaries, a distribution map, discussions, and comprehensive specimen citations.  Both spp. are depicted in composite line drawings.

Wong S. Y., A. W. Meerow & T. B. Croat.  2016. Resurrection and new species of the Neotropical genus Adelonema (Araceae: Philodendron clade).  Syst. Bot. 41: 32–48.

This study corroborates a prior one from the same lab [see under "Wong," this column, in The Cutting Edge 20(4), Oct. 2013] in portraying Homalomena sect. Curmeria (André) Engl. (comprising all the New World spp. of Homalomena) as nested within the immense neotropical genus Philodendron and sister to P. subgen. Pteromischum (Schott) Mayo.  This scenario mandates "patently signficant taxonomic changes" (quoting from the earlier paper), with three principal options:  to submerge Homalomena sect. Curmeria in Philodendron (perhaps as a fourth subgenus) or to elevate it to the rank of genus under the name Adelonema Schott, which would either include Philodendron subgen. Pteromischum or exclude it (as another distinct genus, under the name Elopium Schott).  In our review of the previous paper, we advocated the first of these options (submerging sect. Curmeria in Philodendron) as the least nomenclaturally disruptive and most taxonomically sensible.  After all, sect. Curmeria comprises just 15 or so spp., several of which already have names available in Philodendron; moreover, the taxon barely registers in the public consciousness (even many non-specialist taxonomists are unaware of it), differs from Philodendron in just a few highly technical characters, is uncommon and rarely recognized in the field (being routinely misidentified as Philodendron), is rarely cultivated, and has not held the rank of genus at any time during the past century.  Despite these common-sense considerations, these authors have elected to grant generic recognition to sect. Curmeria, noting somewhat disingenuously that their results "[support] resurrection of Adelonema as a genus distinct from Asian Homalomena" (true, but no more than they support either of the other two available options).  All the "necessary" new combinations in Adelonema are validated in the names of "S. Y. Wong & Croat."  But the real harm in "resurrecting" Adelonema (previously used for a single sp. from 1860–1912) is yet to come, as name-changes are now mandated (according to the cladistic evidence) for all ca. 75 spp. of Philodendron sect. Pteromischum, "which should deserve generic recognition with the name Elopium.." (the authors having evidently rejected the option of including sect. Pteromischum in Adelonema).  As in the case of Adelonema, the name Elopium has only ever been used for a single sp. (from 1865–1878); yet, now (if we accept the taxonomy of these authors) it will have to be visited upon such familiar and widespread spp. as Philodendron guttiferum Kunth, P. inaequilaterum Liebm., and P. lingulatum (L.) K. Koch, to mention just a few.  Moreover, once subgen. Pteromischum has been pried away from Philodendron, there will probably be a strong temptation to also remove the arguably even more distinctive subgen. Meconostigma Engl. (with such well-known horticultural subjects as Philodendron bipinnatifidum Schott ex Endl. and/or P. selloum K. Koch), even if that is not strictly required on cladistic grounds.  None of this has happened yet (though some of it is all but inevitable), on the premise that "much more comprehensive sampling, especially of subgen. Pteromischum, is required..."

Having gotten all of that off our chests, on to the nitty-gritty:  three of the four Costa Rican representatives of "Adelonema" required new combinations under that name, viz., the former Homalomena hammelii Croat & Grayum, H. picturata (Linden & André) Regel, and H. wendlandii Schott; the combination Adelonema erythropus (Schott) Schott was already available for our fourth sp.  The taxon called Homalomena erythropus (Schott) Engl. in Manual Vol. 2 (2003), subsequently distinguished as H. e. subsp. allenii Croat [see The Cutting Edge 11(4): 6, Oct. 2004], is here elevated with zero rationale to sp. rank, as Adelonema allenii (Croat) S. Y. Wong & Croat; this we reject (and would rather use the name A. erythropus), on the grounds that the taxon in question has never been adequately justified even at the rank of subsp.  One thing that we do agree with (based on our field experience) is the division of "Adelonema" into two sections, i.e., A. sect. Adelonema (with unarmed petioles and glabrous leaf-blades) and A. sect. Curmeria (André) S. Y. Wong & Croat (comb. nov.; with often prickly petioles and pubescent leaf-blades); in Costa Rica, the autonymic section houses the former Homalomena hammelii and H. erythropus ("Adelonema allenii"), while sect. Curmeria accommodates H. picturata and H. wendlandii.  Finally, four new spp. are described, though none of these occurs in Costa Rica.  Features a key to the genera of the "Philodendron clade" (also including the two sections of "Adelonema"), a key to the spp. of "Adelonema," and numerous fine color photographs of living material.  And, incidentally:  the genus name Adelonema is neuter, not feminine, as assumed by these authors, so several of its epithets require correction; the only Costa Rican sp. so affected is Adelonema picturatum (not "picturata").



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