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The Cutting Edge
Volume XXVIII, Number 3, July 2021
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Avalos, G., E. Chacón-Madrigal & L. G. Artavia-Rodríguez. 2021. Invasive plants of Costa Rica current status and research opportunities. Pp. 57–76 in, T. Pullaiah & M. R. Ielmini (eds.), Invasive alien species: observations and issues from around the world, Vol. 4: issues and invasions in the Americas and the Caribbean. John Wiley & Sons Ltd., Hoboken, NJ.
The biggest eye-opener comes on the first page: “The service industry and tourism-associated businesses are responsible for 70% of the country’s income, whereas the agricultural sector represents only about 5–11% of the gross domestic product.” Godfrey Daniels! We knew that tourism—and specifically, ecotourism—had become increasingly important in Costa Rica, but whither coffee and bananas? Whatever, the case, the inevitable “focus on biodiversity protection” in the name of ecotourism may provide a modicum of hope for the country’s natural habitats, even if accompanied by an incursion of human hordes (think Monteverde). But we have gone off script: this chapter seeks “to analyze the status of the vascular plant species introduced in Costa Rica and select cases in which these have the potential to become invasive.” It is important to note that, according to the terminology adopted in this paper, only a small percentage of introduced spp. actually qualify as invasive; according to cited sources, introduced plant spp. account for 9.7% of the Costa Rican flora (1048 spp.), but only 0.4% of those (47 spp.) have been deemed invasive (according to a website and databases that “lost support…went offline and are not up to date”). Invasive spp. are said to “represent the fifth largest cause of biodiversity and ecosystem function loss across the world.” In theory, “Costa Rican legislation provides ample opportunity to manage potential invasive species”; however, in practice, “the use, transportation, handling, and propagation of introduced plants is carried out with minimal governmental control.” Sound familiar? The authors consider nine “select cases” of plant spp. with the potential to become invasive in Costa Rica; as with a “greatest hits” album, every fan will gripe about the inclusion of some elements and the omission of others. For what it is worth, here are the the author’s selections: Artocarpus altilis (Parkinson) Fosberg (Moraceae), Cocos nucifera L. (Arecacae), Heliotropium indicum L. (Boraginaceae), Musa velutina H. Wendl. & Drude (Musaceae), Oeceoclades maculata (Lindl.) Lindl. (Orchidaceae), Spathodea campanulata P. Beauv. (Bignoniaceae), Syzygium jambos (L.) Alston (Myrtaceae), Terminalia catappa L. (Combretaceae), and Ulex europaeus L. (Fabaceae). We would wholeheartedly endorse a few of those (Musa, Oeceoclades), and question a few others (Artocarpus, Terminalia), but could do no better (and we even learned a few insidious details we didn’t know about some spp. on the list). The authors also address “Habitats requiring special attention,” viz., lowland tropical rainforests, wetlands, humid savannas, páramo, and islands (Cocos, in particular). The roles of ecological restoration and living plant collections in accelerating the escape of introduced (and potentially invasive) plant spp. are briefly considered. The authors conclude that the lack of information on and monitoring of invasive spp. are “the main obstacles to the development of management and eradication policies” by the government and NGO offices encharged with doing so, which “are faced with scarce resources, a lack of well-trained personnel, and weak environmental legislation at the regional level.” Again we must ask: sound familiar?
Biral, L. 2020. (2784) Proposal to reject the name Maytenus tovarensis (Celastraceae). Taxon 69: 1366–1367.
Maytenus tovarensis Radlk. is a "long-forgotten name," and its type was apparently destroyed at B during the last world war. Its precise identity is deemed "currently unresolvable" by this author; however, he considers it "likely that M. tovarensis is in fact synonymous with" Celastrus liebmannii Standl., the name currently in use for a widespread sp. (and the only one of its genus in Costa Rica). Pittier seemed to think so, which is good enough for us! Anticipating the negative consequences of "any rediscovery of type material or neotypification," the author proposes this pre-emptive resolution. In our opinion, proposals of this sort should never be contemplated (let alone published) until typification has been effected.
——. 2021. A new combination and a new synonym in Haydenoxylon (Celastraceae). Novon 29: 74–76.
A sp. not even described until 1997 (or so we thought!) now undergoes its third name-change since that date—which is perhaps a record for a sp. that is neither a pteridophyte nor an orchid! Gymnosporia haberiana Hammel was its first christening [see The Cutting Edge 4(3): 6, Jul. 1997], but neither element of that binomial would be retained. First to go was the genus name, when Gymnosporia was shown to be polyphyletic and its New World members were transferred to a new segregate genus; our guy thus became Haydenia haberiana (Hammel) M. P. Simmons [see under "McKenna," this column, in The Cutting Edge 19(1), Jan. 2012]. But that didn't last long: shortly thereafter, it was discovered that the genus name Haydenia had already been used (for a fossil fern), and Haydenoxylon haberianum (Hammel) M. P. Simmons was born [see under "Simmons," this column, in The Cutting Edge 21(4), Oct. 2014]. The last-mentioned name was the one used in the recently published (2020) Manual Celastraceae treatment, co-authored by co-PI Barry Hammel (full circle!). But let's backtrack a bit: in both the Manual account of Hydenoxylon haberianum and the protologue of its basionym, Barry had included material from Veracruz, Mexico, in his concept of that sp. It now turns out that, at least according to the paper under review, the same sp. had already been described from Veracruz, in 1985, as Euonymus calzadae Lundell! Hence the new combination Haydenoxylon calzadae (Lundell) Biral, to replace all that came before; and if we count the combination Maytenus calzadae (Lundell) J. S. Ma (2001), this sp. has actually rocked six different names since 1985! By the way: although he regrets never having come to the realization himself (and was never informed of the impending new combination), Barry accepts this new development, and has already implemented the change in the online Manual treatment of Celastraceae.
Bogarín, D. & F. Pupulin. 2021. The orchid flora of Barra Honda National Park, Nicoya, Guanacaste, Costa Rica. Harvard Pap. Bot. 26: 7–99.
These same authors had already published a more informal report on the orchids of Parque Nacional Barra Honda [see The Cutting Edge 14(4): 4–5, Oct. 2007], accounting for 26 spp. in 23 genera in this relatively arid lowland locale, but without citing vouchers. This new, more rigorous treatment ups the ante to 36 spp. in 29 genera, and in addition to specimen citations, provides a dichotomous (though non-indented) key to spp., synonymy and typology, technical descriptions and distribution summaries (for both genera and spp.), and discussions. Two spp. are described as new, both endemic (as far as is known) to Barra Honda: Pelexia barrahondaensis Bogarín & Pupulin [compared most closely to the Honduran endemic P. gutturosa (Rchb. f.) Garay] and Sarcoglottis calcicola Bogarín & Pupulin (compared to S. hunteriana Schltr.). In addition, the new combination Specklinia panamensis (Schltr.) Bogarín & Pupulin is validated, based on Pleurothallis panamensis Schltr., a name (not mentioned in the Manual) that had been deemed synonymous with Specklinia microphylla (A. Rich. & Galeotti) Pridgeon & M. W. Chase (i.e., Pleurothallis microphylla A. Rich. & Galeotti of the Manual). According to these authors, Specklinia panamensis “ranges from Costa Rica (Guanacaste, Alajuela, Puntarenas) up to Panama (Chiriquí),” while specimens answering to the description of the real S. microphylla “have been collected in the Caribbean lowlands of Costa Rica” (which corresponds to the Costa Rican distribution for the sp. indicated in the Manual). Many range extensions are specified in this paper, including minor in-country habitat, elevational, and/or geographic extensions too numerous to mention. Most significant is Tropidia polystachya (Sw.) Ames, ostensibly a new genus for Costa Rica, although these authors reported it themselves in a previous paper [see the third entry under “Pupulin,” this column, in The Cutting Edge 20(2), Apr. 2013], and also alluded to it (as an unidentified “Palmorchis”) in their first Barra Honda paper. But here we finally get an actual voucher citation! Four other records deserve mention: Cohniella brachyphylla (Lindl.) Cetzal & Carnevali, accepted tentatively for material treated as Oncidium cebolleta (Jacq.) Sw. in the Manual (see, e.g., under “Carnevali Fernández-Concha” in The Cutting Edge 17(4), Oct. 2010]; Encyclia macrochila (Hook.) Neumann (we would question the extraparenthetical authority), corresponding, at least in part, to E. cordigera (Kunth) Dressler in the sense of the Manual; Lophiaris oerstedii (Rchb. f.) R. Jiménez, Carnevali & Dressler, accepted somewhat tentatively for Costa Rican material dubbed Oncidium carthagenense (Jacq.) Sw. in the Manual, instead of L. crispiflora (Schltr.) Balam & Cetzal, the name suggested by previous authors [see, e.g., under “Cetzal-Ix,” this column, in The Cutting Edge 19(3), Jul. 2012]; and Sarcoglottis acaulis (Sm.) Schltr., a prior Costa Rican record for which had been rejected (in rather cavalier fashion) in the Manual.
The lengthy introductory portion of this paper, well illustrated with color photos and maps, discusses the abiotic features of the study site, biodiversity and vegetation, and orchid distribution, biogeography, and floristic composition at Barra Honda. A surprising 27.8% of the spp. at the site are terrestrial, and 80% occur (unsurprisingly) in evergreen forest. Each sp. is superbly illustrated, in one form or another. The amount of care that goes into this type of work cannot be overestimated.
Boom, B. M. 2020. Scott Alan Mori (1941–2020): an appreciation. Bot. Rev. (Lancaster) 86: 149–179.
This is a detailed and abundantly illustrated biography and appreciation of the late Scott Mori by one of his long-time colleagues at NY. Having grown up in a small town in southern Wisconsin, Scott's life trajectory was changed forever when he took a Phytogeography course from Hugh Iltis at the University of Wisconsin. During his first trip to Costa Rica in 1967, Scott "spotted a group of trees in the Lecythidaceae," and the rest is history. Enjoy the read!
Bouman, R. W., P. J. A. Keßler, I. R. H. Telford, J. J. Bruhl, J. S. Strijk, R. M. K. Saunders & P. C. van Welzen. 2021. Molecular phylogenetics of Phyllanthus sensu lato (Phyllanthaceae): towards coherent monophyletic taxa. Taxon 70: 72–98.
Okay, we are dyed-in-the-wool lumpers, but Phyllanthus sensu lato (i.e., in the sense of the Manual) is hard to swallow even for us. How can a diminutive, herbaceous weed (e.g., Phyllanthus tenellus Roxb.) and a canopy tree of primary forest (P. skutchii Standl.) possibly be accommodated in the same genus? Well, there is everything in between, but even so… This molecular study actually brings nothing new to the table, in terms of mandating the dissolution of Phyllanthus sensu lato. Sure, it is paraphyletic with respect to Breynia (including Sauropus), Glochidion, and Synostemon, but we already knew that; authors disposed to lump have simply subsumed those genera under Phyllanthus, no big deal. But these authors are hell-bent to split, and this paper is the horse that properly precedes the cart (i.e., a new classification that "is forthcoming"). It will be interesting to see what the authors come up with, especially with respect to circumscribing splinter genera that are easily recognizable, on morphological bases, by non-specialists. They do provide a crude indication of what is planned, and combining that information with our interpretation of the cladograms in this paper permits some speculation regarding the new generic affiliations of those Costa Rican representatives of Phyllanthus sensu lato that were sampled for this study (P. skutchii, unfortunately, is not one of them). These analyses revealed nine major clades in Phyllanthus sensu lato, designated A–I (and corresponding rather poorly to the prevailing subgeneric classification); the letters assigned to the following prospective genera are those of their corresponding clades. We believe that Genus D will include Phyllanthus amarus Schumach. & Thonn., P. stipulatus (Raf.) G. L. Webster, and P. tenellus (and perhaps P. caribaeus Urb.?); Genus E will include Phyllanthus acuminatus Vahl, P. anisolobus Müll. Arg., P. caroliniensis Walter, P. graveolens Kunth, P. niruri L., and P. salviifolius Kunth (and perhaps P. hyssopifolioides Kunth?); Genus F will include Phyllanthus acidus (L.) Skeels (mentioned in the Manual as cultivated in Costa Rica) and P. elsiae Urb. (mentioned as to be expected); and Genus G will include Phyllanthus urinaria L. The authors claim that all of their projected segregate genera already have names, but they don't reveal them, and we don't have the time to figure them out. Breynia (including Sauropus) will definitely ride again, however (it was included in Phyllanthus for the purposes of the Manual). So bring it on! We can't wait to see how close we came. If we are right, we wonder whether a new system that classes such grossly similar spp. as Phyllanthus amarus, P. niruri, and P. urinaria in different genera will be accepted as an improvement over one that includes a range of very different-looking spp. in a single genus.
Breslin, P. B., M. F. Wojciechowski & L. C. Majure. 2021. Molecular phylogeny of the Mammilloid clade (Cactaceae) resolves the monophyly of Mammillaria. Taxon 70: 308–323.
Genus-level classification in the Cactaceae has long been notoriously unstable, and the so-called “Mammilloid clade” is no exception. Even non-specialist botanists will recognize many of the names of genera that have been variously combined with, or segregated from, Mammillaria over the years: Cochemiea, Coryphantha, Escobaria, Neolloydia, and Phellosperma, to mention only a few. Based on their own cladistic analyses, the authors of this paper settle on a system of just three genera: Cochemiea (including Neolloydia and Phellosperma), Coryphantha (including Escobaria), and Mammillaria. To accomplish this, a raft of spp. (we count 22) had to be transferred from Mammillaria to Cochemiea (the necessary new combinations being validated herein), reducing the size of the former genus accordingly. The authors did consider more extreme options: the expansion of Mammillaria so as to be congruent with the entire “Mammilloid clade” was rejected because it would yield “a broadly circumscribed Mammillaria much more difficult to recognize,” while disrupting “our current taxonomy of the group, especially considering the large numbers of species in Coryphantha”; on the other hand, splitting the clade into numerous smaller genera would require at least 15 such segregates from Cochemiea and Coryphantha (in in the sense of this study) alone, of which five would be monospecific (and in any case, many of the historical segregates are themselves non-monophyletic, removing the incentive to cling to those names). So, what does it all mean for Costa Rica? Apart from the aforementioned diminution of Mammillaria, nothing at all: the sole Costa Rican representative of the “Mammilloid clade,” Mammillaria columbiana Salm-Dyck, remains firmly in place.
Caires, C. S. & C. E. B. Proença. 2020. Return to the original concept and new typification of Loranthus spicatus (Loranthaceae), an economically important Neotropical mistletoe. Taxon 69: 1342–1349.
Loranthus spicatus Jacq. is the basionym of Oryctanthus spicatus (Jacq.) Eichler, long the accepted name for a common mistletoe sp. in Costa Rica (and well beyond). This paper emerges from the 2012 Ph.D. dissertation of the first author, embodying a taxonomic monograph of Oryctanthus and Oryctina—one of those Brazilian dissertations which, according to our past expierience, can be difficult or impossible to access. The issue of Loranthus spicatus is a complicated one, but boils down to this: venerable mistletoe specialist Job Kuijt (whose influence on Mesomerican florsistics has been profound) erred in his designation of a Costa Rican specimen (Skutch 3837) as lectotype of L. spicatus, because that specimen conflicts with the protologue in having cylindrical (rather than quadrangular) spikes; these authors thus designate a new lectotype (as well as an epitype) for L. spicatus, with the result that Oryctanthus spicatus now becomes the accepted name for the sp. (also present in Costa Rica) that we have all long known as O. alveolatus (Kunth) Kuijt! So, what name should we now use for the last-mentioned entity? Apparently, Oryctanthus florulentus (Rich.) Tiegh., to judge from the authors' rather cryptic final sentence ["With the new lectotypification of Loranthus spicatus, the specimen previously designated as neotype (A. F. Skutch 3837)…becomes a specimen of Oryctanthus florulentus."]. That sentence becomes even more cryptic when compared with statements in thr first author's dissertation which, we have just discovered, is actually accessible on the Internet, in its entirety, via the following link:
In the dissertation Skutch 3837 is assigned not to Oryctanthus florulentus, rather to O. guatemalensis (Standl.) Standl. & Steyerm., a name that is liberated from synonymy under O. spicatus and applied to most of the material (and all the Costa Rican material) previously (as in the Manual) known by the last-mentioned name. Is the association of the Skutch specimen with O. florulentus merely an error in the paper under review? We think not, because it is reiterated several times, including in the caption of a photo of the specimen itself, in which an annotation label recording Caires's original deterimination as O. guatemalensis is visible. No, clearly, there has been a change of heart since the appearance of the dissertation, but what, exactly, does it mean for us? We have three alternative hypotheses: 1. The name Oryctanthus guatemalensis is now regarded as a synonym of O. florulentus. 2. The concept of O. guatemalensis has been restricted (to Guatemalan material?), and all the Costa Rican (and South American?) material cited under that name in the dissertation is referable to O. florulentus. 3. The Skutch specimen (from the Valle de General) belongs to O. florulentus, but all (or most) of the remaining Costa Rican material cited under O. guatemalensis in the dissertation remains as such—i.e., both O. florulentus and O. guatemalensis are represented in Costa Rica (the two spp. differ, by the way, principally in peduncle length: less than 6 mm in the former, greater than 6 mm in the latter). Certain statements in the present paper suggest that hypothesis #1 merits rejection, but deciding between the other two will likely depend on the publication of an updated version of the dissertation, or critical portions thereof.
So what more can we glean from this dissertation that is relevant to Costa Rican floristics? With respect to Oryctanthus florulentus and O. spicatus, it is worth noting that the former name acquires as a synonym O. phanerolomus (Standl.) Kuijt (type from Belize), which had been synonymized under O. spicatus in the Manual, while O. spicatus inherits as synonyms O. alveolatus and all of the synonyms cited thereunder in the Manual. Oryctanthus cordifolius (C. Presl) Urb. and O. occidentalis (L.) Eichler, the only other spp. treated in the Manual, emerge relatively unscathed. Obviously, the dissertation could not have addressed the subsequently published name Oryctanthus callicarpus Kuijt [see under "Kuijt," this column, in The Cutting Edge 22(1), Jan. 2015], a specious replacement for O. occidentalis subsp. continentalis Kuijt, the name that had been used for all the material of the sp. outside of Jamaica; however, the rejection by Caires of O. o. subsp. continentalis in the dissertation is tantamount to a rejection of O. callicarpus. And finally, a real surprise: an additional (fifth? sixth?) Oryctanthus sp. for Costa Rica! That would be Oryctanthus grandis Kuijt, otherwise known only from western Colombia, but now attributable to Tiquicia on the basis of two specimens (Gómez-Laurito 11254, F; Lent 1187, F) entraining the following parameters: "Bosque muy húmedo, 400–750 m; vert. Carib. Cord. Central (Lagunas Hule y de Río Cuarto). Fl. nov." Hosts are not recorded on the labels of these specimens (both of which are duplicated at CR). In the dissertation, Oryctanthus grandis keys next to O. occidentalis, from which it differs in having sessile leaves, cordate at the base (vs. generally short-petiolate and rarely cordate), and longer spikes (4–15 cm, vs. 0.7–3.4 cm) that are usually glomerate at the branch apices (vs. always axillary). We looked through all the Central American material determined as Oryctanthus occidentalis (as well as genus indets.) at MO, but found no other likely candidates for O. grandis. So the jury is still out! And in case you were wondering: "Oryctina sp. A" of the Manual, later described as Oryctina costaricensis Kuijt (2007), was subsequently transferred to the genus Maracanthus [see under "Kuijt," this column, in The Cutting Edge 21(4), Oct. 2014]. That disposition (then impending) was accepted in this dissertation, which therefore does not treat O. costaricensis.
In closing: we are now faced with a truly nightmarish situation, in which a name (Oryctanthus spicatus) that has been used widely and over a long period for a common sp. is suddenly reapplied to a different sp. that is also common, and has long been known by another name. This is certain to generate widespread and long-lasting confusion, especially considering that it is the second switcheroo involving this name (unbeknownst to us, the name O. spicatus had apparently been used in the sense promoted by this paper—i.e., for the sp. lately known as O. alveolatus—during the pre-Kuijt era). Perhaps some sort of conservation proposal would have been more sensible and may, in fact, still be a possibility. Or maybe that would make matters worse yet. Some authors cling rigidly to the principles of priority and types (as we would generally be inclined to do), while others ignore all of that and turn immediately to conservation or rejection. We find all of this disconcerting; there seem to be no guidelines or rules on how to proceed, so everything depends on the whims of individual authors. A brilliant recipe for nomenclatural instability! And to make matters worse, now we don't really know which or how many spp. of Oryctanthus are present in Cosa Rica.
Cândido, E. S., M. Vatanparast, W. Vargas, L. M. P. A. Bezerra, G. P. Lewis, V. F. Mansano, A. O. Simões, M. J. Silva, C. Stirton, A. M. G. A. Tozzi & A. P. Fortuna-Perez. 2020. Molecular phylogenetic insights into the evolution of Eriosema (Fabaceae): a recent tropical savanna-adapted genus. Bot. J. Linn. Soc. 194: 439–459.
Eriosema was resolved as monophyletic in this study, as were Cajanus and Flemingia (to mention only those genera represented in Costa Rica). So far, so good. The problem child is Rhynchosia, which "is polyphyletic and should be reclassified based on its distinct evolutionary lineages." The prospects for Rhynchosia indeed appear grim. Its largest clade is paraphyletic with respect to the small, Asian genus Paracalyx, while its other major clade, comprising African spp., is sister to Eriosema. Worst of all, the type sp. of Rhynchosia, the Asian R. volubilis Lour., did not come out in either of the aforementioned clades, rather all by itself, as sister to Cajanus; therefore, if the projected reclassification of Rhynchosia were to proceed along the lines suggested by this preliminary study, retention of that genus name for our neotropical spp. could only be achieved by means of a successful conservation proposal. That said, these authors stop short of formally overhauling the classification of this group, pending "further work including a broader sample" [of the nine Rhynchosia spp. occurring in Costa Rica, only R. minima (L.) DC. and R. quercetorum Standl. were included in the present study]. Descending now to a lower taxonomic tier: the two widely recognized vars. of Eriosema simplicifolium (Kunth) G. Don "appeared in different lineages" in these analyses, leading the authors to suggest that the South American E. s. var. micranthum Grear "should be elevated to species rank" (though they pass up the opportunity to validate the necessary new combination). This would relieve us of the need to qualify Costa Rican material as belonging to the autonymic var.
Chase, M. W., M. J. M. Christenhusz, P. Kumar & A. Schuiteman. 2021. (2805) Proposal to conserve Eulophia, nom. cons., against the additional name Geodorum (Orchidaceae: Eulophiinae). Taxon 70: 432–433.
A proposal to conserve Eulophia over Geodorum was promised by another group some six years ago [see under “Bone,” this column, in The Cutting Edge 22(4), Oct. 2015], but never delivered. The authors of the present paper made the same promise just a few months ago [see under “Chase,” this column, in The Cutting Edge 28(2), Apr. 2021], and here it is. Así de fácil!
——, —— & A. Schuiteman. 2020. (2782) Proposal to conserve Calanthe, nom. cons., agains the additional names Phaius, Cyanorkis, and Gastorkis (Orchidaceae, Collabieae). Taxon 69: 1364–1365.
Well, we knew this one was coming [see under "Chase," this column, in The Cutting Edge 28(1), Jan. 2021]. Aquí está!
Clement, W. I., S. Bruun-Lund, A. Cohen, F. Kjellberg, G. D. Weiblen & N. Rønsted. 2020. Evolution and classification of figs (Ficus, Moraceae) and their close relatives (Castilleae) united by involucral bracts. Bot. J. Linn. Soc. 193: 316–339.
Much ado here about the infrageneric classification of Ficus, which interests us not one iota. Of slightly greater relevance, to our small world, is the fact that "Perebea was consistently recovered as paraphyletic" (with respect to Pseudolmedia) in these analyses. As a result, the authors have elected to reinstate the genus Noyera, comprising two South American spp. (both already with names in the last-mentioned genus). The only efffect for Costa Rican floristics is the reduction, by two, of the sp. total for Perebea. The authors also briefly considered "an alternative taxonomic proposal…to expand the circumscription of Perebea to encompass Pseudolmedia," which would have had far more serious consequences for us; however, it was decided that Pseudolmedia "has recognizably distinct morphology that supports maintaining it as a genus for practical reasons."
Croat, T. B., O. O. Ortiz, M. Cedeño-Fonseca & D. Scherberich. 2020. A new Urospatha Schott (Araceae: Lasioideae) from the Caribbean slope of Panama and Colombia. Feddes Repert. 131: 196–202.
The notion of a new sp. of Urospatha, never collected before 1986, from the Atlantic coastal plain of southern Central America, strikes us as outlandish as the discovery of a new sp. of, say, Acrostichum (Pteridaceae), Manicaria (Arecaceae), Montrichardia (Araceae), or Raphia (Arecaceae), under similar circumstances. Introducing Urospatha caribaea O. Ortiz, M. Cedeño & Croat, which may fit the bill, if indeed it is distinct from the standard U. grandis Schott and has not been previously described. Although not documented from Costa Rica, U. caribaea should be sought there, since the Panamanian stations are all in Prov. Bocas del Toro, not all that far from the border. To strain our credulity even further: the authors allude to "yet another probable new species of Urospatha from Central America" based on a Costa Rican collection, Robles 1217 (MO; Parque Nacional Tortuguero), but withhold publication of the suspected novelty because the specimen is "sterile." The MO duplicate may be sterile, but the label notes describe both a spathe and spadix, so a fertile duplicate is likely available, right now, in the CR herbarium. Any takers? A parting comment in defense of the authors: Urospatha is one of those genera—along with most of the others mentioned in first sentence of this paragraph—that nobody ever looks at closely; it is pedestrian, inconvenient to collect, and widely assumed to be monospecific (at least, in our area). Perhaps all such genera merit closer scrutiny!
Eserman, L. A. [et al.]. 2020. (2786) Proposal to change the conserved type of Ipomoea, nom. cons. (Convolvulaceae). Taxon 69: 1369–1371.
We broke with convention, for once, and desisted from enumerating all the authors of this paper, of which there are 41 in all. The authors' names, and their foot-noted addresses, consume an entire page of text, in what is otherwise a two-page conservation proposal! Why did the editors of this journal permit such a thing? C'mon, most of these people must be mere signatories, not authors! Presumably the thinking is that, with more names, the proposal will carry more weight. Never mind that we recognize just two or three of those names, and the ones we recognize are not even associated with Convolvulaceae (to our knowledge, anyway). In our opinion, the overwrought authorship actually detracts from what is otherwise a halfway sensible proposal: to conserve the type of Ipomoea so as to guarantee the use of that genus name for several economically important spp., especially I. batatas (L.) Lam. (the sweet potato). The root of the problem is that Ipomoea, according to its traditional circumscription, is paraphyletic with respect to numerous other genera, e.g., Argyreia, Mina, Rivea, Stictocardia, and Turbina. Not an issue for lumpers like us; we have already embraced proposals [see, e.g., under "Wood," this column, in The Cutting Edge 27(2), Apr. 2020] to lump Stictocardia and Turbina (the only nested genera represented in Costa Rica) back into Ipomoea (where, in fact, names already existed for our spp.). But the authors of this proposal (or at least, one or two of them!) have a splitting agenda (see under "Bouman," this column, for a similar example), and cannot abide a genus of "c. 900 species, without…any infrageneric classification" (so why not get to work on that?). Ipomoea sensu lato, as it turns out, comprises two main clades: one is largely neotropical, and includes most of the economic spp. (the sweet potato, plus several important ornamentals), while the other is almost exclusively paleotropical. Unfortunately, the presently conserved type sp. of Ipomoea is a member of the paleotropical clade, which spells trouble for whomever envisions splitting the genus up, since changing the names of the sweet potato and various morning glories would inevitably be met with stiff resistance. This group therefore proposes to replace the generic type of Ipomoea with I. triloba L., a New World sp. and "one of the most closely related species to I. batatas" (the last-mentioned sp. could not itself be chosen, because Linnaeus had classed it in Convolvulus!). This action, if approved, would retain the name Ipomoea for what we gather to be the overwhelming majority of its neotropical spp. (ca. 600 in all); so this is good news for us, because otherwise most of our names would change (were the splitting option to be widely accepted). We assume, however, that Stictocardia and Turbina will be restored.
These lumper vs. splitter stand-offs are fast becoming the single biggest threats to nomenclatural stability. Two diametrically opposed systems, in simultaneous use by different factions; try explaining that to your horticulture buddies!
Ferreira, J. J. S., A. P. Fortuna Perez, G. P. Lewis & J. Santos Silva. 2021. Characteristics of the fruits of Brazilian species of Stylosanthes Sw. (Leguminosae) and their taxonomic value. Int. J. Pl. Sci. 182: 133–150.
The challenges with Brazilian surnames continue. We feature this contribution because it includes all six taxa of Stylosanthes documented from Costa Rica; one of these, S. guianensis (Aubl.) Sw. var. gracilis (Kunth) Vogel ex ʼt Mannetje according to the Manual, is here ranked as a full sp., S. gracilis Kunth. The authors present a key (dichotomous! indented!), based solely on fruit characters, to 22 Sylosanthes spp. represented in Brazil. Co-PI Nelson Zamora used a fruit character in just one couplet of his Manual key to Stylosanthes, so this new key may prove to be a valuable supplement; some of our readers may wish to give it a shot. The authors also provide a tabular comparison, as well as numerous photogaphic illustrations.
Fonseca-Cortés, A. 2021. Macropsychanthus magnus (Leguminosae: Papilionoideae: Diocleae), a new species from Costa Rica and Nicaragua. Phytotaxa 500: 266–274.
The name Macropsychanthus magnus Fonseca-Cortés (Fabaceae) is validated for a distinctive, lianescent sp. known in Costa Rica by the common name (inexplicably omitted from the Manual Fabaceae treatment) Papa del monte (in reference to its stout, potato-like fruits). The following tale is becoming all too familiar: we first learned this sp., in our early La Selva days, as Dioclea pulchra Moldenke, which soon yielded to the prior and ostensibly synonymous D. malacocarpa Ducke. Recently, Dioclea in the sense of the Manual was divided (with weak justification, we think) into three segregate genera [see the second entry under "Queiroz," this column, in The Cutting Edge 28(1), Jan. 2021], with our guy ending up in the horribly named Macropsychanthus. However, combinations under the latter name were validated for both D. malacocarpa and D. pulchra, leaving us in a state of limbo as to which name to use for Costa Rican material. The answer turns out to be: neither! Dioclea malacocarpa and D. pulchra are indeed maintained as separate spp. (we wondered whether their combinations in Macropsychanthus were made uncritically), both restricted to South America, while Mesoamerican material (Costa Rica and Nicaragua) is distinguished as a third sp., Macropsychanthus magnus. Meanwhile: the common name Papa del monte has held steady, in Costa Rica, for all of these years; scientific names may have certain advantages over common names, but stability over time is clearly not one of them! As denoted by its epithet, Macropsychanthus magnus differs from its two close relatives by its larger flowers, in all of their parts (the distinction between Dioclea malacocarpa and D. pulchra appears more tenuous). The three spp. are compared in a tabular format, and dichotomous (though non-indented) keys are provided, for Costa Rica and Nicaragua, to the genera of the Dioclea clade (Cymbosema, Dioclea, and Macropsychanthus) and the spp. of Macropsychanthus. A distribution map (including Dioclea malacocarpa) is provided, and the new sp. is depicted in a compositie line drawing and color photos from life (with one of D. malacocarpa). Perpetuating an error perpetrated by its original authors, the epithet pulcher (in Macropsychanthus) is consistently misrendered here as "pulchrus."
Gavade, S. K., L. J. G. van der Maesen & M. M. Lekhak. 2020. Taxonomic revision of the genus Flemingia (Leguminosae: Papilionoideae) in India. Webbia 75: 141–218.
What ho, yet another partial revision of Flemingia! Maybe eventually it will all add up to something. This one treats 21 spp. (of an estimated 44 total), including both of those that have been attributed to Costa Rica (as introduced). Like its most recent counterpart [see under "Do," this column, in The Cutting Edge 27(2), Apr. 2020], this effort is seriously compromised by the use of non-indented keys. Apart from that, it appears to be relatively well-executed, with synonymy and typology, technical descriptions, distribution summaries, generous specimen citations, distribution maps, and plenty of illustrations. Still, without a monographic treatment of the genus, we can never be certain of the identity of the spp. in Costa Rica, especially the one dubbed Flemingia macrophylla (Willd.) Kuntze ex Merr.
Ito, Y., N. Tanaka, B. R. Keener & S. Lehtonen. 2020. Phylogeny and biogeography of Sagittaria (Alismataceae) revisited: evidence for cryptic diversity and colonization out of South America. J. Pl. Res. 133: 827–839.
Not much here for us. The authors' "species delimitation analysis suggested" that the two subspp. (one neotropical, one paleotropical) of Sagittaria guayanensis Kunth "could be recognized as two different species," a course of action which they "recommend." Real good. For Costa Rica, this would only mean that our material of S. guayanensis would no longer have to be qualified as belonging to the autonymic subsp.
Jorgensen, S. A. & D. S. Barrington. 2020. Speciation and reticulation in the Polystichum allotetraploids of the Costa Rican Cordillera de Talamanca. Amer. Fern J. 110: 151–164.
Most of this is outside of our province, but we do learn that Costa Rican material heretofore identified (by the second author of this paper, and others) as Polystichum orbiculatum (Desv.) J. Rémy & Fée actually represents a different sp. that is "currently undescribed" (and so remains, though a new name is said to be "under consideration"). According to these authors, true P. orbiculatum is a central Andean, diploid sp. not even involved in the origin of the Costa Rican entity, which is here dubbed the "páramo tetraploid." In Costa Rica, the "páramo tetraploid" (which apparently extends to the northern Andes) is restricted to the very highest elevations (3400–3800 m); just four Costa Rican records are to be found under Polystichum orbiculatum in TROPICOS.
Juárez, P. & J. F. Morales. 2021. Pleurothyrium amissum (Lauraceae), a new endemic tree from the Talamanca montane forest in Costa Rica. Novon 29: 78–84.
"Pleurothyrium sp. 1," one of two provisionally named spp. among the 10 members of its genus ascribed to Costa Rica in the Manual Lauraceae treatment, now receives a Latin binomial: Pleurothyrium amissum P. Juárez & J. F. Morales. Its epithet, meaning "lost," alludes to the fact that the tree from which the type specimen was harvested has been cut down (as of 2015), and no other individuals have been located in the immediate vicinity. Fortunately, the tree that yielded the lone paratype is protected in Parque Nacional Chirripó. A distribution map and a key (dichotomous and indented) to the "currently accepted" spp. of Pleurothyrium in Costa Rica are provided, and the new sp. is illustrated with a composite line drawing and color photos from life.
Kataoka, E. Y. & L. G. Lohmann. 2021. Taxonomic revision of Martinella Baill. (Bignonieae, Bignoniaceae). PhytoKeys 177: 77–116.
Well, this is a surprise, on a couple of levels—first, by virtue of its mere existence, coming as it does just seven years after a "synopsis" of Martinella from the same lab [see under "Zuntini," this column, in The Cutting Edge 21(3), Jul. 2014]. The last-mentioned contribution upped the sp. total for Martinella to three, with the description of a new Brazilian sp., but upheld the traditional notion of a single sp., M. obovata (Kunth) Bureau & K. Schum., in the Mesoamerican region. That was the snapshot portrayed in the recent (2020) Manual Bignoniaceae treatment, but now it is all out the door. Two more new South American spp. of Martinella are described in this proper revision, raising the generic sp. total to five; but more significant, from our perspective, is the second big surprise: the sudden and unexpected northward extension, all the way to Costa Rica, of a sp. that was restricted in the synopsis to the Amazon basin. The sp. in question had been called Martinella iquitoensis A. Samp., but that name now yields to the prior M. insculpta Sprague & Sandwith, which was unaccountably overlooked in the synopsis. In Costa Rica, Martinella insculpta occurs only on the Pacific slope, as expressed by the following parameters: "Bosque muy húmedo, bosques primarios e intervenidos, 50–650 m; vert. Pac. Cord. de Guanacaste, región de Turrubares, P.N. Carara, región de Puriscal (P.N. La Cangreja). Fl. oct." The real Martinella obovata attains higher elevations in Costa Rica and occurs on both slopes, but the two spp. occur sym- or parapatrically in the Cordillera de Guanacaste and Parque Nacional Carara. Martinella insculpta may be distinguished from M. obovata by its quadrangular (vs. cylindrical) mature stems, distally pulvinulate (vs. non-pulvinulate) petioles and petiolules, larger (15–32 × 8.2–23.5, vs. 7.9–13.2 × 3–9.4 cm) and coriaceous (vs. chartaceous) leaflets, and "botryoid" (vs. racemose) inflorescences. Features synonymy, typology, technical descriptions, and discussions at all ranks, a dichotomous (though non-indented) key to spp., distribution and phenology summaries, conservation assessments, distribution maps, generous specimen citations, and an index to exsiccatae (in a supplementary file). The introductory portion (with two composite color plates) addresses distribution, reproductive biology, uses, cytology, and morphology. Each sp. is depicted in a full-page (or nearly so) composite line drawing.
Kechaykin, A. A., A. I. Schmakov, T. Gregor, J. Paule & D. A. German. 2021. (2820) Proposal to conserve the name Potentilla (Rosaceae: Potentilleae) with a conserved type. Taxon 70: 680–681.
The acceptance of Horkelia and Ivesia, “morphologically well-defined genera” of North America, renders the large genus Potentilla paraphyletic. Maintenance of the two aforementioned genera, insistence on monophyletic taxa, and faithfulness to the type principle would restrict the name Potentilla to “a group of 10–12 species,” while “ca. 300 species” would require new combinations under some different generic name. The thinking these days is that, if a type doesn’t suit you, change the type; hence this proposal, which would do so in a way that maintains the name Potentilla for the 300 or so spp. that would otherwise need new names. We have a better idea: lump Horkelia and Ivesia back into Potentilla! That is how we learned them to begin with, and it was never a problem. As to the potential fate of the two Costa Rican representatives of Potentilla: the rare weed P. indica (Andrews) Th. Wolf actually belongs to the same clade as the current type sp., so would wind up in a different genus (apparently Tormentilla L.) if this proposal is approved; however, the prospects for the native P. heterosepala Fritsch are unclear.
Kron, K. A., P. W. Fritsch, L. Lu & W. S. Judd. 2020. New combinations and new and resurrected names in Gaultheria (Ericaceae). Gard. Bull. Singapore 72: 299–317.
The inclusion in Gaultheria of the large, southeast Asian genus Diplycosia [see The Cutting Edge 9(1): 8–9, Jan. 2002] and the minor South American genus Tepuia is here formalized with the validation of 126 new combinations and 17 nomina nova. All this means for us is that the sp. total for Gaultheria, "ca. 115" according to the Manual Ericaceae treatment (2010), abruptly balloons to "ca. 292." That total also includes Pernettya, which was accepted in the Manual despite having already been consigned to synonymy under Gaultheria by many Ericaceae buffs. None of these taxonomic changes requires modifications to the global distribution of Gaultheria indicated in the Manual. And incidentally: if we are not mistaken, the correct name in Gaultheria for Pernettya prostrata (Cav.) DC., the only sp. of Pernettya recognized for Costa Rica in the Manual, must be Gaultheria myrsinoides Kunth.
Levin, G. A. & V. G. Sagun. 2021. (2807) Proposal to conserve the name Acalypha wilkesiana against A. tricolor (Euphorbiaceae). Taxon 70: 435–436.
These authors have discovered that Acalypha tricolor Veitch ex Mast., a name that “has never been accepted,” was published three months prior to A. wilkesiana Müll. Arg., the name that has been consistently used (sometimes at the rank of subsp.) for an ornamental sp. that is cultivated worldwide. The latter is a well-known sp. in Costa Rica, and was mentioned in the Manual as commonly grown there. The validating description for Acalypha tricolor is brief, but “diagnostic of A. wilkesiana,” in the view of these authors; and for good measure, they neotypify the former name accordingly—setting the stage for this proposal.
Londoño-Echeverri, Y., A. M. Trujillo-López & J. A. Pérez-Zabala. 2021. Rediscovering the genus Lubaria (Rutaceae: Galipeinae), a new species and first record for Colombia. Brittonia 73: 77–84.
Lubaria had been a monospecific genus, with L. aroensis Pittier occurring disjunctly in Costa Rica (where it was first found in 1994) and Venezuela. The new sp. described in this paper doubles the total for the genus, and extends the geographic range of the latter to northern Colombia. The Manual Rutaceae treatment (2014) had jumped the gun in already ascribing two spp. to Lubaria!
Mapaya, R. J. & G. V. Cron. 2021. A phylogeny of Emilia (Senecioneae, Asteraceae) – implications for generic and sectional circumscription. Taxon 70: 127–138.
Emilia is shown to be paraphyletic with respect to two minor African genera, which are sensibly and summarily lumped therein. So there is little here for us, other than a small increase in the generic sp. total (now 117, according to this source). These authors assert that Emilia is an Old World genus, with two weedy spp. that are adventive in the Neotropics. If so, we did not get that quite right in the Manual, where one of the two Emilia spp. in Costa Rica (E. fosbergii Nicolson) is indicated as being native in the New World. We always make an effort to establish the native ranges of weedy spp., but many sources are not forthcoming on that issue, and indeed, the truth can be extremely elusive.
Menjívar Cruz, J. E., R. M. Baldini, G. Davidse & P. M. Peterson. 2021. El Salvador grasses: an updated catalogue and nomenclator. Phytoneuron 2021-22: 1–131.
Although this substantial contribution may provide some insights into the current thinking of agrostologists—we note, for example, that Hackelochloa granularis (L.) Kuntze is no longer included in the genus Mnesithea—it includes just one signficant change (as far as we can detect) relevant to Costa Rica floristics: the new combination Rugoloa pilosa (Sw.) Zuloaga var. lancifolia (Griseb.) Baldini & J. Menjívar (the only nomenclatural novelty here validated), which applies to the taxon previously known (as in Manual Vol. 3) as Panicum pilosum Sw. var. lancifolium (Griseb.) R. W. Pohl.
Miranda, C. V., P. B. Schwartsburd, P. H. Labiak & J. Prado. 2021. Three new species of Oleandra (Oleandraceae, Polypodiopsida) from the Neotropics, and notes on the morphological groups among the Neotropical species. Brittonia 73: 143–151.
The three spp. described in this paper are based on a collective total of just five specimens, which raises a red flag right away. One of those specimens, the lone paratype of Oleandra hovenkampii C. V. Miranda & Schwartsb. (Scamman 5914, GH), is Costa Rican (from Orosi, Prov. Cartago); the holotype, and only other specimen cited, is from Peru. That scenario strains credulity, to put it mildly—not the least because the Costa Rican and Peruvian collections both come from very heavily botanized locales. Oleandra hovenkampii is compared with five other spp., none of which occurs in Costa Rica. This is one hypothesis that desperately needs testing! The authors list 23 herbaria from which specimens were “analyzed,” but just five of those are located outside Brazil, and none are in Mesoamerica or other South American countries, or even specialized in the floras of those regions. Among the five Oleandra spp. now attributable to Costa Rica, O. hovenkampii most resembles O. articulata (Sw.) C. Presl and O. bradei Christ in its spreading (vs. appressed) rhizome scales and alternate (vs. verticillate) phyllotaxy, but differs from those spp. by virtue of its lanose (vs. “glabrescent to glabrous”) leaf blades.
Moroni, P., A. Martínez, M. G. Nadra & N. O’Leary. 2021. Back to the starting point: revised typifications of Linnaean names based on Plumier’s Nova plantarum Americanarum genera. Taxon 70: 660–669.
This is what the journal Taxon is all about these days: edifying types on the one hand, disrespecting them on the other. What we have here is an edification project, in which 14 previous lectotypifications of Linnaean names based on Burman illustrations are superseded, on the grounds that those illustrations were not available to Linnaeus in a timely fashion. Well and good. Of the names addressed, just five pertain directly to Costa Rican floristics: Bucephalon racemosa L. [the basionym of Trophis racemosa (L.) Urb.; Moraceae], Mentzelia aspera L. (Loasaceae), Samyda guidonia L. [the basionym of Guarea guidonia (L.) Sleumer; Meliaceae], Thalia geniculata L. (Marantaceae), and Tournefortia hirsutissima L. (Boraginaceae). No changes in usage are implied.
Murillo-Serna, J. S., F. A. Michelangeli & F. Alzate-Guarin. 2020. (2787) Proposal to conserve the name Graffenrieda against Centronia (Melastomataceae: Merianieae). Taxon 69: 1371–1372.
The type of the genus name Centronia has been resolved (based on molecular studies) as nested among spp. of the otherwise monophyletic Graffenrieda. This may not pose a problem for the other spp. currently classed in Centronia, which these authors suggest may be better placed in Meriania or Graffenrieda anyway (though none has been included in molecular analyses). As far as Costa Rica is concerned, our only Centronia sp., C. phlomoides Triana, had been transferred to Meriania even before the publication of the Manual Melastomataceae treatment. It is Graffenrieda, a genus of about 70 spp. (with three in Costa Rica), that these authors are mainly worried about, because the name Centronia has priority. Therefore, should this proposal be rejected, some 70 new combinations in Centronia would be required for spp. currently assigned to Graffenrieda.
Oliveira, R. C., J. Sonsin-Oliveira, T. A. C. dos Santos, M. Simas e Silva, C. W. Fagg & R. Sebastiani. 2021. Lecotypification of Banisteriopsis caapi and B. quitensis (Malpighiaceae), names associated with an important ingredient of Ayahuasca. Taxon 70: 185–188.
Hope we got those Portuguese surnames right! Just when we think we have it figured out, there is always some new twist. Getting down to business: Banisteriopsis caapi (Spruce ex Griseb.) C. V. Morton, the accepted name for a notorious sp. that is rarely cultivated in Costa Rica, is lectotypified, as is one of its synonyms. So much for that.
Ortuño Limarino, T. & T. Borsch. 2021. Gomphrena (Amaranthaceae, Gomphrenoideae) diversified as a C4 lineage in the New World tropics with specializations in floral and inflorescence morphology, and an escape to Australia. Willdenowia 50: 345–381.
It started out in Gomphrena, then progressed through Philoxerus (as in Flora costaricensis), Caraxeron (very briefly), and (most recently) Blutaparon. Now, the coastal sp. treated as Blutaparon vermiculare (L.) Mears in Manual Vol. 4(1) (2020) completes the round trip, reverting to its basionym, Gomphrena vermicularis L., on the basis of molecular studies confirming Blutaparon (or whatever) as "deeply nested within the Gomphrena clade." We use the word "confirming" because this disposition was suggested more than a decade ago [see The Cutting Edge 16(2): 12, Apr. 2009], in a paper that was actually cited in the Manual! We did consider sinking Blutaparon into Gomphrena, but abandoned the idea because the generic transfer had not been formalized; that is now our second biggest regret, after failing to scuttle the embarrassingly bogus genus Conyza (Asteraceae). In the present paper, the transfer of Blutaparon (etc.) to Gomphrena is finally formalized with validation of the necessary new combinations; as previously mentioned, we did not need one of those at sp. rank, but no combinations are made here at infraspecific ranks, so there is at present no autonymic counterpart in Gomprena for Blutaparon vermiculare var. vermiculare, to which Costa Rican material had been tentatively assigned. Those of our readers who may be interested in any of the fancified topics mentioned in the title of this article are encouraged to delve more deeply.
Penagos-Zuluaga, J. C., H. van der Werff, B. Park, D. A. R. Eaton, L. S. Comita, S. A. Queenborough & M. J. Donoghue. 2021. Resolved phylogenetic relationships in the Ocotea complex (Supraocotea) facilitate phylogenetic classification and studies of character evolution. Amer. J. Bot. 108: 664–679.
Let's be honest, how many of us can distinguish all the genera of Lauraceae in the Neotropics? This study confirms that we are not all incompetent or poorly informed: many of these genera really are bogus! Here unmasked as polyphyletic are Aniba, Endlicheria, Licaria (which seemed to us one of the more distinctive entities in the family), and Rhodostemonodaphne (among the genera represented in Costa Rica). The recently resurrected Mespilodaphne [see under "Trofimov," this column, in The Cutting Edge 27(3), Jul. 2020] is claimed as monophyletic, but the authors' own cladogram suggests otherwise. Sure, a few genera in the so-called "Ocotea complex" do pass muster: Pleurothyrium (tentatively), as well as Damburneya [spun off quite recently; see under "Trofimov," this column, in The Cutting Edge 24(1), Jan. 2017] and Nectandra (though its circumscription has been altered drastically over the past several decades). However, even those are deeply nested (with all the rest) in a hopelessly paraphyletic Ocotea. It is all too easy to cherry-pick a few distinctive and monophyletic taxa, but sometimes an indefinable and intractably paraphyletic quagmire is left behind; subsequent workers are left to either do the hard word and break the quagmire up into smaller, monophyletic, and diagnosable genera, or else bite the bullet and lump everything into a single, gargantuan genus. The situation in the Ocotea complex brings to mind similar dilemmas recently brought to light in Campanulaceae subfam. Lobelioideae, Crassulaceae subfam. Sempervivoideae, and Melastomataceae tribe Miconieae. The jury is still out for the first-named taxon [see under "Kagame," this column, in The Cutting Edge 28(2), Apr. 2021], but Melastomataceae authorities bit the bullet and lumped the entire tribe into a single genus, Miconia [see under "Michelangeli," this column, in The Cutting Edge 26(2), Apr. 2019], and Sedum appears destined for a similar enrichment at the hands of Crassulaceae workers [see under "Messerschmid," this column, in The Cutting Edge 28(1), Jan. 2021]. What are the prospects, then, for the Ocotea complex? Reading from the main cladogram in this paper, it would appear that Endlicheria and Rhodostemonodaphne could easily be combined under the former (older) name to yield a monophyletic unit; beyond that, however, the sledding would be tough, and we calculate that Ocotea would have to be divided into a least nine segregate genera to resolve the problem under the splitting option. The fact that the type sp. of Ocotea (the South American O. guianensis Aubl.) was not included in this study does not advance that agenda. The lumping option would, of course, result in all the genera mentioned in this paragraph (excepting Cassytha!) being combined (along with several others not represented in Costa Rica, such as Umbellularia) under one name, logically Ocotea (though Aniba and Licaria have equal priority). Needless to say, we would vote for the lumping option, since we are lumpers at heart, and also because most of the "genera" involved could never be distinguished by the majority of workers, and several are themselves polyphyletic. But it remains to be seen how the Lauraceae specialists will proceed. Despairing that to express the phylogenetic relationships in the Ocotea complex "in the Linnean framework would necessitate the formal recognition of genera that lack diagnostic morphological characters and defy identification," these authors instead provide names under the PhyloCode (such as the one in the title) for "major newly recognized clades without changing standard generic concepts." These names are intended "to facilitate communication about relationships as studies of Lauraceae proceed," and "are meant, for the time being, to complement, not replace, the standard rank-based classification." These last statements give us hope that the authors have not forsaken the ideal of a "Linnean framework" for the Ocotea complex, and will continue working diligently to hammer out a proper classification that features only monophyletic and morphologically diagnosable genera (or a single such genus; they give no indication of having seriously considered the meritorious lumping option). Of course, the taxonomic community at large will never be satisfied with this PhyloCode business, and if these authors are not willing or able to progress beyond it, others will happily take up the cause (consider the potential rewards in new combinations!). The Ocotea complex comprises at least 80% of neotropical Lauraceae spp. (to use the authors' own figure), all of which now find themselves in a state of taxonomic limbo.
Pupulin, F., J. Aguilar, N. Belfort-Oconitrillo, M. Díaz-Morales & D. Bogarín. 2021. Florae costaricensis subtribui Pleurothallidinis (Orchidaceae) prodromus II. Systematics of the Pleurothallis cardiothallis and P. phyllocardia groups, and other related groups of Pleurothallis with large vegetative habit. Harvard Pap. Bot. 26: 203–295.
The first installment of this tediously titled series focused on the genus Echinosepala [see the second entry under “Pupulin,” this column, in The Cutting Edge 28(1), Jan. 2021], while this second one tackles a more heterogeneous assemblage. In fact, four “informal assemblages” are here discerned, encompassing a total of 26 spp. and four “natural hybrids.” It is noteworthy that only 11 of the 26 spp. (and none of the hybrids) accepted in this article were recognized as distinct in the Manual Orchidaceae treatment (2003), all the rest having been described or (in two cases) resurrected from synonymy since that time. Three of those new spp., as well as three new hybrids—all six endemic to Costa Rica—are described in this very paper. The three new spp. are: Pleurothallis callosa M. Díaz & Pupulin, compared to P. cardiothallis Rchb. f., from ca. 1200 m elevation on the Atlantic slope of the northern Cordillera de Talamanca; P. longipetala Bogarín & Belfort, compared inter alia with P. radula Luer, from 1450–2800 m on both slopes of the Cordillera de Talamanca; and P. mesopotamica Bogarín & Belfort, compared with P. rectipetala Ames & C. Schweinf., from ca. 1550–1600 m on the Atlantic slope of the Cordillera de Talamanca. The new hybrids are: Pleurothallis ×karremansiana Pupulin, J. Aguilar & M. Díaz, alleged to involve P. tonduzii Schltr. and P. gonzaleziorum Pupulin, M. Díaz & Pridgeon [see under “Pupulin,” this column, in The Cutting Edge 27(3), Jul. 2020], from ca. 1500–1550 m on the Atlantic slope of the western Cordillera Central; P. ×parentis-certa Pupulin & Bogarín, ostensibly involving P. tonduzii and “a species of the Pleurothallis phyllocardia group,” without locality data; and P. ×subversa Pupulin & Bogarín, proposed as a hybrid between P. compressa Luer and “a large-flowered species close to P. cardiothallis” (most likely P. oncoglossa Luer), from ca. 1300 m on the Atlantic slope of the northern Cordillera de Talamanca. In addition, Pleurothallis ×silvae-pacis Karremans, though originally described as a sp. [see the first entry under “Karremans,” this column, in The Cutting Edge 19(4), Oct. 2012], is here reinterpreted as a “natural cross” between P. eumecocaulon Schltr. and P. phyllocardia Rchb. f. Two other taxonomic innovations, with respect to the Manual, merit attention: Pleurothallis rectipetala and P. triangulabia C. Schweinf. are both liberated from synonymy under P. phyllocardia; the former name is applied to a Costa Rican endemic, ranging from ca. 1300–1450 m elevation on the Atlantic slope of the Cordillera de Talamanca, while P. triangulabia denotes a sp. occurring at (800–)1600–2400 m on both slopes of the Cordilleras de Tilarán and Central, as well as in western Panama. The real Pleurothallis phyllocardia, typified by a Costa Rican specimen, obviously remains a member of the country’s flora, and is relatively widespread in montane habitats. Finally, two spp. heretofore believed endemic to western Panama are reported from Costa Rica for the first time: Pleurothallis anthurioides A. Doucette, from ca. 2000–2050 m on the Pacific slope of the eastern Cordillera de Talamanca; and P. maduroi Luer, from ca. 2150–2200 m on the Atlantic slope of the eastern Cordillera de Talamanca.
The taxonomic portion of this revision is headed by a dichotomous (though non-indented) key to spp. Most inconveniently (for our purposes), the sp. entries are ordered alphabetically within each of the four “informal assemblages.” Each entry features synonymy and typology, a technical description, distribution summaries, an enumeration of “distinguishing features,” and specimen citations. A section at the end entitled “Excluded species” reveals that the Costa Rican record for Pleurothallis scitula Luer cited in the Manual genus discussion of Pleurothallis was based on a misdetermined specimen of P. triangulabia, while an additional Costa Rican voucher cited in this very rag [see The Cutting Edge 15(1): 3, Jan. 2008] could not be located; hence the omission of P. scitula from this account. There are no índices. The introductory portion addresses taxonomic history (with an emphasis on recent phylogenetic studies), the characterization of the informal sp. groups, ecology (especially floral), color dimorphism, and hybridization. As usual with this crew, this work is copiously and brilliantly illustrated throughout, including color photos, excellent composite line drawings, and distribution maps.
Factoring in the five new orchid spp. featured in this issue of the Edge (three in this paper, plus two others under "Bogarín," this column), our much-ballyhooed running count (of new spp. of Orchidaceae described from Costa Rica during the tenure of this blog, i.e., ca. 1993–2021) now stands at 425. Anyone doing the math may have calculated 426, but we do not count such sketchy entities as hybrids, and have therefore deducted Pleurothallis ×silvae-pacis from our tally [the reverse transpired with Epidendrum sandiorum Hágsater, Karremans & L. Sánchez, originally described as a hybrid but later elevated to full sp. status; see the third entry under "Szlachetko" et al., this column, in The Cutting Edge 24(2), Apr. 2017].
Riina, R., B. W. Van Ee, M. B. R. Caruzo, D. S. Carneiro-Torres, R. F. Santos & P. E. Berry. 2021. The Neotropical Croton sect. Geiseleria (Euphorbiaceae): classification update, phylogenetic framework, and seven new species from South America. Ann. Missouri Bot. Gard. 106: 111–166.
Croton sect. Geiseleria (A. Gray) Baill., widespread throughout the Neotropics, comprises 84 spp. (as recognized by these authors), and is characterized by “stellate pubescence, usually dentate or crenate leaves with a pair (or more) of usually stipitate nectary glands at the apex of the petiole, nectary glands often in the sinuses of the leaf margins, pistillate flowers with unequal sepals in many species, and bisexual inforescences with unisexual cymules” (among other things). Much of this paper transcends our narrow sphere of interest, dealing with such topics as the circumscription of the section, infrasectional classification, and the circumscription and description of extralimital spp. The seven new spp. mentioned in the title (none of which concerns us) are described separately, following which a “revised synopsis” of sect. Geiseleria is presented. The synopsis begins with a technical description of the section, followed by a distribution summary, “Notes,” and a dichotomous and indented key to the four subsections recognized by the authors, after which the subsections themselves (three of which are new, at least in terms of status) are briefly described, with their constituent spp. enumerated. Synonymy and typology are provided for the sectional name and those of the subsections, as well as for the spp., all 84 of which are listed in alphabetical order following the subsectional treatment (there are no keys to spp.). Each sp. entry also features a distribution summary and (usually) “Notes.” A section on “Excluded taxa” concludes the proceedings. There are no indices or (apart from types) specimen citations (except for the new spp. and accessions for the molecular analyses). Just six spp. are attributed to Costa Rica: Croton brevipes Pax, C. glandulosus L., C. hirtus L’Hér., C. jutiapensis Croizat, C. ortholobus Müll. Arg. (the sole Costa Rican endemic in the group), and C. trinitatis Millsp. Just one of these, C. glandulosus, comes as a mild surprise to us, because it was not formally treated in the Manual Euphorbiaceae account (2010). It was, however, mentioned there, in the Croton genus discussion, as “conocida de Nic. y Pan.” and thus to be expected in Costa Rica. Having just this week checked all the most pertinent herbarium databases (CR, F, GH, MO, NY, US), we still are unable to find any evidence that Croton glandulosus has ever been collected in Costa Rica; therefore, because no specimens are cited in this paper, we remain uncertain as to whether these authors have seen some Costa Rican collection that has escaped our scrutiny, or are merely repeating an old, debunked literature report.
The opening part of this paper (featuring a distribution map and selected color photos from life) is concerned mainly with phylogenetics.
Rojas-Alvarado, G., M. A. Blanco & A. P. Karremans. 2021. A taxonomic sinopsis and morphological characterization of Myoxanthus (Orchidaceae: Pleurothallidinae). Phytotaxa 507: 211–258.
Coming hard on the heels of a fully realized revision of the Costa Rican members of Myoxanthus from this same lab [see under “Rojas-Alvarado,” this column, in The Cutting Edge 27(3), Jul. 2020], this mere synopsis, though broader in its scope, would figure to hold few surprises for us; and indeed, that is the case. We note only the following: Panamanian material previously included in Myoxanthus scandens (Ames) Luer is here discriminated as a distinct sp., M. panamensis Rojas-Alv. & Karremans (corrected from the authors’ “panamense”). This has the happy result of conferring Costa Rican endemicity upon M. scandens! We’ll see how long that lasts…. The synopsis itself (more of an annotated checklist) comprises an alphabetical listing of the 52 spp. accepted by the authors, punctuated by the expected plates of color photos, and including synonymy and typology (also for the genus), distribution summaries, and notes, followed by sections entitled “Incertae sedis” and “Excluded Taxa.” There are no keys or indices, and descriptions are provided only for the genus and the single new sp. The lengthy and well-illustrated introductory portion addresses taxonomic history and (mainly) the “morphological characterization” promised in the title.
Saini, L. & V. Malik. 2021. Neotypification of Solanum chrysotrichum (Solanaceae). Phytotaxa 497: 298–300.
So it's come to this. Solanum chrysotrichum Schltdl., the accepted name for a well-known Mesoamerican sp., is neotypified by two workers based in India. Who have not even seen the desgnated neotype, only an image of it. We have just one question: why? What interest do these authors have in S. chrysotrichum? Indeed, what knowledge do they have of it? Is it adventive in India? These things are not revealed in this brief article. Should not neotypifications be left to specialists, ideally, monographers of the genus or infrageneric taxon involved? We think so. What's the rush, anyway? Has COVID-19 so impacted the ability of taxonomists to conduct real research, that they must resort to this type of endeavor? Does a paper like this—which involves no science whatsoever—actually earn points toward promotion in the academic world? And if not, what can be the motivation? Why would a reputable scientific journal even accept such a manuscript? We remain mystified, at all levels, by the typification craze that continues to rage all around us. With all of that said, we expect that this particular neotypification will do no harm, only because the designated neotype (imaged in the paper) has been annotated as Solanum chrysotrichum by both Sandra Knapp (BM) and Michael Nee, respected regional authorities on Solanum. Still, we cannot help but wonder whether a duplicate (i.e., potential lectotype) of Schiede 81 (the original type of S. chrysotrichum) may lurk among the many herbaria (more than 20!) to which "further material" of Schiede was distributed (according to TL-2); the authors of this paper give no indication of having considered such a possibility.
Sanín, D. 2018. Serpocaulon (Polypodiaceae). Flora de Colombia No. 32. Inst. Ci. Nat., Univ. Nac. Colombia, Bogotá, Col. 137 pp.
Normally we do not feature floristic works for countries or regions other than Costa Rica, but this one contains one minor bombshell that affects us: the name Serpocaulon loriciforme (Rosenst.) A. R. Sm., long used (along with its basionym in Polypodium) for a familiar sp. in Tiquicia (it is especially common at the Estación Biológica La Selva), is consigned to the synonymy of Serpocaulon dasypleuron (Kunze) A. R. Sm. Some of our readers may find this well-executed and generously illustrated treatment to be useful for general purposes, inasmuch as the 22 Serpocaulon spp. in Colombia include all but a few of the 14 spp. recorded from Costa Rica.
——, J. Mostacero & A. R. Smith. 2021. A new Serpocaulon (Polypodiaceae) from northern South America and a reinterpretation of S. caceresii. Amer. Fern J. 111: 68–85.
The main value of this paper, from our perspective, was to remind us that the name Serpocaulon giganteum (Desv.) A. R. Sm., previously applied to a sp. represented in Costa Rica, has been replaced by S. articulatum (C. Presl) Schwartsb. & A. R. Sm. [see under “Schwartsburd,” this column, in The Cutting Edge 21(2), Apr. 2014], and also to alert us to the first author’s recent treatment of Colombian Serpocaulon (see the foregoing entry), where we learned something else. In the present paper, yet another name that had occasionally been applied to Costa Rican material is scuttled: Serpocaulon caceresii (Sodiro) A. R. Sm., here revealed as a synonym of S. articulatum. The name S. caceresii had been accepted even as recently as 2018 (see previous entry), albeit for a sp. restricted to South America. The new sp. described in this paper largely corresponds to that entity. For us, the bottom line is this: abandon all thoughts of Serpocaulon caceresii and S. giganteum! It seems that all the Mesoamerican material ever branded with either of those names is now correctly Serpocaulon articulatum.
Santamaría-Aguilar, D., R. Aguilar Fernández & H. van der Werff. 2021. Beilschmeidia osacola (Lauraceae) a new species from the Osa Peninsula, Costa Rica. Phytotaxa 498: 197–204.
The Manual Lauraceae treatment, published in 2007, included two provisionally named Beilschmiedia spp.: B. "sp. 1" and B. "sp. 2." Beilschmiedia osacola Aguilar, D. Santam. & van der Werff, described in this paper, is neither of those; indeed, it was first collected only in 2008. The most distinctive attribute of the new sp. is the "small, turbinate to campanulate flowers," with a "relatively deep receptacle which largely encloses the ovary." That feature invites comparison with Cryptocarya; however, the fruits of the new sp. "are seated unprotected on the pedicel," vs. "enclosed by the accrescent floral tube" in Cryptocarya. Within Beilschmiedia, the new sp. is compared most closely (in narrative and tabular form) with B. costaricensis (Mez & Pittier) C. K. Allen. A distribution map is provided, as well as a composite line drawing and photos from life of Beilschmeidia osacola. Curiously, although only the holotype is formally cited, at least three other collections would appear to exist, according to the vouchers provided for the illustrations.
—— & L. P. Lagomarsino. 2021. Two new species of Otoba (Myristicaceae) from Colombia. PhytoKeys 178: 147–170.
The first author’s work toward a Flora mesoamericana treatment for Myristicaceae continues to bear fruit, even in a genus that was revised only last year [see under “Jaramillo-Vivanco,” this column, in The Cutting Edge 27(3), Jul 2020]. The present paper affects us only insofar as one of the new Colombian spp., Ocotea scottmorii D. Santam., is segregated from O. acuminata (Standl.) A. H. Gentry, which the authors “now consider to be endemic to the Caribbean slope of Costa Rica and Panama.” Annotate your copies!
Schiavinato, D. J., D. G. Gutiérrez & A. Bartoli. 2021. Typifications and taxonomical rearrangements in North and Central American Tagetes (Asteraceae, Tageteae). Phytotaxa 507: 81–97.
In the course of their revisionary work on Tagetes, these authors “detected the need for typifications” in the genus. The result is this paper, in which lectotypes (plus one epitype) are designated for 12 names. Just one of these, Tagetes lucida Cav., was used as an accepted name in the Manual, albeit for a sp. merely mentioned as occasionally cultivated in Costa Rica. However, six synonyms (some of which were not cited in the Manual) of accepted names used in the Manual are lectotypified (and, in one case, epitypified): one of Tagetes foetidissima DC., one of T. lucida, and three of T. tenuifolia Cav. The net effect for Costa Rican floristics is zilch.
Schneider, J. V., T. Jungcurt, D. Cardoso, A. M. Amorim, M. Töpel, T. Andermann, O. Poncy, T. Berberich & G. Zizka. 2021. Phylogenomics of the tropical plant family Ochnaceae using targeted enrichment of nuclear genes and 250+ taxa. Taxon 70: 48–71.
"Targeted enrichment"? So it's come to that. Ouratea, the largest neotropical genus of Ochnaceae, survived this scare and "came out as monophyletic"; however, "the neotropical and palaeotropical taxa of Sauvagesia form independent clades, requiring the re-erection of formerly separate genera." The latter quote comes from the abstract, but in the main text the authors hedge their bets, allowing that "it is beyond the scope of the present study to judge if this [splitting] is a sound solution to the problem or whether a broad concept [lumping] is to be preferred…" Indeed, Sauvagesia could be maintained largely according to its prevailing circumsciption merely by absorbing the obscure, dispecific South American genus Adenarake, which is basal to the neotropical clade of the former genus. Whatever the case, Costa Rican floristics does not stand to be affected by any of this—except possibly with respect to the generic sp. total for Sauvagesia—because the type sp. of said genus (Sauvagesia erecta L.) is a member of the neotropical clade.
Taylor, C. M. 2021. Rubiacearum americanarum magna hama pars XLVIII: a new view of Palicourea sect. Axillares and description of some new species (Palicoureeae). Novon 29: 85–111.
The taxon here treated as Palicourea sect. Axillares (Hook. f.) Borhidi comprises 14 spp., “found from southern Central America through the Andes of northwestern and western South America with a center of diversity in eastern Colombia and western Venezuela.” The group “is characterized by the combination of laminar, well-developed, bilobed stipules and mostly capitate, pseudoaxillary or sometimes terminal, mostly sessile inflorescences with numerous well-developed bracts that enclose the flowers but without enlarged involucral bracts.” The “new view” espoused in this paper involves minor changes to the circumscriptions of the section and some of its members, as well as the description of four new spp., but these developments have no impact on Costa Rican floristics. Just two of the 14 spp. occur in Costa Rica: Palicourea axillaris (Sw.) Borhidi and Palicourea pseudaxillaris (Wernham) C. M. Taylor. In the Manual Rubiaceae treatment (2014), co-written by the author of this paper, these spp. were treated as Psychotria aubletiana Steyerm. and Psychotria cooperi Standl., respectively; however, we have already absorbed the new nomenclature, and there are no other significant changes to report. This is a rigorous though fundamentally synoptic contribution, featuring synonymy, typology, and distribution summaries for the section and each sp., a detailed description of the section, a dichotomous and indented key to spp., often extensive “Notes,” and a section entitled “Taxon of unclear identity.” There are no indices, and descriptions and specimen citations are provided only for the newly described spp. The introductory portion addresses taxonomic history, the circumscription of the section, morphology, and ecology. Only the new spp. are illustrated (with basic composite line drawings).
Torres-Montúfar, A., H. Flores-Olvera & H. Ochoterena. 2021. A taxonomic treatment of Rogiera (Guettardeae, Rubiaceae). Syst. Bot. 46: 186–210.
The genus Rogiera comprises 10 spp., according to this revision, ranging from Mexico to western Colombia; however, nine of its spp. occur in Mexico, and four are endemic to that country. This is a fine piece of work, but of limited utility to us, because just one sp., Rogiera amoena Planch. (the only member of the genus to extend beyond what has been called "Megamexico") occurs in Costa Rica. The overall geographic range of R. amoena was specified in the Manual Rubiaceae treatment (2014) as "S Méx.–O Pan.," but we must now amend that to "Méx.–O Col." (and ditto for the genus). Features synonymy, typology, and technical descriptions at all ranks, a dichotomous and indented key to spp., distribution and phenology summaries, discussions of varying length, distribution maps, and "representative" (but often extensive) specimen citations. There are no indices. The brief introductory part (featuring a composite color plate of living material) deals almost exclusively with taxonomic history. Each sp. is illustrated with a composite line drawing. No new names or combinations are validated in this paper.
Turner, I. M. 202. Heyne, Roth, Roemer and Schultes, and the plant names published in Novae plantarum species praesertim Indiae orientalis. Taxon 70: 365–428.
All 411 names published by Albrecht Wilhelm Roth (1757–1834) in the work italicized above (published in 1821) are here “listed with their correct citation and typification.” These names were based largely on material collected by Benjamin Heyne (1770–1819), mainly in southern India, and many were published first in the Systema vegetabilium of Roemer and Schultes (to whom Roth had provided a manuscript copy of his book). Due to the provenance of Heyne’s material, this paper is of limited interest to us. Myriads of lectotypes and neotypes are newly designated, of which four apply to the accepted names (or their basionyms) of spp. represented in Costa Rica: Crotalaria spectabilis Roth (Fabaceae), Eleocharis montana (Kunth) Roem. & Schult. (Cyperaceae), Eragrostis rufescens Schrad. ex Schult. (Poaceae), and Gmelina arborea Roxb. ex Sm. (Verbenaceae). Synonyms of the two last-mentioned names are also typified, as are one or more synonyms of each of the following names accepted in the Manual for spp. occurring (at least in cultivation) in Tiquicia: Cyathula prostrata (L.) Blume (Amaranthaceae), Echinochloa colona (L.) Link (Poaceae), Ficus benjamina L. (Moraceae), Fimbristylis cymosa R. Br. (Cyperaceae), Ipomoea aquatica Forssk. (Convolvulaceae), Justicia betonica L. (Acanthaceae), Lindernia crustacea (L.) F. Muell. (Scrophulariaceae), Linum usitatissimum L. (Linaceae), Merremia umbellata (L.) Hallier f. (Convolvulaceae), Nymphoides indica (L.) Kuntze (Menyanthaceae), Ocimum basilicum L. (Lamiaceae), Panicum maximum Jacq. (Poaceae), Rubus niveus Thunb. (Rosaceae), and Urochloa reptans (L.) Stapf (Poaceae). But we have saved the biggest bombshell for last: the only name change (as far as we could discern) required for a member of the Costa Rican flora. This affects the sp. we have known as Panicum rudgei Roem. & Schult. (Poaceae), and was not provoked by any of the typifications debuted in this paper; rather, the author discovered that Roemer and Schultes, in proposing Panicum rudgei as a replacement name for the illegitimate P. scoparium Rudge, had cited the prior Panicum cayennense Lam. in synonymy. That action renders the name Panicum rudgei illegitimate, and a synonym of P. cayennense (the accepted name for a sp. that also occurs in Costa Rica). The oldest available name for the sp. previously known as P. rudgei is Panicum pilosissimum Roth, which we must now get used to.
Uotila, P., A. F. Sukhorukov, N. Bobon, J. McDonald, A. A. Krinitsina & G. Kadereit. 2021. Phylogeny, biogeography and systematics of Dysphanieae (Amaranthaceae). Taxon 70: 526–551.
The “Johnny-come-lately” genus Dysphania has already been debunked as paraphyletic, with respect to the monospecific North American genus Cycloloma. No problem, the lone sp. of Cycloloma is here formally transferred to Dysphania. For Manual users, this requires only that the sp. total for Dysphania be augmented by one (these authors tote 46 spp).