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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XXVII, Number 1, January, 2020

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

Aguilar-Sandí, D. 2019. El árbol Myroxylon peruiferum (Fabaceae), nuevo registro en la flora de Costa Rica. Cuad. Invest. UNED 11: 424–429.

Myroxylon peruiferum L. f. was first reported from Costa Rica several years ago [see under "Sartori," this column, in The Cutting Edge 23(2), Apr. 2016] on the basis of a single, sterile collection from the Zona Protectora El Rodeo on the Pacific slope of the Valle Central; however, we were not persuaded by the evidence, and preferred "to await fertile Costa Rican collections before accepting M. peruiferum as a member of the flora." Taking the proverbial bull by the horns, this author proactively sought and collected the desired material in the central Pacific region of Costa Rica, and here confirms the existence of two Myroxylon spp. in the country. These are carefully distinguished textually and with color photos, and all known Costa Rican specimens of M. peruiferum are cited and their distribution mapped. Summarizing in Manualese yields the following statement for M. peruiferum: "Bosque húmedo, (250–)800–1250 m; vert. Pac. Valle Central, vecindad de San Mateo, cuenca del Río Grande de Candelaria. Fl. feb." The only flowering record is apparently from an individual cultivated on the campus of the Universidad de Costa Rica, near San José. Detailed data are not provided for Myroxylon balsamum (L.) Harms (a missed chance), but it is said to occur principally in the "zona sur del país (vertiente pacífica) entre 0 y 100 m de altitud."

Incidentally, Manual co-PI (and Fabaceae author) Nelson Zamora had arrived independently at a similar conclusion, following a recent visit to K and BM. He also reports that, luckily, both the Manual voucher and illustration purporting to represent Myroxylon balsamum, as well as at least the lion's share of the description, do indeed pertain to that sp., rather than M. peruiferum.

This journal uses two different titles, one English and one Spanish. We flipped a coin!

Biral, L. 2019. A new combination in Monteverdia (Celastraceae). Brittonia 71: 401–402.

In our review of a previous paper (helmed by this author) segregating Monteverdia from Maytenus [see under "Biral," this column, in The Cutting Edge 25(1), Jan. 2018], we expressed reluctance to implement the new system for the impending Manual Celastraceae treatment because two spp., Maytenus grisea Lundell and M. guyanensis Klotzsch ex Reissek. remained unplaced. This note resolves that situation for M. guyanensis. Only.

Blanco, M. A. & J. E. Jiménez. 2019. Duckweeds (Araceae: Lemnoideae) growing on wet, vertical rocks behind a waterfall in Costa Rica, with a new country record of Wolffiella oblonga (Phil.) Hegelm. Adansonia sér. 3, 41: 193–200.

A thriving duckweed community was discovered by the authors on the rock walls of a natural grotto (since partially collapsed) behind a well known waterfall, near Bagaces in Prov. Guanacaste. The authors claim (and we have no cause to argue) that this is "the only place in the world where duckweeds have been documented growing on wet rocks behind a waterfall." Who would even think (or dare!) to look in such a place? Indeed, we have been to the very site (in the so-called "Comelco" region), many years ago, and did not chance to encounter this population. This is also the first report of the genus Wolffiella "growing in a hygropetric [new term for us!] manner"; among duckweeds, only Lemna spp. had been found previously in that situation. Wolffiella oblonga, widespread in the Neotropics and previously recorded from both Nicaragua and Panama, becomes the second Costa Rican representative of its genus, after W. welwitschii (Hegelm.) Monod. Sparingly intermixed with W. oblonga in this population were Lemna aequinoctialis Welw. and (at an unusually low elevation) L. valdiviana Phil. Color photos depict the habitat, plus plants of L. valdiviana and W. oblonga.

——, K. M. Neubig, L. Endara, K. Silvera, I. E. Molgo & B. S. Carlsward. 2019. Obituary. William Mark Whitten (1954–2019). Lankesteriana 19(2): I–IX.

Several other obituaries of orchid biologist Mark Whitten, who passed away on 11 April 2019 [not 12 April, as reported under "News and Notes" in The Cutting Edge 26(3), Jul. 2019], have been (or will soon be) published, but this one—by a group of his former students—will do very nicely (and in any case, the others are here cited!). Features several well-chosen photos, enumerations of the taxa described by and named for Whitten, and a comprehensive listing of his publications. We did not know Mark, and clearly that was our loss.

Bortolus, A., P. Adam, J. B. Adams, M. L. Ainouche, D. Ayres, M. D. Bertness, T. J. Bouma, J. F. Bruno, I. Caçador, J. T. Carlton, J. M. Castillo, C. S. B. Costa, A. J. Davy, L. Deegan, B. Duarte, E. Figueroa, J. Gerwein, A. J. Gray, E. D. Grosholz, S. D. Hacker, A. R. Hughes, E. Mateos-Naranjo, I. A. Mendelssohn, J. T. Morris, A. F. Muñoz-Rodríguez, F. J. J. Nieva, L. A. Levin, B. Li, W. Liu, S. C. Pennings, A. Pickart, S. Redondo-Gómez, D. M. Richardson, A. Salmon, E. Schwindt, B. R. Silliman, E. E. Sotka, C. Stace, M. Sytsma, S. Temmerman, R. E. Turner, I. Valiela, M. P. Weinstein & J. S. Weis. 2019. Supporting Spartina: interdisciplinary perspective shows Spartina as a distinct solid genus. Ecology 100(e02863): 1–6.

Couldn't they just have branded themselves as, say, the "Spartina Preservation Group (SPG)," and spared us all the pointless typing? It is rare that we review a paper in Ecology (maybe we were cowed by the numbers!), but this one should be required reading for taxonomists in ivory towers: other people can be impacted and provoked by our decisions! And this was predictable in the case of the genus Spartina, recently judged (on the basis of relatively weak support) to be nested within Sporobolus and consequently subordinated to the latter genus [see under "Peterson," this column, in The Cutting Edge 22(2), Apr. 2015]—despite the fact that the name Spartina has priority over Sporobolus and "dominates the literature" (quoting here from the perpetrators themselves). The myriad authors of the paper under review (among whom we recognize only Stace as a practicing taxonomist) present a strong case, noting that "some Spartina species are among the most studied plant species"; they reckon that, despite the large discrepancy in sp. totals, at least "eight times more papers per species" have been published on Spartina (with 17 spp.) than on Sporobolus sensu stricto (with ca. 200 spp.). The authors do not dispute the cladistic evidence (though it is not compelling!), bending over backward to "recognize the value of molecular work"; nor do they take particular issue (though they might well have!) with the proposed conservation of Sporobolus over Spartina [water under the bridge anyway, since that proposal has already been recommended; see under "Applequist," this column, in The Cutting Edge 23(3), Jul. 2016]. Rather, their proposed resolution hinges on ranking: they suggest that traditional Spartina, deemed monophyletic and retained as a subgenus in the molecular study, could just as well resume its role as a full-fledged genus, providing only that additional genera be segregated from the remainder of Sporobolus sensu lato. In their own words: "There is clearly no rationale so compelling to subsume the name Spartina as a subgenus that could rival the striking, global, iconic history and use of the name Spartina for over 200 years. The name Spartina rings as a signature name in ecology and biology as well as across a striking range of stakeholders and disciplines, inlcluding history, literature, and anthropology." And for the time being: as pointed out by the authors, the botanical community has so far ignored these proposed taxonomic and nomenclatural changes ["Web of Science shows no (zero) listings using Sporobolus alterniflorus in the title between 2015 and 2019, while Spartina alterniflora has 189 listings"], a trend that will likely continue. Hanging onto traditional Spartina need not be interpreted as defiance of IAPT, merely tacit acceptance of other segregate genera in Sporobolus sensu lato.

Carrascal Prasca, D., F. Herazo Vitola & T. B. Croat. 2019. A new variety of Anthurium clavigerum (Araceae) from Sucre Department, Colombia. Novon 27: 151–155.

Recognition of this new var. of Anthurium clavigerum Poepp. will require that populations of that sp. in Costa Rica (and everywhere else outside the region specified in the title) be qualified as belonging to the autonymc var. Moving right along...

Cedeño-Fonseca, M., J. M. Flores-Leitón, A. Quesada-Román & R. Flores. 2020. Inventario florístico en un bosque amenazado por la expansión agrícola en la reserva del Centro Turístico Los Chocuacos, Costa Rica/Floristic inventory in a forest threatened by agricultural expansion in the reserve of Centro Turístico Los Chocuacos, Costa Rica. Revista Ci. Amb. 54(1): 33–57.

The private reserve named in the title comprises a mere scrap (25 ha) of primary and secondary forest in "bosque húmedo," at 250–300 m elevation in the lower Valle de Coto Brus. A floristic inventory resulting from "random walks" about the property between November 2017 and March 2018 yielded a list (included here as an appendix) of 266 spp. of vascular plants, representing 75 families and 219 genera. The most diverse families at the site were Fabaceae (with 31 spp.), Araceae (17 spp.), and Malvaceae and Rubiaceae (14 spp. each). "Trees" and "terrestrial" predominated in the habit and habitat categories, respectively. However, all of these statistics were no doubt influenced by the expertise of the authors (e.g., the first being specialized on Araceae) and seasonality (November–March being mostly dry). In both of those regards, it is perhaps significant that taxa such as pteridophytes (one sp.!), Cyperaceae (three spp.), and Poaceae (five spp.) are suspiciously underrepresented on the list. One notable discovery emerged from this work: Eugenia veraguensis Rod. Flores & A. Ibáñez (Myrtaceae), a sp. previously believed endemic to central Panama, now vouchered for Costa Rica (Bosque húmedo, bosques primarios y secundarios y orillas de quebradas, 50–100 m; vert. Pac., Valle de Coto Brus. Fl. feb., mar.) by Cedeño 772 [or 773?] (USJ) and Cedeño 1349 (USJ); that one of the authors of this paper is also a co-author of the sp. name lends a great deal of credence to the identification! The authors also collected material of an Aristolochia (Aristolochiaceae) sp. "nueva para la ciencia que actualmente se está describiendo"; we assume this to be one of five such spp. currently included in the Manual Aristolochiaceae draft treatment under provisional names. Color photos of selected habitats and spp. (including the aforementioned Eugenia) are provided. Regarding the alleged publication date of this article, see our remarks under "Salazar," this column.

Dalström, S. & A. P. Karremans. 2019. Obituary. Carlyle A. Luer (1922–2019) - in memory of a legend, a teacher and a friend. Lankesteriana 19(3): IX–XVIII.

The authors' contributions to this paper are separate, that of Dalström—who knew and collaborated (mainly as an illustrator) with Luer for nearly 40 years—being the longer of the two. Carlyle Luer, a surgeon by training and profession, retired in the "mid nineteen-seventies" to become an orchid giant. Indefatigable and "not the easiest individual to work with," Luer would dedicate the rest of his life to traveling and collecting Orchidaceae all over the Neotropics, and in the process describe 5604 "taxa," 5568 of those (including 3271 new spp.) in subtribe Pleurothallidinae, on which (at the suggestion of Calaway H. Dodson) he specialized. A notoriously difficult taxon comprising mostly small and/or small-flowered plants, Pleurothallidinae included "only about 1650 accepted species" in 1975, when the self-taught Luer began to work on the group; in other words, he essentially tripled the sp. total singlehandedly! According to Karremans (and few will argue), these accomplishments made Carl Luer "the most prolific plant taxonomist of modern times." Inevitably, many taxa (including three genera) were also named in honor of Luer; these are enumerated at the end of Karreman's portion, together with a handful honoring Luer's wife Jane or their son George (and it should be noted that about one-third of the names ostensibly honoring Carl Luer use the epithet luerorum, suggesting that they actually were meant to honor two or more members of the Luer family). This particular epitaph leans heavily on personal recollections, omitting some critical biographical details (such as the exact date of Luer's birth and death). But there will be many others. Illustrated with numerous (mostly) color photos.

Feuillet, C. 2019. New combinations in Heliotropium for four New World species of Tournefortia (Heliotropiaceae, Boraginales). Phytoneuron 2019-63: 1–2.

Workers on the taxon we used to know as Boraginaceae have concluded (among many other things) that the genus Tournefortia must be dispensed with, the bulk of it to be submerged in Heliotropium [see, e.g., under "Boraginales Working Group," this column, in The Cutting Edge 23(4), Oct. 2016]. To that end, this author engages in a bit of bookkeeping by validating new combinations in Heliotropium for four of the affected spp. that still lack them. Two of these occur in Costa Rica: Tournefortia bicolor Sw., which becomes Heliotropium album (Splitg. ex de Vriese) Feuillet (the epithet bicolor being preoccupied in Heliotropium); and T. cuspidata Kunth, now H. cuspidatum (Kunth) Feuillet. Still, as this author himself points out, "only a few species of Tournefortia sect. Tournefortia have a name in Heliotropium"; he sheds no light on why he sealed the deal for just four such spp., ignoring all the rest. A reminder to our readers: for both practical and theoretical reasons, traditional Tournefortia will not be sundered in the soon-to-be-published Manual Boraginaceae treatment, nor will "Heliotropiaceae" rear its ugly head.

Fonseca, L. H. M. & L. G. Lohmann. 2019. An updated synopsis of Adenocalymma (Bignonieae, Bignoniaceae): new combinations, synonyms, and lectotypifications. Syst. Bot. 44: 893–912.

Although the genus Adenocalymma boasts (according to this paper) some 76 spp., just one occurs in Costa Rica; thus, there is not a whole lot here for us. All the same, it is worth noting that these authors advocate a broad circumscription of Adenocalymma apurense (Kunth) Sandwith—first espoused by the late Alwyn H. Gentry in his Flora of Panama Bignoniaceae treatment (1973)—that includes (among other things) A. inundatum Mart. ex DC. They fail to indicate (nay, they misrepresent) the fact that Gentry later, in his Flora de Veracruz account (1982), reversed his stance and embraced A. inundatum as a distinct sp. That view was maintained in the Flora costaricensis (2000) Bignoniaceae treatment (co-authored posthumously by Gentry), where the name Adenocalymma inundatum was applied to the sole Costa Rican member of the genus, and is perpetuated in the Manual draft treatment of the family. Moreover, a fairly recent paper [see under "Udulutsch," this column, in The Cutting Edge 20(4), Oct. 2013] with a title eerily similar to that of the paper under review also accepted A. inundatum; therefore, we see no compelling reason to change. Features synonymy, typology, nomenclatural and/or taxonomic notes (as warranted), and distribution summaries at all ranks, plus a rigorous description of the genus (but none for the spp.). There are no keys or indices, but three composite plates of color photos from life.

Godoy, S. M., J. F. M. Silva, P. M. Ruas, M. R. Ritter, G. B. N. Paula, E. M. D. Maffei, B. D. Góes & C. F. Ruas. 2019. Multilocus approach reveals a complex evolutionary history of the invasive mile-a-minute plant, Mikania micrantha (Asteraceae), in its natural habitat. Bot. J. Linn. Soc. 191: 188–215.

"Mile-a minute plant"? Who comes up with these things? This study documents "relatively high genetic diversity" in the widespread ruderal sp. Mikania micrantha Kunth, samples of which were resolved in two lineages "that do not show morphological differences" and are "without geographical signature." Seek and ye shall find! The authors reference the "non-monophyletic origin of M. micrantha," even though their only published cladograms depict a sisterly relationship for the aforementioned lineages. They wonder whether M. micrantha may represent "a cryptic species complex," but defer an answer to the usual "further studies."

Guiggi, A. & M. Mariotti. 2018. W. C. Wercklé exsiccata collection (Cactaceae) in Herbarium Mortolensis (HMGBH): study and revision. Haseltonia 25: 2–4.

This sounds intriguing, from a Costa Rican perspective, but actually it is not. Just "five exsiccata relating to three taxa [i.e., names] of Costa Rican epiphytic cacti" were involved in the "revision," and two of those names were never validly published (i.e., they are nonexistent, for our purposes). The third name, Rhipsalis wercklei A. Berger, has long been acknowledged as a synonym of R. micrantha (Kunth) DC. (as also indicated here). The specimens are depicted photographically.

Hazlett, M. E. 2019. In memoriam: Donald Lavern Hazlett (3 September 1949–5 January 2019). J. Bot. Res. Inst. Texas 13: 599–600.

An illustrated obituary, by his wife, of recently deceased botanist Don Hazlett [see under "News and Notes" in The Cutting Edge 26(2), Apr. 2019].

Janzen, P. 2019. Orchideenfunde - Eindrücke aus Costa Rica. Orchidee (Hamburg) 70: 380–389.

The saga of "a two-week trip in a rental car" in ecotourist-overrun Costa Rica is published in a "scientific" journal. Seriously? Anyway, we get a lot of decent color photos, including some of animals and one of seldom-seen Volcán Arenal. So much for that.

Karremans, A. P. 2019. To be, or not to be a Stelis. Lankesteriana 19: 281–333['343'].

Quite some time ago (and prior to the publication of the Manual Orchidaceae treatment), the distinctive genus Stelis was enlarged and adulterated [see The Cutting Edge 9(1): 9–10, Jan. 2002] with an admixture of various taxa that had previously been recognized as distinct genera (Salpistele) or included in Pleurothallis (e.g., Crocodeilanthe, Dracontia, and Physosiphon). As it turns out, that classification has not been embraced wholeheartedly by the orchidophile community, with many workers succumbing to the temptation to "cherry-pick" some of the more well-marked monophyletic clades in the assemblage (including Stelis sensu stricto) and retain those at generic rank. However, as emphasized by this author, that course of action "necessarily entails the recognition of many ill-defined genera that no user would be happy about," and "that will not be more accepted and better defined or recognizable than Stelis s.l." This is a familiar dilemma nowadays, faced, e.g., also by those who would seek to salvage Chamaesyce and Pedilanthus from Euphorbia sensu lato [see The Cutting Edge 10(1): 10, Jan. 2003], or Perovskia and Rosmarinus from Salvia sensu lato [see, e.g., under "Will," this column, in The Cutting Edge 25(2), Apr. 2018]. According to the broader view, Stelis boasts some 1243 spp., and is the largest genus in subtribe Pleurothallidinae. This paper seeks to reinforce that view with the formal implementation of a new infrageneric classification comprising nine subgenera (subg. Stelis, with 1030 spp., being by far the largest) and various sections. These are of no concern to us, nor do we appear to be impacted by any of the new combinations or nomina nova validated in the illustrated nomenclator that comprises the bulk of the paper. A new subgenus of Pleurothallis is also debuted, in a section on "Excluded taxa," where we also learn that several taxa persistently associated with Stelis [see, e.g., The Cutting Edge 9(1): 9–10, Jan. 2002] have been "consistently shown" by DNA data to belong in other genera; these taxa inlcude Pleurothallis sect. Alatae Luer (= Anathallis), P. subgen. Effusia Luer (= Pabstiella), and the genus Effusia (Luer) Luer (= Pleurothallis).

Curiously, the pagination of this article skips ten pages in the middle, p. 289 being followed immediately by p. 300. However, since it is the last article in the volume, no others are affected.

Luteyn, J. L. 2019. Typification in and contributions to a revision of Psammisia (Ericaceae: Vaccinieae). J. Bot. Res. Inst. Texas 13: 397–449.

This paper, by Manual Ericaceae co-author James L. Luteyn, is bursting at the seams with information and insights, but most of it pertains to taxa of South America, where Psammisia is best represented. The only item we could find that is relevant to our interests is the author's neotypification of Psammisia ulbrichiana Hoerold, the accepted name for one of just three spp. in the genus (out of ca. 70) that extend northward to Costa Rica.

Maas, P. J. M., L. Y. T. Westra, L. W. Chatrou, N. Verspagen, H. Rainer, N. A. Zamora & R. H. J. Erkens. 2019. Twelve new and exciting Annonaceae from the Neotropics. PhytoKeys 126: 25–69.

We were remiss in not reviewing this paper in our last issue, or even the one before that (it was published on 2 July). Perhaps we wanted to pretend that it didn't exist. Among the "exciting" novelties described by these authors are two that occur in Costa Rica: Guatteria rubiginosa N. Zamora & Maas (which also enters western Panama) and the endemic G. turrialbana N. Zamora & Maas. These constitute rather an embarrassment for us, because both involve Manual co-PI Nelson Zamora and were disdained (swept under the rug?) in a recent revision of Guatteria [see under "Maas," this column, in The Cutting Edge 23(1), Jan. 2016], as well as in the Manual draft treatment of Annonaceae—both co-authored by Nelson! That they are finally seeing the light of day, so late in the game, will, unfortunately, have few repercussions for the Manual: due to time constraints, neither Guatteria rubiginosa nor G. turrialbana will be keyed or treated formally, merely mentioned under G. talamancana N. Zamora & Maas and G. lucens Standl. (respectively)—the spp. with which they are compared in this paper. Here we also get (for each sp.) a full-page, composite plate of color photos from life.

Mandáková, T., J. Zozomová-Lihová, H. Kudoh, Y. Zhao, M. A. Lysak & K. Marhold. 2019. The story of promiscuous crucifers: origin and genome evolution of an invasive species, Cardamine occulta (Brassicaceae), and its relatives. Ann. Bot. (Oxford) 124: 209–220.

According to its authors, this study "provides unequivocal evidence" that the widely adventive east Asian Cardamine occulta Hornem. (which, we now know, occurs in Costa Rica) is an allooctoploid derived from two other east Asian spp., C. kokaiensis Yahara et al. and C. scutata Thunb. We remain largely in the dark with respect to its overall geographic distribution [see under "Šlenker," this column, in The Cutting Edge 25(4), Oct. 2018].

Matos, F. B., P. H. Labiak & R. C. Moran. 2019. The Decursivum Group of Elaphoglossum sect. Polytrichia (Dryopteridaceae): non-subulate species nested in an otherwise subulate-scaled clade. Ann. Missouri Bot. Gard. 104: 400–428.

The big deal here is that seven spp. previously classed in the autonymic section of Elaphoglossum are revealed (using molecular techniques) as nested within E. sect. Polytrichia Christ. While we certainly do not care about anything on that order, we do glean some new information from the authors' taxonomic treatment of these previously misplaced spp. (dubbed the "Decursivum Group"). Just three of the seven occur in Costa Rica: Elaphoglossum albomarginatum A. R. Sm., E. decursivum Mickel, and E. lonchophyllum (Fée) T. Moore, only the last of which was attributed to the country in Flora mesoamericana Vol. 1 (1995). Costa Rican (and Panamanian) material of E. albomarginatum was referred in the last-mentioned work to E. pallidiforme Mickel, there qualified as "relacionada con E. albomarginatum" (and here consigned, without comment, to synonymy thereunder). Elaphoglossum decursivum (type from Oaxaca) was not mentioned at all in Flora mesoamericana, although dozens of specimens are cited in this paper from southern Mexico to western Colombia and Hispaniola; we can only assume that most of those had been mis- or unidentified. The taxonomic treatment features brief, formal descriptions of the Decursivum Group and each of its spp., a dichotomous and indented key to spp., synonymy and typology, distribution summaries and maps, discussions, comprehensive specimen cigtations, and an index to exsiccatae. The introductory portion is mainly devoted to taxonomic history, the molecular phylogenetic analysis, and morphology. All seven spp. are illustrated with composite line drawings and/or color photos.

Moroni, P. & N. O'Leary. 2019. Insights into the systematics of tribe Duranteae (Verbenaceae): a taxonomic revision of the New World genus Bouchea. Ann. Missouri Bot. Gard. 104: 355–399.

It seems to us that the first half of the title obscures rather than highlights the true nature of this contribution, and would better have been omitted: this is a revision of Bouchea, plain and simple. Why muddy the waters? With that said, there are a few changes here for us. First, this revision recognizes a total of 13 spp. in Bouchea, vs. just nine as according to the Manual Verbenaceae treatment (2015) by Ricardo Rueda (HULE). Second, and more significantly: although these authors concur with Ricardo in attributing just one Bouchea sp. to Costa Rica, they use a different name for it, Bouchea nelsonii Grenzeb. In the Manual, that name had been listed in synonymy under B. prismatica (L.) Kuntze, accepted there as the correct name for Costa Rican material; however, these authors maintain that the "two taxa can be distinguished with a high degree of accuracy," based (according to their key) on pubescence, inflorescence density, and the length of the mericarp beak. Bouchea nelsonii ranges only from southern Mexico to Costa Rica. Somewhat confusingly, B. prismatica sensu stricto is also attributed to "Central America," and mapped even in Costa Rica, though no Costa Rican specimens are cited (and two appendices indicating the distribution of spp. by country omit Costa Rica from the range of B. prismatica, or vice versa). Includes synonymy, typology, technical descriptions, distribution summaries and maps, and discussions or notes at all ranks, a dichotomous and indented key to spp., phenology summaries, a list of excluded taxa (two), and indices to synonyms and exsiccatae, as well as the aforementioned appendices. The introductory part is devoted to taxonomic history and morphology. Two new combinations are validated and three lectotypes designated, but none of that concerns us. Each sp. is illustrated with a composite line drawing, and a few (not B. nelsonii) also with color photos from life.

Moura, T., R. P. Campos & R. E. Gereau. 2019. Typification of the Linnaean name Atropa arborescens (Solanaceae). Novon 27: 235–237.

Atropa arborescens L. is the basionym of Acnistus arborescens (L.) Schltdl., the name used in Manual Vol. 8 (and just about every other botanical work up to now) for a sp. that is well known throughout its geographic range. As noted by these authors, the monospecific genus Acnistus has been shown to be nested within the larger (ca. 25 spp.), otherwise South American Iochroma; indeed, apparently unbeknownst to these authors, an abortive attempt was recently made [see under "Shaw," this column, in The Cutting Edge 25(4), Oct. 2018] to transfer Acnistus arborescens to Iochroma (by an author who was ignorant of its basionym!). Here, Atropa arborescens is lectotypified on a Plumier plate, the only element cited in the Linnaean protologue. Given the limited utility of plates, in conjunction with the authors' agenda to "clarify the identity of A. arborescens," we wonder why they did not also designate a good, modern collection as epitype.

Ossenbach, C. 2019. Obituary. Robert Dressler (1927–2019)–a botanist for all seasons. Lankesteriana 19(3): I–VIII.

This excellent contribution, written by a respected historiographer who knew his subject personally, is among the first of many obituaries that will inevitably be published on behalf of the recently deceased orchid mastermind [see under "News and Notes" in our last issue]. The author skillfully combines basic historical data with personal reminiscences, and those of others who knew Dressler well, spicing the mix with photos (mostly color) taken or at least contributed by Dressler's wife Kerry. Fittingly portrayed on the cover of this magazine issue, Robert Dressler moved (with Kerry) to Costa Rica in March 2005 to assume the role of director of scientific research at the Jardín Botánico Lankester. There he would remain until his death. As indicated in our review of a recently published biography [see under "Pridgeon," this column, in The Cutting Edge 23(3), Jul. 2016] from which the title of this paper was "borrowed," Bob had been born in southern Missouri, graduated from both high school and college in the Los Angeles area, and obtained his Ph.D. at Harvard University under Reed Rollins (in our last issue, we stated mistakenly that he had earned his Ph.D. at the University of Southern California). What followed is mostly common knowledge, but those needing to refresh their memories should read this epitaph (and/or any of the others soon to appear).

Page, C. N. 2019. New and maintained genera in the taxonomic alliance of Prumnopitys s.l. (Podocarpaceae), and circumscription of a new genus: Pectinopitys. New Zealand J. Bot. 57: 137–153.

Alas, Podocarpus of yore continues to decay into ever-smaller and more ill-defined segregate "genera." A common trend these days! In this paper, Prumnopitys in the sense of Manual Vol. 2 (2003)—itself a comparatively recent Podocarpus segregate, at least in our minds—is fragmented into three parts, largely on the basis of "first-hand observations of predominately [sic] vegetative features as the main essential 'key' characters." Just three spp. (of the nine attributed to Prumnopitys in the Manual, or the 12 accepted here for the titular "alliance") remain in Prumnopitys s. str., and the endemic Costa Rican Prumnopitys standleyi (J. Buccholz & N. E. Gray) de Laub. is not among them; rather, it joins five putative congeners in the new genus of the title, becoming Pectinopitys standleyi (J. Buchholz & N. E. Gray) C. N. Page (comb. nov.). Not surprisingly (to anyone who knows the last-mentioned taxon), Pectinopitys is characterized by its "strongly regimented arrangement of leaves into one plane, seen on most lateral shoots." Unfortunately, although a great deal of morphological information is presented in this paper, there is no key to genera, nor are formal genus descriptions provided (except in the case of the new genus). Lip service is paid to cytogenetic and molecular "backing," but neither field appears to offer convincing support for the new classification (and tellingly, no cladograms are reproduced here). So we remain skeptical (hey, Missouri is the "Show-me State"!). Several other nomenclatural innovations (e.g., at subgeneric and infraspecific ranks) are of no interest to us.

Queiroz, R. T., T. M. Moura, R. E. Gereau, G. P. Lewis & A. M. G. Azevedo-Tozzi. 2019. Resolving nomenclatural ambiguity in South American Tephrosia (Leguminosae, Papilionoideae, Millettieae), including the description of a new species. Austral. Syst. Bot. 32: 555–563.

Germane tangentially to Costa Rican floristics are the lecto- or neotypifications of two synonyms of Tephrosia cinerea (L.) Pers. and one of T. vogelii Hook. f. (the latter merely mentioned in the Manual). Also of passing interest is a dichotomous key to the spp. of Tephrosia in South America. We cannot help but wonder why any of this was deemed appropriate (by either the authors or the editors) for an Australian journal.

Salazar, H. 2020. Alberto Manuel Brenes: el primer botánco costarricense/Alberto Manuel Brenes: the first Costa Rican botanist. Revista Ci. Amb. 54(1): 190–199.

This article is dated "2020," and even the Web site (which we just checked) states that it was published on 1 Jan 2020, but we are writing this review on 24 Dec 2019 and have had the paper in hand for a week already. So something is fishy. This latest biography of Alberto Brenes is at least the third to appear during the new millennium [see, e.g., The Cutting Edge 10(2): 12, Apr. 2003]. We doubt there is much new to be said about the man, but are not about to undertake a meticulous comparison of these various works. As this author notes in a closing statement, "hay personas que nunca mueren." Readers will learn here, among other things, that holotypes of all the orchid spp. (146!) described as new by Rudolf Schlechter based on Brenes's collections were destroyed in Berlin during World War II; that Brenes "fue un entusiasta recolector de hongos," in addition to vascular plants; and that Brenes played key roles in several major botanical "expeditions" to Costa Rica, including those of Walter Kupper and Otto Porsch. His membership in the "hall of fame" of Costa Rican biologists is surely guaranteed for all eternity.

Saldivia, P., O. M. Vargas, D. A. Orlovich & J. M. Lord. 2019. Nomenclatural priority of the genus Linochilus over Piofontia (Asteraceae: Astereae). Phytotaxa 424: 158–166.

We had received some advance warning of this development, from a a source apparently unconnected with this team. It boils down to this: the genus name Piofontia Cuatrec. (1943) was applied by the second author of this paper to some 60 spp. previously included in Diplostephium [see under "Vargas," this column, in The Cutting Edge 25(4), Oct. 2018], but one of those spp. happens to be the generic type of Linochilus Benth. (1845), which has priority by nearly a century. Because "the incorrect application of Piofontia is still very recent..., [and] has not yet been in use," the authors consider that "the quick publication of the present article" (validating 61 new combinations in Linochilus) "will help to avoid further confusion..." It would figure that Linochilus costaricensis (S. F. Blake) Saldivia & O. M. Vargas (comb. nov.) must become the correct name for one of the two affected Costa Rican spp., while the situation for the other (treated as "Diplostephium sp. A" in Flora mesoamericana) is ambiguous; however, we have it on good authority that further changes are in order for both of our spp.

Salmaki, Y., G. Heubl & M. Weigend. 2019. Towards a new classification of tribe Stachydeae (Lamiaceae): naming clades using molecular evidence. Bot. J. Linn. Soc. 190: 345–358.

The titular tribe is confirmed as monophyletic, but beyond that there are problems: the widespread and well-known genus Stachys was recovered as paraphyletic (was there ever a doubt?) with respect to 10 (by our count) other established genera. The authors ruminate over two familiar options (see also under "Bortolus" and "Karremans," this column): to split the entire assemblage into smaller monophyletic genera (12 in all) "without clear morphological synapomorphies," or to lump all the nested genera (some "widely recognized and well-defined") into a broadly defined Stachys. At this stage they take no stand, preferring to indulge in "naming clades" (an activity that leaves us cold). From a purely New World perspective, this one is a no-brainer: go for the big lump! All of the nested genera are restricted to the Old World (or Oceania), none is familiar to us, and all but three are mono- or oligospecific. Sideritidis, by far the largest (with ca. 140 spp.) and the only one with a name we recognize, is already flagged in The plant-book (2017) as "~ Stachys." So our vested interest falls squarely on the side of lumping, although Old World workers would likely take a different view; moreover, it is conceivable (though not at all clear from the information presented in this paper) that our five Costa Rican spp. (none of which was included in this study) would remain in Stachys even under the splitting scenario (in which case, we wouldn't care what transpires among the other clades!).

Santamaría-Aguilar, D., J. E. Jiménez & R. Aguilar. 2019. Otoba vespertilio (Myristicaceae), una especie nueva de Mesoamérica. Brittonia 71: 369–380.

The new sp., Otoba vespertilio D. Santam. & J. E. Jiménez (its epithet alluding to presumed fruit dispersal by bats), is compared most closely with the Ecuadorean O. cyclobasis T. S. Jaram. & Balslev, though in Costa Rica it had been confused principally with O. acuminata (Standl.) A. H. Gentry. The numerous differences between those three superficially similar spp. are conveniently tabularized. For Costa Rica, the distribution and phenology of O. acuminata may now be revised as follows (in Manualese): "Bosque muy húmedo, 50–900 m; vert. Carib. Cords. Central y de Talamanca. Fl. mar., jun.–set."; and those of the new sp. summarized as: "Bosque húmedo y muy húmedo, 0–200 m; vert. Carib., Baja Talamanca. Fl. feb., abr.–jun." Both spp. also range into Panama. Includes distribution maps, a dichotomous key to all four spp. of Otoba occurring in Mesoamerica, comprehensive specimen citations for both O. acuminata and O. vespertilio (facilitating the foregoing summaries), a composite drawing of the latter, and color photos from life of all three Costa Rican representatives of the genus. Full disclosure: material of the new sp. had also been misidentified (including by two of your editors!) as Otoba novogranatensis Moldenke, and even Virola multiflora (Standl.) A. C. Sm.! Fortunately, however, the Manual vouchers for all the spp. mentioned in this paragraph were determined correctly.

Taylor, C. M. 2019. Rubiacearum americanarum magna hama pars XLV: more new species and taxonomic changes in Palicourea (Rubiaceae, Palicoureeae) and Psychotria subg. Heteropsychotria. Novon 27: 165–195.

All of the 32 assorted spp. dealt with in this paper (which mainly continues the long-gestating theme of transferring taxa from Psychotria to Palicourea) are extralimital to our interests, i.e., restricted to areas north or south of Costa Rica. However, one of them, Palicourea galeottiana M. Martens, was reported from Costa Rica [as Psychotria galeottiana (M. Martens) C. M. Taylor & Lorence] in both the Flora mesoamericana (2012) and Manual (2014) Rubiaceae treatments, each involving the author of this article. The same Costa Rican voucher was cited in both works: Paul 1039, a fruiting collection from 1090 m elevation on the Pacific slope of the Cordillera de Tilarán in the Monteverde region. Now, presumably with respect to the Paul collection (though it is not cited here), the author states the following: "that specimen is in poor condition and its identity cannot be confirmed, but further study shows that it does not belong to the Palicourea galeottiana group." So it would appear that we should expunge Palicourea (formerly Psychotria) galeottiana (otherwise ranging from southern Mexico to central Nicaragua) from the Costa Rican flora (although the aforementioned voucher remains so determined in TROPICOS!).

Tucker, G. C. & K. N. Gandhi. 2019. Nomenclatural notes on Cyperus odoratus and related species (Cyperaceae). Harvard Pap. Bot. 24: 349–357.

Another of those contributions that raises more questions than it answers, at least for us. Here the authors (among other things) resurrect the names Cyperus ferax Rich. and C. acicularis (Schrad. ex Nees) Steud. from synonymy under C. odoratus L., where they had been cited in Manual Vol. 2 (and elsewhere), and apply them to distinct spp. It turns out, however, that C. acicularis, with the authorities just indicated, is an illegimate later homonym, here replaced by the nomen novum Cyperus strongii G. C. Tucker & Gandhi. Now, the newly liberated C. ferax apparently does not occur in Central America, according to the specimens cited here. Cyperus strongii does, however, based on one specimen each cited from Belize, Guatemala, and Costa Rica. The Costa Rican record of C. strongii is from Isla del Coco (Trusty & Kesler 468, FTG), raising the following questions: is C. strongii restricted, in Costa Rica, to Isla del Coco, or is it more widespread in the country? And if the latter, does the real Cyperus odoratus occur in Costa Rica at all, or do we only have C. strongii? We gain no insights from this particular paper, which cites no specimens of C. odoratus, so must await some future installment. One final observation: if the real Cyperus odoratus does indeed occur in Costa Rica, we expect that our material would have to be qualified as belonging to the autonymic var., as these authors recognize (in their dichotomous key to Cyperus subg. Diclidium) another var. restricted (as far as we can tell from the information available to us) to Texas.

Turner, I. M. 2019. Warszewiczia and sixteen other names in the Rubiaceae authored by J. C. Klotzsch: when and where were they published? Willdenowia 49: 151–154.

The author of this paper has determined that 17 names in Rubiaceae commonly cited as having been published by Johann Friedrich Klotzsch (whence the "C."?) in an issue of Flora dated December 1853 were in fact published at least two months earlier (by the same author) in a more obscure German journal. Just one protologue citation in the Manual Rubiaceae treatment (2014) is affected: that of Warszewiczia coccinea (Vahl) Klotzsch, to be changed from "Flora 36: 716. 1853" to "Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1853: 497. 1853." Also, a lectotype is newly designated for the basionym of W. coccinea (and there are seven other new type designations for names that do not concern us).

Vasques, D. T., A. Ebihara, R. Y. Hirai, J. Prado & I. Motomi. 2019. Phylogeny of Hymenophyllum subg. Mecodium (Hymenophyllaceae), with special focus on the diversity of the Hymenophyllum polyanthos species complex. Pl. Syst. Evol. 305: 811–825.

Hymenophyllum polyanthos (Sw.) Sw., one of the commonest spp. of its genus in Costa Rica, is generally specified (as in Flora mesoamericana Vol. 1) as essentially pantropical. This study, however, portrays H. polyanthos as polyphyletic, comprising at least eight independent lineages. Several samples from Bolivia and Costa Rica, hypothesized as "representative of [the] type lineage," are "closely related to" several other neotropical spp., including H. myriocarpum Hook., H. undulatum (Sw.) Sw., and (presumably) H. fendlerianum J. W. Sturm. Based on this, the authors "suggest that the type lineage for H. polyanthos may be restricted to the Neotropics," hedging their bet only because Antillean material was (short-sightedly) not studied (the basionym of H. polyanthos is typified by a Jamaican specimen). One other separate lineage of H. polyanthos is represented in the New World, though apparently only in South America (e.g., Chile and Brazil). Bottom line for us: if all of this plays out as the authors seem to believe it will, Costa Rican material of Hymenophyllum polyanthos will retain that name.

Wolf, P. G., C. A. Rowe, S. P. Kinosian, J. P. Der, P. J. Lockhart, L. D. Shepherd, P. A. McLenachan & J. A. Thomson. 2019. Worldwide relationships in the fern genus Pteridium (bracken) based on nuclear genome markers. Amer. J. Bot. 106: 1365–1376.

We have been struggling to come to grips with the latest taxonomic developments affecting the genus Pteridium (Dennstaedtiaceae), of which three spp. were recognized in Robbin Moran's (1995) Flora mesoamericana treatment: Pteridium arachnoideum (Kaulf.) Maxon, P. caudatum (L.) Maxon, and P. feei (W. Schaffn. ex Fée) Faull (with only the first two attributed to Costa Rica). This latest contribution follows rather closely on the heels of two others from different teams [see, e.g., under "Schwartsburd," this column, in The Cutting Edge 25(1), Jan. 2018], and agrees with those in recognizing two widespread diploid spp.: Pteridium aquilinum (L.) Kuhn, "predominant in the northern hemisphere," and P. esculentum (G. Forst.) Cockayne, "predominantly southern hemisphere in distribution." In addition, "two allotetraploid taxa resulting from hybridization" of the two aforementioned spp. are recognized: Pteridium caudatum (mainly from Central and South America) and P. semihastatum (Wall. ex J. Agardh) S. B. Andrews (from southeastern Asia and northern Australia). So we are beginning to arrive at an understanding of this new taxonomy, and a sort of consensus would appear to be emerging. Of the three Flora mesoamericana spp., only P. caudatum survives at that rank, and is the only taxon explicitly attributed to Costa Rica (via a map and a specimen citation) in this paper (which is by no means a taxonomic revision). Pteridium arachnoideum and P. feei are both apparently maintained at the rank of subsp., of P. esculentum and P. aquilinum, respectively; however, neither can be confirmed as occurring in Costa Rica from this source. In conclusion: the only clear deviations of this new taxonomy from that of Flora mesoamericana are the demotions of Pteridium arachnoideum and P. feei to subspecific rank.



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