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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XIX, Number 4, October 2012

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

Aedo, C. 2012. Revision of Geranium (Geraniaceae) in the New World. Syst. Bot. Monogr. 95: 1–550.

This monumental contribution accounts for the 137 spp. of Geranium (nearly 40% of the world total) that are either native or well-established in the New World. Just 15 non-native spp. are treated formally. Within the study region, Geranium is most diverse in Mexico and Andean South America, being rather poorly represented in the intervening area. That includes Costa Rica, from which only two Geranium spp. were recorded in the 2010 Manual Geraniaceae treatment by Francisco Morales (INB). The Costa Rican sp. total remains the same, and although a few nomenclatural changes have been made, they have, for the most part, already been noted in these pages: the name Geranium bolivarianum Dayton is to be deleted from synonymy under G. costaricense H. E. Moore [see "Annotate Your Copy" in The Cutting Edge 18(1), Jan. 2011], and the name Geranium guatemalense R. Knuth (accepted in the Manual) becomes a synonym of G. seemannii Peyr. [see "Annotate Your Copy" in The Cutting Edge 18(3), Jul. 2011]. The only new development for us is that two subspp. of G. seemannii, both widespread in the Mesoamerican region, may now be attributed to Costa Rica: the autonymic subsp. and G. s. subsp. repens (H. E. Moore) Aedo (comb. nov.), with somewhat longer petals and fruits. Features complete synonymy and typology and technical descriptions at all ranks, dichotomous and indented, artificial keys to spp. and subspp. (one for North and Central America and another for South America), phenology and distribution summaries, comprehensive specimen citations (with the more lengthy entries deferred to an appendix), discussions, distribution maps, a rather long section on "Doubtful and excluded names," and indices to exsiccatae and scientific names. The sp. entries are nested non-alphabetically under three subgenus headings (with all but 10 of the spp. classed in just one of the subgenera). The introduction highlights taxonomic history, adventive spp. (those that are not well-established), infrageneric divisions (though there is no key to the subgenera), and morphology (with numerous illustrations, including SEM micrographs). Each sp. is illustrated with an excellent composite line drawing. One new combination is validated (mentioned previously), and numerous epi-, lecto- and neotypes are designated.

Borchsenius, F., L. S. Suárez Suárez & L. M. Prince. 2012. Molecular phylogeny and redefined generic limits of Calathea (Marantaceae). Syst. Bot. 37: 620–635.

The results of these analyses confirm previous studies [see, e.g., The Cutting Edge 13(3): 11, Jul. 2006] in portraying Calathea in the traditional sense as polyphyletic, with the smaller of its two major clades (the one including the type sp.) sister to the clade harboring Ischnosiphon and Pleiostachya. Rather than lump the two last-mentioned genera (as well as Monotagma) into Calathea—an option that is scarcely mentioned here and would involve fewer than 80 potential new combinations—the authors choose to split traditional Calathea in twain, even though this means that about 87% of the ca. 285 spp. involved must receive a new genus name. In this regard, the authors are eager to oblige, validating 246 new combinations at sp. rank (all attributed to "Borchs. & S. Suárez") under the obscure generic name Goeppertia Nees, seldom if ever used since it was first published in 1831. The new Calathea s. str. comprises "relatively large plants with tall flowering shoots and inflorescences placed well above the base of the plants"; synapomorphies include "compound inflorescences that usually consist of two to several, similar, partial inflorescences" and "corolla lobes [that] are always reflexed or rolled back." So defined, the following Costa Rican spp. remain in Calathea: Calathea allenii Woodson, C. brenesii Standl., C. carlae H. Kenn., C. crotalifera S. Watson, C. guzmanioides L. B. Sm. & Idrobo, C. lasiostachya Donn. Sm., C. lutea (Aubl.) Schult., C. platystachya Standl. & L. O. Williams, C. recurvata H. Kenn. ("Calathea sp. A" of the Manual), C. retroflexa H. Kenn., C. similis H. Kenn., C. spiralis H. Kenn., C. tarrazuensis H. Kenn., and C. verruculosa H. Kenn. ("Calathea sp. B" of the Manual). Some readers may recognize that, with a few exceptions (Calathea allenii, C. guzmanioides and C. lutea), the spp. in the foregoing list are also characterized by laterally flattened inflorescences with distichous bracts. The remaining 30 Costa Rican spp. (out of 44 total) heretofore treated under the genus name Calathea are henceforth referred to Goeppertia, comprising "plants [that] tend to be smaller in size" and (we would add) having generally cylindrical or fusiform to ellipsoidal, ovoid, or obovoid inflorescences with spiraled bracts; synapomorphies for this group include "a simple inforescence (with a few exceptions…) and corolla lobes straight to spreading…" The only Costa Rican sp. in the latter group for which a combination in Goeppertia does not yet exist is the recently described Calathea jondule H. Kenn. & Hammel [see under "Kennedy," this column, in The Cutting Edge 19(1), Jan. 2012]; also, because no combinations are validated in this paper for infraspecific taxa, those wishing to recognize subspp. or vars. (as was done for Calathea macrosepala K. Schum. and C. silvicola H. Kenn. in Helen Kennedy’s Manual treatment) will apparently have to make the indicated combinations themselves (should they decide to go the Goeppertia route). It seems odd that the authors of this paper did not validate a new combination in Calathea for the Panamanian Sanblasia dressleri L. Andersson (actually, a nomen novum would be required), despite the fact that they formally synonymized the monospecific Sanblasia L. Andersson under Calathea. And even though their results showed that "Ischnosiphon was paraphyletic with respect to Pleiostachya," they refrained from sinking the latter genus, concluding that "further analysis is necessary" (in any case, most names in Pleiostachya already have combinations in Ischnosiphon).

Burt-Utley, K. & J. F. Utley. 2012. New species and notes on Begonia (Begoniaceae) from México and Central America. Phytoneuron 2012-74: 1–25.

Ten new Begonia spp. are described from the Mesoamerican region in this Flora mesoamericana precursor, including four that occur in Costa Rica. Three of the four Costa Rican spp. are evidently endemic: Begonia liesneri Burt-Utley & Utley, collected just twice at 1400–1450 m elevation on the Atlantic slope of the northern Cordillera de Talamanca, compared especially with B. tonduzii C. DC.; Begonia aguabuenensis Burt-Utley & Utley, likewise known from just two collections, at 1000–1300 m in the vicinity of Cerro Caraigres (Fila Aguabuena), and most similar to the Guatemalan B. manicata Brongn. ex F. Cels var. peltata L. B. Sm. & B. G. Schub.; and Begonia sukutensis Burt-Utley & Utley, known only from the type collected at 350–550 m on the Atlantic slope of the eastern Cordillera de Talamanca, compared with B. corredorana C. DC. and B. quaternata L. B. Sm. & B. G. Schub. The fourth new sp. in Costa Rica, Begonia wilburii Burt-Utley & Utley (corrected by us from "wilburi"), has been found at elevations of 1600–3350 m on the Pacific slope of the northern Cordillera de Talamanca, but also occurs disjunctly in western Panama (Prov. Chiriquí); characterized by 1-flowered female inflorescences and "apparent" dioecy, B. wilburii is sharply delimited from its Mesoamerican congeners and "appears most closely related" to B. ursina L. B. Sm. & B. G. Schub., endemic to Colombia. A draft treatment of Begoniaceae for the Manual, submitted some time ago by Alexánder Rodríguez and Armando Estrada (CR), includes six provisionally named and (ostensibly) undescribed spp., just one of which (their "Begonia sp. C") clearly equates with a sp. described in this paper (B. sukutensis). Therefore, it is conceivable that a few additional Begonia spp. remain to be described from Costa Rica. Also to be sought in Costa Rica is another of the spp. described in this paper, Begonia panamensis Burt-Utley & Utley, reported from 1700–2300 m elevation on the Pacific slope of westernmost Panama (Prov. Chiriquí) and most similar to B. copeyana C. DC.; indeed, because B. panamensis has been collected (twice) from Cerro Pando, which straddles the Costa Rican border, it will need to be treated in full for the Manual. All ten new spp. are illustrated with photographic images of type specimens. The "notes" pertain to previously described spp. of Mexico and Guatemala.

Carvalho, F. A. & S. S. Renner. 2012. A dated phylogeny of the papaya family (Caricaceae) reveals the crop’s closest relatives and the family’s biogeographic history. Molec. Phylogen. Evol. 65: 46–53.

The controversial (at least in our besotted brains) separation of Vasconcellea from Carica, leaving the cultivated C. papaya L. as the sole member of the latter genus [see The Cutting Edge 8(1): 3, Jan. 2001], receives emphatic support from this molecular study, analyzing DNA sequences (both nuclear and plastid) from every sp. in the family Caricaceae. Indeed, the two aforementioned genera are not even particularly close relatives, as has generally been supposed; rather, Carica enjoys a sister-group relationship with Horovitzia and Jarilla, mono- or oligospecific (respectively) genera of northern Mesoamerica (Jarilla being herbaceous). Of course, none of this mandates the segregation of Vasconcellea from Carica, though the lumping alternative would now involve adding not only Horovitzia and Jarilla to the mix, but Jacaratia as well (almost all the spp. involved already have names in Carica, by the way). We must admit that our review of a previous molecular investigation [see, e.g., The Cutting Edge 12(3): 6, Jul. 2005] took somewhat out of context the reported lack of morphological evidence to support the separation of Vasconcellea from Carica; in an attempt to find ways to identify sterile material, that study allowed only vegetative characters. The rather obvious difference of 5-locular vs. unilocular ovaries, pointed out in the present paper, makes the distinction somewhat more tolerable for us. On the other hand, the repeated mention of "wild" or "natural" populations of Carica papaya in Costa Rica is cause for consternation on the part of your editors [see under "Brown" et al., this column, in The Cutting Edge 19(2), Apr. 2012]. Now we are experiencing a crisis of confidence: are the populations (e.g., in Parque Nacional Carara) that we have assumed to represent Carica cauliflora Jacq. [okay, Vasconcellea cauliflora (Jacq.) A. DC.] nothing more than wild populations of C. papaya? If so, then what is the real C. cauliflora? And if not, then where are these wild populations of C. papaya? We promise to get this resolved (at least for our own sakes!) in time for the publication of the Manual Caricaceae treatment (in Vol. 4).

Chakrabarty, T. & N. P. Balakrishnan. 2009. Transfer of Indian species of Breynia, Glochidion and Sauropus to Phyllanthus (Phyllanthaceae) – new combinations and new names. J. Econ. Taxon. Bot. 33: 712–716.

Well, now we have seen it with our own eyes and can vouch for the terrible truth: as we previously reported (see under "Annotate Your Copy" in our last issue), the combination Phyllanthus androgynus (L.) J. A. González, validated in Manual Vol. 5 (p. 376), is an illegitimate isonym, predated (by almost exactly a year) by the identical combination appearing in this paper and attributed to its authors. Why it took nearly three years for this issue to reach our shores is anyone’s guess. Evidently, a similar period of time elapsed before it arrived in the hands of the people in charge of IPNI (which we had been checking regularly for this very name). This begs the question: how reliable, in determining the date of actual publication, is the date printed on a journal or book?

Dehgan, B. 2012. Jatropha (Euphorbiaceae). Fl. Neotrop. Monogr. 110: 1–274.

A lifetime of research on the genus Jatropha comes to full fruition with this effort, covering the 86 New World spp. (about two-thirds of the world total) in this nominally pantropical genus. The sp. entries are arranged according to a finely subdivided infrageneric classification which includes (at least in the New World) two subgenera, eight sections (one described here as new), and 13 subsections (six described as new), except for two still unclassified spp. deferred to an "Addendum." From a Costa Rican perspective, the situation at sp. rank is little changed from the 2010 Manual Euphorbiaceae treatment by José González (LSCR), which is not cited here and evidently was not seen by the author. In fact, José may have been slightly ahead of the game, as he included Jatropha stevensii G. L. Webster, portrayed in this monograph as a Nicaraguan endemic, based on Costa Rican material apparently not available to this author. Two other minor departures from José’s treatment are worth mentioning, both involving infraspecific classification: Costa Rican material of Jatropha gossypiifolia L. may now be qualified as belonging to J. g. var. staphysagriifolia (Mill.) Müll. Arg. (widespread from Mexico to Ecuador), while Jatropha integerrima Jacq. (mentioned in José’s genus discussion as occasionally cultivated) is represented in Costa Rica by its autonymic var. Features synonymy, typology, and technical descriptions at all ranks, a dichotomous and indented key to infrageneric taxa and similar keys (nested below the sections or subsections) to spp. and infraspecfic taxa (the latter being also keyed below the appropriate spp.), distribution summaries, discussions, representative specimen citations, a lengthy section on "Doubtful names and excluded taxa" (mostly names now referred to Cnidoscolus or Manihot), and indices to exsiccatae, common names, and scientific names. The generously illustrated and tabularized introductory section addresses taxonomic history, morphology and anatomy, pollination, karyology, distribution and evolution, and relationships. Most of the spp. and infraspecific taxa (and all of those occurring in Costa Rica) are illustrated with composite line drawings. N.B.: we have followed IPNI (not to mention Müller Argoviensis) in correcting the original staphysagrifolia (used in this monograph) to staphysagriifolia.

De Laubenfels, D. J. 2009. Nomenclatural actions for the New World cypresses (Cupressaceae). Novon 19: 300–306.

This paper, in our own in-house journal, somehow slipped through the cracks for us (see under "Lombardi," this column). It’s all just as well, however, as the main intent was to propose a new genus name, Neocupressus de Laub., for most of the New World spp. that had previously been included in Cupressus. It turns out that the author was beaten to the punch on this by about five months and that Hesperocyparis Bartel & R. A. Price is the correct name for the taxon in question [see under "Adams," this column, in The Cutting Edge 17(1), Jan. 2010]; Neocupressus is accordingly illegitimate, as are all of the combinations made thereunder at infrageneric rank.

——, D. J., C. E. Husby & M. P. Griffith. 2012. Further nomenclatural action for the cypresses (Cupressaceae). Novon 22: 8–15.

Confusion and disagreement continue to reign supreme in the group of plants formerly lumped under the genus name Cupressus, perhaps in part because most of the active participants appear to have horticultural backgrounds. We are assuming that Hesperocyparis lusitanica (Mill.) Bartel (see the previous entry) remains the correct name for the sp. (cultivated in Costa Rica) treated as Cupressus lusitanica Mill. in the Manual. However, the combination H. l. var. lindleyi (Klotzsch ex Endl.) de Laub. is here validated, based on a name that was synonymized under C. l. var. lusitanica in the Manual; H. l. var. lindleyi is said to differ from the autonymic var. by its "thicker branchlets (1.5–2.1 mm vs. 1.2–1.5 mm) and the larger seed cones (15–23 mm vs. 10–18 mm)." Even more disconcerting, another name synonymized (under Cupressus lusitanica) in the Manual, Cupressus benthamii Endl., is characterized as quite distinct morphologically from Hesperocyparis and possibly belonging either to Callitropsis or Cupressus s. str.; no firm position is taken on the matter, although all the possible combinations have been made. It is difficult to comprehend which of these taxa may be represented in Costa Rica, especially as none of these authors seems inclined to annotate or even cite herbarium material.

Donnell, A. A., H. E. Ballard, Jr. & P. D. Cantino. 2012. Callianthe (Malvaceae): a new genus of Neotropical Malveae. Syst. Bot. 37: 712–722.

The new genus of the title was revealed in an effort to define the boundaries between Abutilon and Bakeridesia, two genera of Malvaceae (in the traditional, narrow sense) that have been suspected of being polyphyletic. Such has proven to be the case, with "most of the pluriovulate species of Abutilon…and the species of Bakeridesia subgenus Dipteron together form[ing] a previously unrecognized clade…described here as a new genus, Callianthe Donnell." The new genus, as it turns out, is more closely related to the likes of Briquetia and Hochreutinera than to either Abutilon or Bakeridesia in their new, restricted senses. Callianthe is formally described and compared in tabular form with Abutilon, Bakeridesia, and Gaya (its closest relative); a dichotomous, indented key is also provided to separate Abutilon, Bakeridesia, and Callianthe. Forty new combinations at sp. rank are validated (all in the name of the first author) in Callianthe, with 11 of these (by our count) coming from Bakeridesia and the rest from Abutilon. An additional 21 names (all but one in Abutilon) are listed "that will likely be transferred to Callianthe once more source information is available and/or nomenclatural issues are resolved"; thus, the genus may be expected to comprise perhaps 60 spp. Callianthe is principally a South American taxon, so the consequences for Costa Rica are minimal; Bakeridesia vulcanicola (Standl.) D. M. Bates, our sole representative of its genus, is unaffected, and just two naturally occurring Costa Rican Abutilon spp. are transferred to Callianthe: Abutilon purpusii Standl. and A. striatum G. F. Dicks. ex Lindl. (a combination is also validated for A. brenesii Standl., but that name was synonymized under A. purpusii in the Manual Malvaceae treatment by the late Paul Fryxell). The so-called "flowering maples" of commerce also belong in Callianthe, to which Abutilon vexillarium E. Morren, mentioned in the Manual genus discussion of Abutilon, has accordingly been transferred (as "Callianthe vexillaris," which we think must be corrected to C. vexillaria). Incidentally, the results of this study "further confirm that Bastardia and Bastardiopsis are nested among Abutilon species in the well-supported Abutilon s. s. clade"; however, for reasons that are not explained, the indicated new combinations in Abutilon are not validated for this group, meaning that, at least for the time being, we are stuck with Bastardia viscosa (L.) Kunth in the Costa Rican flora.

Eshbaugh, W. H. 2012. The taxonomy of the genus Capsicum. Pp. 14–28 in, V. M. Russo (ed.), Peppers: botany, production and uses. CABI, Oxfordshire, UK/Cambridge, MA. 280 pp.

This is the obligatory taxonomic chapter in a book otherwise devoted almost entirely to agronomic topics. Thus, it is not surprising that the five spp. of Capsicum (Solanaceae) most important in cultivation are the main focus here. These five spp. are distinguished in a dichotomous key, and each is briefly characterized. The nomenclature employed by the author agrees in virtually every respect with that used by Lynn Bohs (UT) in the Manual Solanaceae draft treatment. All 36 Capsicum spp. accepted by the author (an established authority on the genus) are listed in a table that includes their New World distributions and chromosome numbers, and the "hypothetical distribution of domesticated Capsicum peppers at the time of European discovery of the New World" is mapped for the five principal spp. Relationships within the genus are expressed by means of a cladogram (based on molecular analyses) and hybridization chart. The relationship among three of the main cultivated spp.—Capsicum annuum L., C. chinense Jacq., and C. frutescens L.—"has not been resolved," and it is possible that these may eventually be combined in a single sp. (under the name C. annuum).

Fuentes-Bazan, S., P. Uotila & T. Borsch. 2012. A novel phylogeny-based generic classification for Chenopodium sensu lato, and a tribal rearrangement of Chenopodioideae (Chenopodiaceae). Willdenowia 42: 5–24.

Chenopodium, once boasting about 100 spp., has been reduced to (we think) about 60 spp. by recent taxonomic changes, including those realized in this article. Of course, we were already familiar with the removal of C. ambrosioides L. (and related spp.) to the genus Dysphania [see The Cutting Edge 15(4): 5–6, Oct. 2008], which is the only change that directly affects Costa Rica, since C. album L., the only other sp. recorded from the country, is the generic type and not going anywhere. But we are gratified to finally see the cladograms on which these taxonomic decisions have been predicated. The present authors now accept a total of seven genera (one described as new) for Chenopodium in its traditional sense, and validate all the necessary new combinations at sp. and subsp. rank. Perusing the cladograms, we concur that they had little choice and made the best possible decision, with the information available; the major constraint is clearly the nesting within Chenopodium s. l. of Atriplex, a well-known and well-marked genus with upwards of 250 spp. (and also, to a lesser extent, the nesting of the much smaller but also well-known genus Spinacia). Includes a dichotomous (though non-indented) key to genera, synonymy and typology for each genus (and also each sp. and subsp. name for which a new combination is validated), and technical descriptions of the genera (it would have been nice to have sp. totals and distributions indicated as well). The genera are arrayed in four different tribes, and while tribal classification is officially beyond our pale, we are surprised to see spp. that were included in the same genus no more than a decade ago (and in some cases, up till this moment!) now relegated to different tribes.

Gomes-da-Silva, J., F. A. C. Vargens, R. C. O. Arruda & A. F. Costa. 2012. A morphological cladistic analysis of the Vriesea corcovadensis group (Bromeliaceae: Tillandsioideae), with anatomical descriptions: new evidence of the non-monophyly of the genus. Syst. Bot. 37: 641–654.

This study, involving a mere 23 spp., suggests that the large (ca. 280 spp.) genus Vriesea (apparently excluding recent segregates such as Werauhia) is paraphyletic, with spp. generally assigned to other genera (e.g., Guzmania and Mezobromelia) nested within it. The authors, fully cognizant of the limitations of their research, venture no nomenclatural changes.

Harley, R. M. & J. F. B. Pastore. 2012. A generic revision and new combinations in the Hyptidinae (Lamiaceae), based on molecular and morphological evidence. Phytotaxa 58: 1–55.

Our review of a previous paper helmed by these same authors [see under "Pastore" et al., this column, in The Cutting Edge 19(1), Jan. 2012] indicated that Hyptis had been revealed as paraphyletic, with Asterohyptis and Marsypianthes (among other genera) nested within, and that further splitting of Hyptis was looming as the likely resolution for this dilemma. We listed "at least eight" Costa Rican spp. that figured to remain in Hyptis under the splitting scenario, and six others that "would have to be transferred to at least three segregate genera." Now that the fan has been hit and all the new combinations validated, we find that the best-case scenario has prevailed for Costa Rican floristics: all five of the spp. unaccounted for in our lists (of the total of 19 Hyptis spp. treated in Amy Pool’s Manual Lamiaceae account) will remain in Hyptis, these being H. brachiata Briq., H. obtusiflora C. Presl ex Benth., H. personata Epling, H. savannarum Briq., and H. vilis Kunth & C. D. Bouché; and we will have to contend with just three segregate genera, the minimal possibility. As an arguable bonus, Asterohyptis and Marsypianthes are preserved. The subdivision of Hyptis (now reduced to "around 144" spp.), proceeding largely (though by no means exclusively) along sectional lines, yielded a total of 11 splinter genera, as presaged in the earlier paper. Those of us who work on the Costa Rican flora will need to get used to the following three newly deployed genus names: Cantinoa Harley & J. F. B. Pastore [including the former Hyptis mutabilis (Rich.) Briq. and H. spicigera Lam.], Condea Adans. (including the former Hyptis verticillata Jacq.), and Mesosphaerum P. Browne [including the former Hyptis pectinata (L.) Poit., H. suaveolens (L.) Poit., and H. urticoides Kunth, as well as H. oblongifolia Benth., mentioned in the Manual genus discussion]. The new combinations are all validated in the names of the authors, and all are straightforward except in the case of Hyptis spicigera, which becomes Cantinoa americana (Aubl.) Harley & J. F. B. Pastore (the prior epithet americana, blocked in Hyptis, being again available in Cantinoa). Features a dichotomous (though non-indented) key to the 19 genera of subtribe Hyptidinae, an informal diagnosis of each genus, synonymy and typology for each genus and the sp. for which new names or combinations are validated, and an extremely useful appendix indicating the current status of each "earlier name." Four pages of photos of live plants of selected spp. are included at the end of the paper, along with a rather crude cladogram (allowing one to ponder possible alternatives to the system proposed here).

Inda, L. A., M. Pimentel & M. W. Chase. 2012. Phylogenetics of tribe Orchideae (Orchidaceae: Orchidoideae) based on combined DNA matrices: inferences regarding timing of diversification and evolution of pollination syndromes. Ann. Bot. (Oxford) 110: 71–90.

Just one incidental conclusion of this study is of interest to us: "Habenaria…is grossly paraphyletic, with Gennaria, Herminium and Pecteilis embedded in it." The three embedded genera, with a combined total of about 40 spp. (mostly in Herminium), are all restricted to the Old World.

Karremans, A. P. & M. Muñoz García. 2011. Pleurothallis silvae-pacis, a new species. Orchid Rev. 119: 155–159.

Pleurothallis silvae-pacis Karremans hails from 1500–2000 m elevation in the private, Bosque de Paz Biological Reserve [see The Cutting Edge 10(3): 6–7, Jul. 2003], on the Atlantic slope of Volcán Poás. The new spp. is believed most closely related to the Ecuadorian Pleurothallis wigginsii C. Schweinf., and among Costa Rican spp. is most similar to P. eumecocaulon Schltr. and P. imitor Luer. The three last-named spp. have been transferred to the segregate genera Acronia (P. imitor and P. wigginsii) and Ancipitia by Luer [see The Cutting Edge 11(3): 10–11, Jul. 2004]; however, these authors question the distinction of those taxa even at sectional rank, so class their novelty conservatively in Pleurothallis. Illustrated with a composite line drawing (by the first author) and two close-up color photos of flowers in life

—— & C. M. Smith. 2012. A note on genus Dracontia (Orchidaceae: Pleurothallidinae), with a new species. Harvard Pap. Bot. 17: 13–17.

Progress continues on the genus Dracontia (see under "Karremans," this column, in our last issue), a recent segregate of Pleurothallis in the sense of Carlyle Luer’s (2003) Manual treatment of the latter genus, inspired by the first author’s unpublished Wageningen University Master’s thesis (2010) on Stelis "and closely related genera." The new sp. is Dracontia hydra Karremans & C. M. Sm., based on six specimens, all collected by the usual JBL crew at 1050–1550 m elevation on the Atlantic slope of the northern Cordillera de Talamanca. Evidently, no one else has stumbled upon it in this heavily botanized region. The comparison is with Dracontia conochila (Luer) Luer (Pleurothallis conochila Luer of the Manual), from which D. hydra differs in the size (larger), morphology, and pubescence of its flowers. Our gain of one sp. is offset by the loss of another, as Dracontea perennis (Luer) Luer (Pleurothallis perennis Luer of the Manual) is revealed as conspecific with D. dracontea (Luer) Luer (Pleurothallis dracontea Luer of the Manual), the latter being the older name. Illustrated with composite line drawings of both Dracontia dracontea and D. hydra.

Koenen, E. J. M. & J. J. F. E. de Wilde. 2012. A taxonomic revision of the reinstated genus Leplaea and the newly recognized genus Neoguarea (Meliaceae, Sapindales): the exclusion of Guarea from Africa. Pl. Ecol. Evol. 145: 209–241.

The Western Hemisphere gains an endemic genus! The Manual treatment of Guarea (2007) by Alexánder Rodríguez (INB) dutifully included five African spp. in the generic total of "Ca. 75," following the reigning authorities on the group. However, morphological studies by the present authors, allegedly "supported by molecular data" (not presented here), reveal that the African contingent is in fact distinct. As a result, the African spp. previously included in Guarea are transferred to the two other genera mentioned in the title (Neoguarea being upgraded from sectional rank), with two spp. of Leplaea described as new (eight spp. total are involved). So the Eastern Hemsiphere (Africa, specifically) actually gains two endemic genera in this deal. Interestingly, these African spp. lack the one character that makes Guarea so easily identifiable in sterile condition in the New World: paripinnate leaves with a terminal bud.

Lehnert, M. 2012. A synopsis of the species of Cyathea (Cyatheaceae-Polypodiopsida) with pinnate to pinnate-pinnatifid fronds. Phytotaxa 61: 17–36.

This represents the second installment in a series of synoptic treatments of neotropical Cyathea and, like its predecessor [see under this author, this column, in The Cutting Edge 18(4), Oct. 2011], is useful to us mainly for the dichotomous keys to spp. and vars. (compromised by being non-indented). We count 57 spp. that are treated here, or rather listed (alphabetically), with synonymy, typology, and distribution summaries. Many of the spp. in the study group were formerly (as in Flora mesoamericana) included in the genus Cnemidaria, recently submerged in Cyathea [see, e.g., The Cutting Edge 13(4): 9–10, Oct. 2006]. As was the case with the first paper in the series, there is little that is new here for Costa Rican floristics, and no taxa are added to the flora. New combinations in Cyathea are validated in the author’s name for several taxa transferred from Cnemidaria, including (for Costa Rica) the former Cnemidaria cocleana Stolze and Cnemidaria mutica (Christ) R. M. Tryon var. grandis (Maxon) Stolze. Cnemidaria mutica vars. chiricana (Maxon) Stolze and contigua (Underw. ex Maxon) Stolze sensu Flora mesoamericana are here spun off to a separate sp., Cyathea chiricana (Maxon) Domin, under which a new combination is validated for var. contigua. Also represented in Costa Rica among this group are Cyathea choricarpa (Maxon) Domin, C. horrida (L.) Sm., and C. mutica (Christ) Domin var. mutica, all erstwhile members of Cnemidaria, as well as Cyathea stolzei A. R. Sm. ex Lellinger and C. ursina (Maxon) Lellinger.

Lombardi, J. A. 2009. Three new species of Salacia from Mesoamerica (Celastraceae, Salacioideae). Novon 19: 372–379.

We had this issue of Novon marked for review, but it must have gotten buried among our papers and overlooked. All three of the Salacia spp. described in this paper occur in Panama, but only Salacia mennegana Lombardi makes it to Costa Rica (as well as western Colombia). The name S. mennegana is attributed to "J. Hedin ex Lombardi," but we have deleted the "ex" citation because a different spelling (mennegae) was suggested in Manual contributor Jennifer Hedin’s Ph.D thesis (where the designation "Salacia sp. nov. A" was used in the sp. heading). The new sp. is compared to two South American congeners, Salacia opacifolia (J. F. Macbr.) A. C. Sm. (the name used in Hedin’s Manual draft, yet to be published) and S. insignis A. C. Sm. Salacia mennegana is known from Costa Rica by just two collections, both made by Danish botanist Karsten Thomsen on the Península de Osa. All three new spp. are depicted in composite line drawings.

López, A. & O. Morrone. 2012. Phylogenetic studies in Axonopus (Poaceae, Panicoideae, Paniceae) and related genera: morphology and molecular (nuclear and plastid) combined analyses. Syst. Bot. 37: 671–676.

Axonopus is paraphyletic, but not in a manner that portends trouble for Costa Rican floristics. Two small Brazilian genera, Centrochloa (one sp.) and Ophiochloa (two spp.), are nested within and summarily transferred to Axonopus, with the three new combinations at sp. rank validated by the authors. So just bump up the sp. total in the Manual accordingly.

Lovo, J., R. C. Winkworth & R. Mello-Silva. 2012. New insights into Trimezieae (Iridaceae) phylogeny: what do molecular data tell us? Ann. Bot. (Oxford) 110: 689–702.

The news is not good for Neomarica, the only genus of tribe Trimezieae occurring naturally in Costa Rica, with spp. falling into three of the principal clades resolved by these analyses. Thus Neomarica is undoubtedly polyphyletic, but the authors forego taxonomic revision pending "further molecular analyses." We cannot even guess as to the potential fate of N. variegata (M. Martens & Galeotti) Henrich & Goldblatt, our only indigenous sp., which was not included in the study; however, N. caerulea (Ker Gawl.) Sprague, cultivated in Costa Rica, groups rather closely with the generic type (in one of the anaylses, at least). Similarly polyphyletic (with spp. split among three clades) is the genus Trimezia; however, T. steyermarkii R. C. Foster, cultivated and sparingly naturalized in Costa Rica (see under "Leaps and Bounds"), was not sampled

Morales, [J.] F. 2012. Nuevas especies de Sapotaceae para Costa Rica. Darwiniana 50: 107–113.

The names Chromolucuma cespedesiiformis J. F. Morales and Pouteria spicata J. F. Morales are formally made available for two spp., both endemic to the humid lowlands of the southern Pacific slope in Costa Rica, that were treated under provisional names (Chromolucuma sp. A and Pouteria sp. E, respectively) in the author’s Manual draft of Sapotaceae. The newly christened spp. are compared most closely with the South American Chromolucuma rubriflora Ducke and the sympatric endemic Pouteria triplarifolia Standl. & L. O. Williams ex T. D. Penn. (respectively), and both are illustrated with composite line drawings. Dichotomous (though non-indented) keys are provided for all the Chromolucuma spp. [including the recently combined C. congestifolia (Pilz) Alves-Araújo & M. Alves; see under "Alves-Araújo," this column, in our last issue] as well as for an unspecified subset of the Pouteria spp. (including P. spicata) of Nicaragua, Costa Rica, and Panama, and a distribution map does double duty for both novelties. Note that we have corrected the epithet of the new Chromolucuma from the original "cespedisiiformis," which embodies an orthographical error [the sp. being named for its resemblance to juvenile plants of the genus Cespedesia (Ochnaceae)]. We enclose the author’s first initial in brackets because it is omitted from the by-line (though included in both versions of the abstract).

Motes, M. 2011. Eric Christenson (1956–2011). Orchid Rev. 119: 75.

While about 85% of this obituary is appropriately reverent, we are inevitably drawn to the opening paragraph, in which the author shoots from the hip in a most colorful fashion: "In his abbreviated career, Eric cut a broad swathe through the staid world of taxonomy. A Rabelaisian or Falstaffian figure, many will learn of his demise with mixed emotions. Notorious for quickly taking umbrage, Eric would fire off obscenity-laced missives to those who irritated him. By the end of his career Eric had managed to offend the vast majority of his peers." Let it be said that we were not among those who Eric had managed to offend, and in fact got along quite well with him during our limited interactions over the years (mainly when Eric was the editor of Selbyana); however, we had heard the rumors about his darker side, and are most grateful to have been spared!

Nicolas, A. N. & G. M. Plunkett. 2012. Untangling generic limits in Azorella, Laretia, and Mulinum (Apiaceae: Azorelloideae): insights from phylogenetics and biogeography. Taxon 61: 826–840.

The only genus of the three mentioned in the title recorded from Costa Rica is Azorella, spp. of which occur in each of the seven major clades discriminated in this analysis. To summarize, we can do no better than to quote ourselves, from a review in these pages of another recent article by these same authors [see under their names, this column, in The Cutting Edge 17(2), Apr. 2010]: "Azorella appears polyphyletic, with the generic type (A. filamentosa Lam.) in a clade far removed from that harboring A. biloba (Schltdl.) Wedd., the sole Costa Rican representative." A taxonomic resolution, still on the horizon, will involve "major changes in generic circumscriptions and the possibility that new genera may need to be erected or that existing ones may have to be subsumed."

Pupulin, F., A. P. Karremans & B. Gravendeel. 2012. A reconsideration of the empusellous species of Specklinia (Orchidaceae: Pleurothallidinae) in Costa Rica. Phytotaxa 61: 1–20.

It turns out that the novel adjective "empusellous" derives from the monospecific genus Empusella (Luer) Luer, one of numerous new segregates created by its author in his wholesale deconstruction of traditional Pleurothallis [see The Cutting Edge 11(3): 10–11, Jul. 2004]. The sole member of Empusella is the sp. that was treated in Carlyle Luer’s (2003) Manual contribution under the name Pleurothallis endotrachys Rchb. f. The present authors opt for a compromise resolution that synonymizes Empusella under Specklinia, while acknowledging the "different interpretations of the circumscription" of the latter genus. However, they do recognize an "empusellous" clade that includes 14 spp., characterized generally by "intense bright orange flowers with a fruity odor," the coloration being "unique within Specklinia, uncommon in Pleurothallidinae, and not at all common among Orchidaceae in general." Rather than being hummingbird-pollinated, as is aparently the case with most orange-flowered orchid spp., flowers of the "empusellous" group are pollinated by fruit flies (Drosophila). Four "empusellous" spp. are dealt with in this paper, all of which occur in Costa Rica and were subsumed under Pleurothallis endotrachys in the Manual, and one of which is described as new. The new sp. is Specklinia remotiflora Pupulin & Karremans, known from numerous collections (hooray!) from ca. 1750–2500 m elevation on both slopes of the eastern Cordillera de Talamanca, as well as Panama and Colombia. It is compared, quite logically, with S. endotrachys (Rchb. f.) Pridgeon & M. W. Chase. Two other segregates from the last-mentioned sp., Specklinia pfavii (Rchb. f.) Pupulin & Karremans (comb. nov.) and S. spectabilis (Ames & C. Schweinf.) Pupulin & Karremans (comb. nov.), occur at lower elevations, the former at ca. 500 m elevation in "southern Costa Rica and western Panama" (though no collections with explicit locality data are cited), the latter at 450–1350 m elevation from southern Mexico (Chiapas) to Panama (with the cited Costa Rican collections hailing from the Atlantic slope of the Cordillera de Central, the Llanura de Santa Clara, and the Pacific slope of the Cordillera de Tilarán). Their basionyms in Pleurothallis both appear in synonymy under P. endotrachys in the Manual, together with P. platyrachis (Rolfe) Rolfe, here synonymized under Specklinia pfavii. The real Specklinia endotrachys sensu stricto (of these authors) is a Costa Rican endemic, limited to elevations of 700–1100 m in the Cordilleras de Guanacaste and de Tilarán ("from the Pacific watershed," according the the authors, but three of the four cited collections with locality data are explicitly from the Atlantic slope). Features a dichotomous key to the four spp. already mentioned, along with full synonymy and typology for each, technical descriptions, specimen citations, distribution summaries, notes, and distribution maps. Superlative composite line drawings are provided for all four spp., and fresh flowers of each are depicted photographically (as are some of the pollinators!).

Sánchez Agudo, J. Á., M. M. Martínez-Ortega, S. Cafferty & E. Rico. 2012. A contribution toward clarifying the nomenclature of Veronica L. (Plantaginaceae). Taxon 61: 867–870.

Eighteen names in Veronica are epi-, lecto-, or neotypified, in such a way as to "support the current use of the names concerned and therefore contribute to establishing the correct nomenclature of the genus." Neotypified thusly is Veronica persica Poir., the only name dealt with that is accepted for a sp. occurring in Costa Rica (though two synonyms of V. arvensis L. are lectotypified in like manner)

Schubert, H. K., M. S. Taylor, J. F. Smith & A. J. Bornstein. 2012. A systematic revision of the genus Manekia (Piperaceae). Syst. Bot. 37: 587–598.

We had long since adjusted to the substitution of the name Manekia for the genus we had learned many years ago as Sarcorhachis [see The Cutting Edge 6(1): 7, Jan. 1999] and, with just one sp. involved in Costa Rica, we were expecting no surprises here. But we are surprised nonetheless: the transformation is now complete for our Sarcorhachis naranjoana (C. DC.) Trel., briefly Manekia naranjoana (C. DC.) Callejas ex N. Zamora, Hammel & Grayum, now M. incurva (Sieber ex Schult.) T. Arias, Callejas & Bornst. The final stage of this transformation may be explained, frankly, by the fact that all the new combinations in Manekia were rushed into print before a taxonomic revision of the group had been undertaken. Said revision has now been accomplished and, through the magic of "canonical discriminant analyses," six spp. have become just three. One of the casualties, needless to say, is our M. naranjoana, formerly conceived as restricted to southern Central America but now extended to Peru, Venezuela, and the Lesser Antilles, whence the prior M. incurva (based on a specimen from Martinique). Features synonymy and typology, technical descriptions, distribution summaries, and "notes" for both the genus and spp., a dichotomous and indented key to spp., a phenology summary and comprehensive specimen citations for each sp., and a distribution map. There are no illustrations or indices.

Smith, C. M. & F. Pupulin. 2012. Contributions to a taxonomic revision of Masdevallia (Orchidaceae: Pleurothallidinae) in Costa Rica. A note on Masdevallia attenuata. Harvard Pap. Bot. 17: 25–38.

The conclusions of this study mandate some name changes for Costa Rican Masdevallia spp., but result in no net change in sp. total. These authors demonstrate convincingly that, contrary to a presumption by Manual Orchidaceae contributor Carlyle Luer, the name Masdevallia attenuata Rchb. f. (1871) cannot be said to have a holotype; rather, it was described on the basis of two syntypes, requiring that lectotypification be effected. That is accomplished in this article, in which the only element (a drawing) that "really corresponds to the protologue" is designated as the lectotype of M. attenuata. This seemingly benign action affects the names of two spp. in the Costa Rica flora, because, as it turns out, the two original syntypes represent two different spp., and the new lectotype is not the same element as Luer’s presumed holotype. In other words, Luer has been applying the name Masdevallia attenuata incorrectly (according to the new lectotypification). The lectotype of M. attenuata, according to these authors, actually corresponds to Masdevallia scabrilinguis Luer (1979; characterized by its author as "similar to M. attenuata"), which thus falls into synonymy. That would leave Masdevallia attenuata sensu Luer without a name, were it not for the existence of Masdevallia fonsecae Königer (included in synonymy under M. attenuata in the Manual), here consecrated as the correct name for the sp. So, in the long-festering polemical war between Königer and Luer [see The Cutting Edge 10(3): 7, Jul. 2003], chalk one battle up for Königer. To summarize for Manual users: the sp. called Masdevallia attenuata in Vol. 3 becomes M. fonsecae (and "Ecua." should evidently be deleted from the distribution summary), while the sp. called M. scabrilinguis is now M. attenuata (with M. scabrilinguis as a synonym). By the way, the spp. dealt with in this paper would fall within Luer’s recent segregate genus Acinopetala [see The Cutting Edge 13(4): 5–6, Oct. 2006]; however, as that has not received "unanimous support" and "some of the species…discussed here [i.e., Masdevallia fonsecae] have no combination in Acinopetala," the authors retain the traditional classification. Illustrated with photographic reproductions of original and historical material, as well as composite line drawings and photographs of live flowers of both Masdevallia attenuata (in the sense of its new lectotype!) and M. fonsecae.

Snow, N. & P. M. Peterson. 2012. Nomenclatural notes on Dinebra, Diplachne, Disakisperma and Leptochloa (Poaceae: Chloridoideae). Phytoneuron 2012-71: 1–2.

This brief contribution validates new combinations, in several recent segregates of Leptochloa, that were overlooked (accidentally or deliberately) in a previous paper involving the same authors (see under "Peterson," this column, in our last issue). Germane to Costa Rican floristics are the combinations in Dinebra and Diplachne (respectively) for infraspecific taxa of the spp. treated in the Manual as Leptochloa panicea (Retz.) Ohwi and L. fusca (L.) Kunth. These combinations were made at the rank of subsp. in the previous paper, but at the rank of var. in the present one. One is left with the impression that the authors wanted to stamp their names on all the possible new combinations, but were prevented by ICBN Art. 34.2 from doing so in a single paper. Devious indeed! For the record, the rank of subsp. was employed, in each of these cases, in the Manual Poaceae treatment by Francisco Morales (INB)

Sousa S., M. & S. Sotuyo. 2012. El género Muellera L. f. en Mesoamérica y norte de Sudamérica. Acta Bot. Mex. 100: 15–40.

We recently reported on a molecular study by a Brazilian group [see under "Silva" et al., this column, in The Cutting Edge 19(2), Apr. 2012] in which the genus Muellera (Fabaceae), traditionally (as in the Manual) considered dispecific, was expanded to accommodate 24 additional spp., most of which were transferred from Lonchocarpus. We indicated that Muellera frutescens (Aubl.) Standl. [called M. monilis (L.) M. J. Silva & A. M. G. Azevedo by the Brazilian workers] remained the only Costa Rican representative of its genus. The present paper, accepting apparently the same expanded circumscription of Muellera (but with some original molecular evidence), treats 13 spp. from the region specified in the title, three of which are described as new. Again, the only sp. explicitly attributed to Costa Rica is M. frutescens (that name being reinstated in place of M. monilis, but with no explanation as to why the decision was made). Lonchocarpus unifoliolatus Benth. is also considered to belong to Muellera (with the combination duly validated in the name of "M. Sousa"); but although that sp. was included in Manual co-PI Nelson Zamora’s (INB) Fabaceae treatment as occuring in "S Méx., CR y Pan." (with a Costa Rican voucher cited), it is here characterized as a Mexican endemic. Unfortunately, the Manual usage of the name L. unifoliolatus is not addressed, nor has the determination of our Costa Rican voucher been changed in TROPICOS; we can only imagine that these authors regard the name Lonchocarpus monofoliaris Schery (type from Prov. Bocas del Toro, Panama), cited in synonymy under L. unifoliolatus in the Manual, as applying to a distinct sp. (in a different genus!). Features synonymy (at sp. rank) and typology, a detailed description of the genus, descriptions and specimen citations for the new spp. and selected others, and a dichotomous, indented key to the spp. Selected spp. (including the new ones) are depicted in crude composite line drawings.

Verloove, F. & E. Sánchez Gullón. 2012. A taxonomic revision of non-native Cenchrus s.str. (Paniceae, Poaceae) in the Mediterranean area. Willdenowia 42: 67–75.

We were surprised to see Cenchrus incertus M. A. Curtis (1835), an accepted name in the Manual, included in synonymy under C. spinifex Cav. (1799). However, reading the fine print, we learn that the correct application of the latter name "is not uncontested." Furthermore, even as recent evidence (second-hand!) is cited in support of the proposed synonymy, influential authorities on New World grasses are quoted as favoring conservation of C. incertus, should the need arise. Under the circumstances, we’ll put our money on C. incertus prevailing. The authors also attribute Cenchrus longispinus (Hack.) Fernald to Central America, but cite no specimens from the region, and we were unable to scare up any specimen records or literature reports to substantiate the notion.


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