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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XV, Number 1, January 2008

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

Bachelier, J. B. & P. K. Endress.  2007. Development of inflorescences, cupules, and flowers in Amphipterygium and comparison with Pistacia (Anacardiaceae).  Int. J. Pl. Sci. 168: 1237–1253.

This study, comprising “the first detailed account of the branching pattern of the female inflorescence and the structure of the samaroid unit” of the small, Mesoamerican genus Amphipterygium, “strongly supports” the inclusion of this taxon in Anacardiaceae.  It likewise supports our decision to treat Amphipterygium in Anacardiaceae for the Manual, rather than in the small, splinter family Julianiaceae, to which it has often been assigned [see, e.g., The Cutting Edge 14(4): 5–6, Oct., 2007].  Lucky thing, since the latter option is no longer available to us (see under Hammel et al., this column).

Berg, C. C.  2007. Proposals for treating four species complexes in Ficus subgenus Urostigma section Americanae (Moraceae).  Blumea 52: 295–312.

The four complexes in question are designated by the binomials Ficus americana Aubl., F. aurea Nutt., F. citrifolia Mill., and F. pertusa L. f., of which all but F. aurea were used for accepted spp. in the Manual Moraceae treatment by José González (LSCR).  The author of this paper has dealt with these complexes mostly by lumping.  Least affected are Ficus americana and F. citrifolia, the concepts of which are unchanged with respect to Costa Rican material; however, the inclusion of numerous extralimital elements extends their overall geographic ranges somewhat, and consigns our material of F. americana to the autonymic subsp.  Central American material of Ficus americana subsp. americana is further distinguished as belonging to the “‘eugeniifolia’-form,” one of three “informal entities” briefly discussed.  The concept of Ficus pertusa also suffers minimally in Costa Rica, absorbing only F. trachelosyce Dugand, among names accepted in the Manual (though its epithet survives in the name of an “informal entity”).  The most radical changes involve the Ficus aurea complex, the name of which was mentioned in the Manual genus discussion as applying to a sp. of south Florida and the Antilles, and said to have been misapplied in Costa Rica to F. isophlebia Standl., F. jimenezii Standl., F. laterisyce W. C. Burger, and F. tuerckheimii Standl.  The last four names are here lumped summarily into F. aurea (together with Ficus cabusana Standl. & Steyerm., synonymized under F. nymphaeifolia Mill. in the Manual), with only the “‘isophlebia’-form” and “‘tuerckheimii’-form” meriting mention even as “informal entities.”  Features a dichotomous (non-indented) key to the subspp. of Ficus americana, plus distribution maps for the subspp. and “informal entities” of F. americana.

Blanco, M. A. & S. Martén-Rodríguez.  2007. The Stained-Glass Palm, Geonoma epetiolata.  Palms 51: 139–146.

This semi-popular article discusses the morphology, biology, and conservation of Geonoma epetiolata H. E. Moore, “the only neotropical palm that exhibits substantial leaf mottling.”  Rarely encountered in the wild, G. epetiolata occurs only in the understory of very wet, Atlantic-slope forests at 300–1500 m elevation in the Cordillera Central of Costa Rica and in central and eastern Panama.  The average annual rainfall at one Costa Rican station is 6000–7000 mm.  Several alternative hypotheses are considered to explain the mottled leaves of this sp. (maximizing the capture of photosynthetically active radiation, protection from ultraviolet radiation or herbivores, camouflage), but none is fully embraced.  Despite the fact that G. epetiolata is difficult to grow, wild populations have been persecuted by “unscrupulous collectors,” and “it is said that the type population [in Panama] has been extirpated.”  Even so, the sp. may be “locally abundant,” at least in Costa Rica, where populations are nominally protected within Parque Nacional Braulio Carrillo and some private reserves.  Weighing these factors, the authors recommend that Geonoma epetiolata “should probably be listed as Vulnerable according to the IUCN Red List criteria.”  Illustrated with vivid color photographs of living plants in the wild.

Brummitt, R. K.  2007. Report of the Nomenclature Committee for Vascular Plants:  59.  Taxon 56: 1289–1296.

The following recommendations are of only passing interest (now that we know they are not binding in any way):  the conservation of Gnaphalium purpureum L. (Asteraceae) with a conserved type is recommended [safeguarding the well-known Gamochaeta americana (Mill.) Wedd.]; the conservation of Prescottia Lindl. (Orchidaceae) with that spelling is recommended (vs. the original but incorrect and rarely used Prescotia); the rejection of Cymbidium muricatum Sw. (Orchidaceae) is recommended (protecting the well-established Dichaea morrisii Fawc. & Rendle); the conservation of the very recent Xanthocyparis Farjon & Hiep against the much older (but neglected) Callitropsis Oerst. (Cupressaceae) is surprisingly recommended [meaning that taxonomists wishing to divide traditional Cupressus into New and Old World genera would now have to use the name Xanthocyparis for the New World taxa, even though all the indicated new combinations were recently validated in Callitropsis; see The Cutting Edge 14(1): 7, Jan. 2007]; the conservation of Duroia L. f. (Rubiaceae) against Coupoui Aubl. is recommended unanimously, as are the conservation of Caladium Vent. (Araceae) with a conserved type (precluding its substitution for Colocasia Schott) and the conservation of Canna tuerckheimii Kraenzl. (Cannaceae) against C. sylvestris Roscoe (recently confirmed as a synonym).

Clayton, J. W., E. S. Fernando, P. S. Soltis & D. E. Soltis.  2007. Molecular phylogeny of the tree-of-heaven family (Simaroubaceae) based on chloroplast and nuclear markers.  Int. J. Pl. Sci. 168: 1325–1339.

The big news here is that Hans Peter Nooteboom’s (1962) notoriously “broad circumscription of Quassia is unwarranted given several well-supported clades corresponding to traditional generic units.”  Said traditional units include Simaba and Simarouba, both of which were synonymized by Nooteboom (along with several other traditional genera) under Quassia, which thereby became a genus of some 40 spp.  That taxonomy is tactfully characterized as “unnecessarily conservative” by the present authors, who suggest that “Quassia should be limited to just two species”:  Q. africana (Baill.) Baill. and Q. amara L.  Truth be told, every major Mesoamerican floristic work known to us has ignored Nooteboom and continued to treat Simaba and Simarouba as distinct from Quassia; however, some references, most notably The plant-book (2. ed., 1997), have obediently accepted Quassia s. l.

Dressler, R. L.  2007. Sobralia macrophylla:  y otras sobralias de brácteas tubulares, con dos especies nuevas, S. rarae-avis y S. sororcula/Sobralia macrophylla and other sobralias with tubular bracts, with two new species, S. rarae-avis and S. sororcula.  Orquideología 25: 31–49.

Of the two new spp. mentioned in the title(s), only Sobralia rarae-avis Dressler (Orchidaceae) occurs in Costa Rica.  Not an outright addition to the flora, S. rarae-avis corresponds to the sp. that was called S. madisonii Dodson in the Manual (the voucher for which is here cited as a paratype).  The latter name is now restricted to an Ecuadorian sp. that is very similar vegetatively, but differs in various floral details.  The lack of decent flowering material from both Costa Rica and Ecuador had precluded a meaningful comparison in time for the Manual treatment, but these deficiencies have since been overcome.  Sobralia rarae-avis is known from three Costa Rican collections, from roughly 500–900 m elevation (by our estimate) on the Atlantic slope of the Cordilleras de Tilarán (Monteverde region) and Central (including the private Rara Avis reserve, whence the epithet).  The author claims that the sp. is also known from Panama, but cites no supporting voucher.  Illustrated with color photos of living material and a composite line drawing of floral details.  Fully bilingual (Spanish/English).

Hammel, B. E., M. H. Grayum, C. Herrera & N. Zamora (eds.).  2007. Manual de plantas de Costa Rica.  Vol. VI.  Dicotiledóneas (HaloragaceaePhytolaccaceae).  Monogr. Syst. Bot. Missouri Bot. Gard. 111: 1–933.

This series continues with the first dicot volume to be published, and the fourth altogether of a projected total of seven.  And where better to attack the dicots than directly in the mid-section?  The 54 families treated in this volume account for a total of 1396 Costa Rican spp. in 296 genera.  Thus, in both sp. and page number, this is (contrary to expectations) our largest volume yet.  The flagship family for Vol. 6 is Melastomataceae, featuring 303 spp. of mostly understory trees, followed rather distantly by Lauraceae (146 spp.) and Malvaceae (98 spp.)  Astute readers may recall that Piperaceae were supposed to be included in this volume [see The Cutting Edge 10(2): 1, Apr. 2003], but they would not have fit here (and we hope they can be accommodated in Vol. 7!).  These same readers may also note the absence of particular families within the alphabetical purview of Vol. 6, e.g., Hippocastanaceae and Hippocrateaceae; never fear, these omitted groups will appear in future volumes, subsumed within other families (Sapindaceae and Celastraceae, respectively, in the case of these exemplars), in accordance with recent phylogenetic studies.  Illustrated with 343 line drawings, 54 black-and-white photos, and eight pages of color photos.  The basic features of the new volume are the same as for Vols. 2 and 3 [see The Cutting Edge 9(1): 6, Jan. 2004]; however, in a break with tradition, three new spp. names [in Eschweilera (Lecythidaceae)] are validated (in an appendix) and three existing names [in Ocotea (Lauraceae)] are lectotypified.

Many contributors share the honors for Vol. 6, including:  Frank Almeda (CAS; Melastomataceae); William R. Anderson (MICH; Malpighiaceae); Fred R. Barrie (F/MO; Myrtaceae); Garrett E. Crow (NHA; Haloragaceae, Lentibulariaceae, Menyanthaceae, Nymphaeaceae); Armando Estrada (CR; Lythraceae, Passifloraceae pro parte); Paul Fryxell (RSA, Malvaceae); Jorge Gómez-Laurito (USJ; Monimiaceae); José González (LSCR; Hamamelidaceae, Hernandiaceae, Hydrophyllaceae, Krameriaceae, Lacistemataceae, Lauraceae pro parte, Magnoliaceae, Moraceae, Nyctaginaceae, Oleaceae, Onagraceae); Bruce K. Holst (SEL; Myrtaceae pro parte); Quírico Jiménez (formerly INB; Meliaceae, Menispermaceae, Myristicaceae, Olacaceae); M. L. Kawasaki (F; Myrtaceae pro parte); David Kenfack (MO; Carapa); Ricardo Kriebel (CAS; Melastomataceae pro parte); L. R. Landrum (ASU; Myrtaceae pro parte); J. Francisco Morales (INB, Hydnoraceae, Hydrangeaceae, Juglandaceae, Lepidobotryaceae, Loasaceae, Loranthaceae, Myricaceae, Myrsinaceae, Ochnaceae, Orobanchaceae, Oxalidaceae, Phytolaccaceae); Scott A. Mori (NY; Lecythidaceae); Amy Pool (MO; Lamiaceae); Luis J. Poveda (JVR; Lauraceae pro parte); Alexánder Rodríguez (INB; Guarea, Muntingiaceae, Passifloraceae pro parte); Joaquín Sánchez (CR; Loganiaceae); Armando Soto (INB; Papaveraceae); and Gina Umaña (formerly CR; Melastomataceae pro parte).  Our profound gratitude to one and all!

Iltis, H. H. & T. S. Cochrane.  2007. Studies in the Cleomaceae V:  a new genus and ten new combinations for the Flora of North America.  Novon 17: 441–451.

It seems that Cleome is being subjected to the same sort of treatment as Capparis (see below), viz., splintering.  Based on a “comprehensive synopsis of New World Cleomaceae,” as yet unpublished, these authors propose to convert traditional Cleome into an unstated number of smaller genera.  As in the case of Capparis, Cleome s. str. will be limited to Old World spp., at least one of which (C. viscosa L.) occurs adventively in Costa Rica.  We know that the remainder of Cleome s. l. has decayed into at least four genera, as that is the number addressed in this paper.  One of these, Cleoserrata Iltis, is described as new; it comprises five spp., including the redundantly named Cleoserrata serrata (Jacq.) Iltis (formerly Cleome serrata Jacq.), well known in Costa Rica.  Just one of the other three segregate genera is definitely represented in Costa Rica, that being Tarenaya Raf., by the cultivated (and escaped) T. hassleriana (Chodat) Iltis (based on Cleome hassleriana Chodat).  Tarenaya is said to harbor “the majority of New World species classified in the genus Cleome,” though we learn little else of its composition.

-- & X. Cornejo.  2007. Studies in the Capparaceae XXX:  Capparicordis, a new genus from the neotropics.  Brittonia 59: 245–254.

The new genus is of no concern to us, but a cryptic incidental remark by the authors portends turmoil:  “...we herewith establish the genus Capparicordis Iltis & Cornejo, and reduce Capparis s.s. to a small Old World genus centering on the generic type, C. spinosa L.”  We can detect no further mention of the alleged reduction in this paper, and no action taken that would accomplish same.  However, it would appear that, at some point in the foreseeable future, all the 16 or so spp. in Costa Rica presently referred to Capparis will find themselves in a different genus (or genera).  For a tad more insight, see the next entry.

-- & ---. 2007. Studies in the Capparaceae XXXII:  a new combination in Quadrella.  Novon 17: 452–453.

Okay, here is where the events presaged in the foregoing article begin to materialize (though we seem to have missed XXXI!).  Quadrella (DC.) J. Presl comprises an unspecified number of spp., ranging from Texas and Florida through the Mesoamerican region to Colombia, Venezuela, and the West Indies, segregated from Capparis s. l.  The only combination validated here is Q. incana (Kunth) Iltis & Cornejo, for a sp. said to range from Texas to northern Honduras; but unless the sp. has also been split up, Capparis incana Kunth ranges southward to Costa Rica.

Kårehed, J. & B. Bremer.  2007. The systematics of Knoxieae (Rubiaceae)—molecular data and their taxonomic consequences.  Taxon 56: 1051–1076.

The largely African genus Pentas is shown to be non-monophyletic and is split into four smaller genera.  Although the new taxonomy is formalized, with numerous new combinations validated, Pentas lanceolata (Forssk.) Deflers, cultivated in Costa Rica (and pantropically), is effectively the type sp. of Pentas and thus unaffected.

Knapp, S.  2007. Lectotypification of Cavanilles’ names in Solanum (Solanaceae).  Anales Jard. Bot. Madrid 64: 195–203.

At least four of these designations pertain to names accepted for spp. native to or cultivated in Costa Rica, viz., Solanum betaceum Cav., S. elaeagnifolium Cav., S. lanceolatum Cav., and S. lentum Cav. [the basionym of Lycianthes lenta (Cav.) Bitter].  None of the lectotypifications affects current usage.  Illustrated with color(!) photos of the lectotypes.

Kocyan, A., L.-B. Zhang, H. Schaefer & S. S. Renner.  2007. A multi-locus chloroplast phylogeny for the Cucurbitaceae and its implications for character evolution and classification.  Molec. Phylogen. Evol. 33: 553–577.

“The first molecular phylogeny for the family” Cucurbitaceae is inferred from analysis of chloroplast DNA sequences, involving all but seven of the 130 genera currently accepted.  Most of the insights gained from this study pertain to suprageneric classification, of little interest to us.  Descending to our level, several genera appear to be paraphyletic, including:  Cucumis (with respect to several oligotypic Old World genera), Psiguria (with respect to Gurania and the monospecific, South American Helmontia), and perhaps Sicyos (with respect to Microsechium).  The situation with Cucumis (cultivated and adventive in Costa Rica) has already been resolved by subsuming the smaller genera therein (see Ghebretinsae et al., 2007; Novon 17: 176–178).  Apparently, the Psiguria and Sicyos problems will require additional research.

Krapovickas, A. & W. C. Gregory.  2007. Taxonomy of the genus Arachis (Leguminosae).  Bonplandia (Corrientes) 16, Supl.: 1–205.

If the title sounds familiar, there’s a simple explanation:  this is nothing more than a translation into English (by D. E. Williams & C. E. Simpson) of the 1994 Spanish-language revision of peanuts and company by Krapovickas & Gregory [see The Cutting Edge 1(4): 6–7, Oct. 1994].

Kreier, H.-P., A. F. Rojas Alvarado, A. R. Smith & H. Schneider.  2007. Hyalotrichopteris is indeed a Campyloneurum (Polypodiaceae).  Amer. Fern J. 97: 127–135.

And Campyloneurum is indeed a Polypodium!  Editorializing (partially) aside, the first stage of our fantasy transformation is now complete, as the once-and-future Polypodium anetioides Christ, way out on a limb in the monospecific Hyalotrichopteris W. H. Wagner, is returned to the Campyloneurum fold, wherein it is comfortably nested, according to these cladistic analyses of cpDNA sequence data.  Lack of suitable material from this rare sp. (restricted to Costa Rica and Panama) had prevented its inclusion in previous such studies.

Kuijt, J.  2007. First record of the genus Oryctina (Loranthaceae) in Mesoamerica:  O. costaricensis, a new species from Costa Rica.  Novon 17: 476–478.

So now we have a name for “Oryctina sp. A” of the Manual, viz., O. costaricensis Kuijt, endemic to the Península de Osa.  In the bargain, we get an expanded description and discussion and two excellent, composite line-drawings (this genus having been added to the Manual too late for a drawing to be made).  The new sp. is compared with the Venezuelan Oryctina badilloi (Ferrari) Kuijt and O. pedunculata (Kuijt) Kuijt, from which it differs in several respects; all three of these sp. would belong to the genus Maracanthus Kuijt, should it be resegregated (as the author has suggested), in which case O. costaricensis would require a new combination.  More intriguing than the new sp., at least for us, is the author’s allegation of the occurrence on the Península de Osa of Psittacanthus acinarius (Mart.) Mart. (Loranthaceae), a sp. not mentioned in the Manual and for which we can find no Costa Rican records in any of our usual sources.  Could it be the same as “Psittacanthus sp. A” of the Manual?

Liu, H.-M., X.-C. Zhang, W. Wang, Y.-L. Qiu & Z.-D. Chen.  2007. Molecular phylogeny of the fern family Dryopteridaceae inferred from chloroplast rbcL and atpB genes.  Int. J. Pl. Sci. 168: 1311–1323.

This study supports the recent new fern classification of Alan R. Smith (UC) and colleagues [see The Cutting Edge 13(4): 9–10, Oct. 2006] with respect to Dryopteridaceae, except that Didymochlaena is here excluded from that family.  Arachniodes would appear to be paraphyletic, and Dryopteris and Polystichum polyphyletic; however, as few if any New World members of these genera were included in the study, the potential consequences for Costa Rican floristics cannot be assessed (not by us, anyway).

McDade, L. A. & E. A. Tripp.  2007. A synopsis of Costa Rican Ruellia (Acanthaceae), with descriptions of four new species.  Brittonia 59: 199–216.

This precursor to the Manual Acanthaceae treatment (which will involve at least the first author) accounts for 22 spp. of Ruellia in Costa Rica, by means of a dichotomous, indented key and a “quick guide” to sp.-groups according to eight “distinctive traits” (these features largely comprising the synoptic portion of the paper).  Three of the four new spp. are endemic to the Península de Osa:  Ruellia exilis McDade & E. A. Tripp, compared with various spp., including R. pittieri Lindau, R. stemonacanthoides (Oerst.) Hemsl., and R. malacosperma Lindau; Ruellia mira McDade & E. A. Tripp, “very distinctive,” but somewhat resembling R. praeclara Standl. and R. tubiflora Kunth; and Ruellia odorata E. A. Tripp & McDade, most similar to R. palustris Durkee.  The fourth new sp., Ruellia norvegigratiosa McDade & E. A. Tripp (the odd epithet acknowledging financial support of INBio by the Norwegian government), is known by a single collection from 600–700 m elevation on the Pacific slope of the Cordillera de Talamanca in the Dota region; it is distinguished from the same group of spp. mentioned above under R. exilis.  As promised in a recent paper by the second author [see The Cutting Edge 14(4): 13, Oct. 2007], the Costa Rican spp. of Blechum are transferred to Ruellia on the basis of unpublished molecular data that shows them nested therein.  The combination Ruellia costaricensis (Oerst.) E. A. Tripp & McDade is validated for the former Blechum costaricense Oerst., while Ruellia blechum L. is already available for Blechum pyramidatum (Lam.) Urb.  Three new synonymies are proposed, at least tentatively:  Ruellia cooperi Leonard under R. biolleyi Lindau, R. tonduzii Lindau under R. pittieri Lindau, and R. campestris (Oerst.) Hemsl. (a name not in recent use) under R. geminiflora Kunth.  It is also suggested that the widely used Ruellia metallica Leonard may prove synonymous with R. barbillana Cufod., a name that has been applied only to its type collection.  Finally, minor, in-country range extensions are reported for both Ruellia stemonacanthoides (though we don’t detect anything different here) and R. golfodulcensis Durkee (to the Atlantic slope).  All the new spp. are illustrated with composite line drawings and SEM micrographs of pollen and laminar glands.

Meerow, A. W., J. L. Reveal, D. A. Snijman & J. H. Dutilh.  2007. (1793) Proposal to conserve the name Amaryllidaceae against Alliaceae, a “superconservation” proposal.  Taxon 56: 1299–1300.

Cladistic studies have shown that Alliaceae, Amaryllidaceae, and Agapanthaceae form a monophyletic group that may be treated optionally as a single family, according to the most recent classification of the Angiosperm Phylogeny Group [see The Cutting Edge 10(3): 5–6, Jul. 2003].  These authors take no position as regards the one- vs. three-family options; rather, they propose to require that the familiar name Amaryllidaceae be used instead of Alliaceae (the oldest of the three) in the event of a merger.

Morales, J. F. & J. González.  2007. Buddleja filibracteolata (Buddlejaceae), una nueva especie para Costa Rica.  Anales Jard. Bot. Madrid 64: 161–163.

Buddleja filibracteolata J. A. González & J. F. Morales is described on the basis of a single specimen, from 1000–1200 m elevation on the the flanks of Cerro Caraigres, on the Pacific slope of the northern Cordillera de Talamanca.  The new sp. is compared with B. crotonoides A. Gray, with which it shares valvate sepals; it differs from the latter in its much shorter, denser, spiciform inflorescences, conspicuous, filiform bracts and bisexual (vs. unisexual) flowers with a much longer calyx.  We have good reason to believe that the real B. crotonoides also occurs in Costa Rica (specifically, on the Pacific slope of Volcán Barva and in the Cerros de Escazú), even though these authors portray it as “distribuido desde México a Nicaragua.”  Furthermore, we know of at least one other collection of the new sp. from the type locality.  Illustrated with a photo of the type specimen, supplemented with line drawings of floral details.

Mori, S. A.  2007. Tropical botanist and blueberry expert retires.  Gard. News New York Bot. Gard. 41(5): 7.

May we take this opportunity to congratulate longtime NY curator and Manual Ericaceae co-contributor James L. Luteyn on his (early!) retirement to upstate Michigan, and to thank him for polishing off his Manual manuscript before he left.  May he prosper in the tranquility of Beaver Island!  And don’t miss this appreciation of the man by NY colleague (and Manual Lecythidaceae contributor) Scott A. Mori.

Mytnik-Ejsmont, J. & P. Rutkowski.  2006. Taxonomic position of Zhukowskia Szlach., R. González T. & Rutk. (Cyclopogoninae, Spirantheae, Orchidaceae).  Biodivers. Res. Conservation 1-2: 7–10.

In response to molecular studies showing the recent segregate Zhukowskia [see The Cutting Edge 8(2): 14, Apr. 2001] to be nested within Sarcoglottis, these authors (with a vested interest in the matter) proffer statistical phenetic analyses that “confirm the legitimacy” of the former genus.  A dichotomous key separates the five spp. of Zhukowskia, including Z. smithii (Rchb. f.) Szlach., R. González & Rutk., treated as Sarcoglottis smithii (Rchb. f.) Schltr. in the Manual.

Nickrent, D. L.  2007. Cytinaceae are sister to Muntingiaceae (Malvales).  Taxon 56: 1129–1135.

This according to molecular analyses, and with “strong support.”  Cytinaceae comprises two genera of the traditional Rafflesiaceae, now dissembled among four separate families of disparate affinities (with Rafflesiaceae s. str. now limited to three tropical Asian genera).  Cytinaceae consists of the Central American Bdallophyton (which occurs rarely in Prov. Guanacaste, Costa Rica) and the largely African Cytinus.  Morphological synapomorphies with Muntingiaceae “include trichome types, mature pollen released as tetrads, inferior ovaries, intrusive parietal placentation with many ovules, and fruit type.”  We will clearly have to rethink our Manual treatment of Rafflesiaceae s. l., also represented in Costa Rica by the genus Apodanthes, lately referred to Apodanthaceae.  But the relationships of Apodanthaceae have still not been resolved, and no members of that family were included in the present study; indeed, according to previous studies Apodanthaceae have “varied between Malvales and Cucurbitales,” suggesting that an intimate affinity with Cytinaceae is not out of the question.

Oakeley, H. F.  2007. A new infrageneric classification of Lycaste (Lindl.) and a checklist of species for the genus Lycaste.  Orchid Digest 71: 196–208.

This gaudily and gratuitously illustrated article seems intended mainly as a promo for an upcoming book by the author (advertised on the final page), in which all the new taxa included in this checklist are to be validated (Code requirements such as Latin descriptions and/or type citations being omitted from this paper).  About 31 spp. of Lycaste (Orchidaceae) are here accepted (approximating previous estimates), as well as a fair number of vars. and hybrids.  Most of the seven Lycaste spp. treated in the Manual continue to be recognized under the names used therein, but a few changes must be noted.  The sp. called Lycaste brevispatha (Klotzsch) Lindl. & Paxton in the Manual is apparently undescribed, with L. brevispatha itself a synonym of L. candida Rchb. f.  Another name used in the Manual, Lycaste macrophylla (Poepp. & Endl.) Lindl., is here restricted to a South American sp.; Costa Rican material that had gone by that name is now apportioned among at least four different spp., with names elevated from subsp. rank under L. macrophylla.  Whether intentionally or not, these new combinations appear to be validly published in this paper, so we will mention them here:  Lycaste measuresiana (B. S. Williams) Oakeley, L. panamanensis (Fowlie) Oakeley, L. puntarenasensis (Fowlie) Oakeley, and L. xanthocheila (Fowlie) Oakeley.  Lycaste measuresiana and L. panamanensis are attributed to Costa Rica for the first time, as far as we know, while L. macrophylla subsp. desboisiana (Cogn.) Fowlie, accepted in the Manual, is nowhere accounted for.  Also attributed to Costa Rica (in addition to various vars. and hybrids, which do not particularly interest us) are Lycaste luminosa Oakeley and L. xytriophora Linden & Rchb. f., both mentioned and rejected in the Manual genus discussion, as well as another still-undescribed sp.

O’Leary, N., M. E. Múlgura & O. Morrone.  2007. Revisión taxonómica de las especies del género Verbena (Verbenaceae):  serie Pachystachyae.  Ann. Missouri Bot. Gard. 94: 571–621.

The genus Verbena, though lately of global distribution, is New World in origin, and with a total of about 45 spp. (according to these authors).  About two-thirds of these spp. belong to the basically North American ser. Verbena, the remainder to the basically South American ser. Pachystachyae Schauer (characterized, as according to its name, by generally stouter, more densely flowered inflorescences).  This revision treats the 18 spp. of ser. Pachystachyae, 13 of which belong to the autonymic subser., the rest to the newly described subser. Pseudoracemosae N. O’Leary.  The former subser. includes Verbena litoralis Kunth, the only sp. of the genus that grows wild (though adventively) in Costa Rica.  Two vars. of V. litoralis are accepted, of which our material would figure to represent the autonymic one (though no Costa Rican specimens are cited).  The distribution summaries of three other taxa (Verbena bonariensis L. var. bonariensis, V. glabrata Kunth, and V. rigida Spreng.) include Costa Rica implicitly, but no Costa Rican specimens are cited for any of these, and we find no such records in any of our sources.  Features synonymy and typology at all levels, formal descriptions of the genus and all its spp. and vars., dichotomous, indented keys to the sers. and subsers. of Verbena and the spp. of ser. Pachystachyae, and, for each sp. or var., a distribution summary, discussion, and list of exsiccatae.  Most of the taxa are illustrated with composite line drawings.  At the end are found sections on dubious and “invalid” taxa, as well as indices to exsiccatae and synonyms.  The brief introductory part addresses taxonomic history, subdivision of the genus, various aspects of morphology and anatomy, and phenology.  In Spanish (with English abstract).

Ossenbach, C.  2007. “Señor” Endrés:  profiling a mysterious collector of orchids.  Orchids (West Palm Beach) 76: 594–601.

This represents a careful compilation of just about every detail that is known, or even so much as surmised, about the obscure A. R. Endres (or perhaps it is correctly “Endrés,” as the gentleman’s signature, reproduced here, seems to indicate).  While the author has been characteristically thorough, his opening sentence (“Almost nothing is known about the origins and life of A. R. Endrés”) proclaims that this will be no mother lode of new information.  Nevertheless, we do learn a few intriguing details about the man’s life:  he worked for a while on Koschny’s finca in the San Carlos region, establishing a plantation of “native rubber” (we would assume this means Castilla elastica Sessé ex Cerv.) and nutmeg (does the adjective “native” apply here as well, viz., Virola koschnyi Warb.?); he may also have collected in Panama, Peru, and Brazil; and he collected birds as well as plants.  Includes a list of orchid spp. collected by Endres, plus a list of “plants other than orchids” collected by or dedicated to him.  Richly endowed with gratuitous orchid illustrations, as typical of these hobbyist journals, but also featuring a photo of Avenida Central, San José, ca. 1871 (around the time of Endres’s residence).

Peterson, P. M. & I. Sánchez Vega.  2007. Eragrostis (Poaceae: Chloridoideae: Eragrostideae: Eragrostidinae) of Peru.  Ann. Missouri Bot. Gard. 94: 745–790.

From this unlikely source, we learn two things that we probably should already have known.  First, Eragrostis tenella (L.) P. Beauv. ex Roem. & Schult. reverts to being the correct name for the sp. called E. amabilis (L.) Wight & Arn. ex Nees in the Manual (and, before that, in Flora mesoamericana Vol. 6 and Standley’s Flora of Costa Rica); “reverts,” because E. tenella was the name used for this sp. in Richard W. Pohl’s Flora costaricensis treatment (1980).  We have no idea what is going on here.  Second, the name Eragrostis glomerata (Walter) L. H. Dewey, accepted in the Manual for a sp. indicated as New World native, is now regarded as a synonym of E. japonica (Thunb.) Trin., an Asian sp. introduced in our region.

Pool, A.  2007. A revision of the genus Pithecoctenium (Bignoniaceae).  Ann. Missouri Bot. Gard. 94: 622–642.

Pithecoctenium is a mainly South American genus of lianas, comprising just four spp.  In addition to its habit, it is characterized by sharply hexangular stems, bifoliolate leaves with a terminal tendril, white or pale yellow, bent corollas, and (generally) echinate fruits.  Just one sp. extends beyond South America, P. crucigerum (L.) A. H. Gentry, which ranges from Mexico to northern Argentina and Uruguay, as well as in the Greater Antilles.  This has been much collected in Costa Rica, so there is little new here for us.  Features full synonymy (with many new synonyms) and typology (with numerous newly designated lectotypes), detailed genus and sp. descriptions, and an index to exsiccatae.  Each sp. entry includes distribution and phenology summaries, enumerations of common names and uses (where known), a discussion, and representative specimen citations.  The introduction considers morphology, taxonomic history, anatomy, palynology, distribution (with maps), and uses.  Apart from a reproduction of a plate designated as a lectotype, there are no illlustrations.

Pupulin, F. & J. D. Zúñiga.  2007. The upside-down Pleurothallis (Orchidaceae) of Mesoamerica, with a new species from Costa Rica.  Orchids (West Palm Beach) 76: 690–695.

Pleurothallis bogarinii Pupulin & J. D. Zúñiga sp. nov. is distinguished (though only in the Latin diagnosis) from the otherwise very similar P. scaphipetala Luer on the principal basis of floral details.  The new spp. is based on just two collections, from roughly 100–500 m elevation (by our estimate) on the Caribbean slope of the Cordillera Central.  Illustrated by an excellent composite line drawing, as well as a color photo from life of the actual holotype plant.  Together with Sobralia rarae-avis (see under Dressler, this column), this pushes our running total of orchid spp. described from Costa Rica since the inception of this newsletter up to 251.

Retana-Salazar, A. P.  2007[‘2006’]. Las 100 especies de Alexander Francisco Rojas-Alvarado.  MES 2(2): 1–4.

This seems as good a place as any to introduce the new periodical MES (an acronym for “Métodos en Ecología y Sistemática”), edited by the author of this paper.  MES, the organ of the San José-based NGO Instituto Cenroamericano de Investigación en Biología y Conservación (ICIBC), is published on-line three times per year.  By way of satisfying nomenclatural-code requirements for hard-copy publication, authors of taxonomic papers are required to purchase at least 25 reprints, to be distributed to selected libraries, all either general or zoological in nature, and just two of which are outside Costa Rica (Germany and Mexico).  This reprint ploy is fast becoming the proverbial foot-in-the-door for on-line taxonomic journals (see also under Soto Arenas et al., this column).  The vast majority of interested individuals will thus have to seek articles in MES, as well as general information about the journal, at the following site:


This particular paper celebrates the publication, in the same issue (see following three entries), of the hundredth new sp. (all ferns) described by Costa Rican pteridologist Alexander Rojas (USJ), beginning only in 1996.  Actually, the number has reached 101, including a few hybrids, and all of said novelties are conveniently enumerated here, in chronological order, with formal bibliographic citations (which is where we learn that “MES” is preferred for this journal, rather than “Métod. Ecol. Sist.”).  We congratulate Alex on this most impressive achievement!

Rojas-Alvarado, A. F.  2007[‘2006’]. Novelties in Tectaria (Tectariaceae) in the Neotropics.  MES 2(1): 6–9.

The only new taxon is Tectaria rivalis (Mett. ex Kuhn) C. Chr. var. gemmipara A. Rojas, occurring at ca. 400–1000 m elevation on the Pacific slope of the Cordillera de Talamanca and, disjunctly, in Depto. Chocó, Colombia.  It differs form the autonymic var. in having gemmae (asexual propagules) in the axils of the distal pinnae and, often, the costae of the proximal pinnae.  The remaining “novelties” comprise range extensions, of which just two are of interest to us:  Tectaria faberiana A. Rojas is reported from Depto. Chocó, Colombia, marking the loss of another Costa Rican endemic; and Tectaria murilloana A. Rojas, otherwise known from Panama to Ecuador, can now be added to the Costa Rican flora on the basis of G. Herrera et al. 2905, from 600 m elevation on the Atlantic slope of the Cordillera de Guanacaste.  The new var. is depicted in detailed photos showing the diagnostic gemmae.

--.  2007[‘2006’]. Two new species of Adiantum (Filicales: Pteridaceae) from Costa Rica and Nicaragua.  MES 2(2): 5–12.

Both new sp. are restricted in Costa Rica to elevations below 300 m on the northern Atlantic slope, mainly on the Llanura de Tortuguero.  The endemic Adiantum caribense A. Rojas is compared mainly with A. obliquum Willd. from which it differs in its slenderer rhizome, smaller leaf-blades, sparsely scaly (vs. ± glabrous) abaxially, and glabrous (vs. puberulent) indusia.  Adiantum reptans A. Rojas, which extends into Nicaragua, differs from A. latifolium Lam. in substantially the same manner, at least as regards relative leaf-blade size and abaxial laminar indumentum.  It has also been confused with A. terminatum Kunze ex Miq., under which name it was recently reported as new to the Estación Biológica La Selva [see The Cutting Edge 14(3): 3, Jul. 2007].  Both new spp. are illustrated with composite line drawings of their types.

--.  2007[‘2006’]. Una nueva especie de Notholaena (Pteridaceae) para Costa Rica.  MES 2(2): 18–21.

Notholaena revoluta A. Rojas, restricted to a few hills (limestone, we think) in Parque Nacional Palo Verde (one of the driest parts of Costa Rica), has been confused with N. candida (M. Martens & Galeotti) Hook., from which it differs inter alia in its shorter, nearly concolorous (vs. strongly bicolored), and more narrowly margined rhizome scales, slenderer stipes, and occurrence at lower elevations (50–250 m, vs. 450–1700).  We don’t know whether this means that N. candida must be deleted from the Costa Rican flora.  The new sp. has also been mistaken for Aleuritopteris farinosa (Forssk.) Fée, another sp. of higher elevations.  Illustrated with a composite drawing of the holotype.  Incidentally, two paratypes of N. revoluta are erroneously cited:  U. Chavarría et al. 1665 and 1666 are correctly 1965 and 1966, respectively.

Rzedowski, J., R. Medina Lemos & G. Calderón de Rzedowski.  2007. Segunda restauración de Bursera ovalifolia y nombre nuevo para otro componente del complejo de B. simaruba (Burseraceae).  Acta Bot. Mex. 81: 45–70.

The “nombre nuevo” does not concern us, but the “segunda restauración,” rather surprisingly, does.  The name Bursera ovalifolia (Schltdl.) Engl. is resurrected from recent synonymy under B. simaruba (L.) Sarg., and applied to a sp. resegregated from the latter complex and freshly characterized.  The citation of two Costa Rican specimens (both duplicated at MO), supplemented by excellent illustrations, suffices to nail this as a mystery sp. that we have collected numerous times in the Guanacaste region and apparently misidentified, generally as Bursera permollis Standl. & Steyerm. (the latter restricted, according to this paper, to Guatemala and Honduras).  Bursera ovalifolia, ranging from southern Mexico to Costa Rica, differs from the more widespread B. simaruba (according to this paper and our own experience) in having generally fewer, proportionately broader leaflets and smaller fruits (for those fortunate enough to have pistillate material).  It is apparently rather more similar to B. standleyana L. O. Williams & Cuatrec., an epiphytic(!) sp. endemic to very wet lowland forests on the Pacific slope of Costa Rica, which these authors portray as only dubiously distinct.  Whatever the case, we are in complete agreement that Costa Rican material here called Bursera ovalifolia should be maintained as distinct from B. simaruba—an argument made pointedly on herbarium labels by such field cognoscenti as Bill Haber and Dan Janzen.

Features a dichotomous, indented key to the Mesoamerican spp. of the Bursera simaruba complex (which, surprisingly to us, does not include B. schlechtendalii Engl.) and, for B. ovalifolia, synonymy and typology, a rigorous description, specimen citations, an extensive discussion, and both a composite line drawing and a black-and-white photo of living material.

Schuettpetz, E. & K. M. Pryer.  2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes.  Taxon 56: 1037–1050.

Here we present some selected gleanings:  “two distinct Alsophila clades” emerge within the “scaly tree fern clade”; “the genus Dennstaedtia is strongly supported...as paraphyletic”; “Athyrium is not monophyletic”; “nearly all genera previously segregated from Asplenium (e.g., ...Loxoscaphe) nest well within this large genus”; “Thelypteris...is definitely paraphyletic to the cyclosoroids”; “Blechnum is definitely not monophyletic”; Lastreopsis is a “paraphyletic genus”; “Bolbitis is resolved as polyphyletic”; and “the grammitid ferns...are nested firmly within the Polypodiaceae.”  Space does not permit analyses of the potential repercussions of these conclusions for the Costa Rican flora, and in any case, most of the implied taxonomic changes remain to be formally implemented.

Sitko, M., P. Tukałło & M. Górniak.  2006.  Introduction to the phylogenetic analysis of Maxillaria Ruiz & Pav. (Maxillariinae, Orchidaceae).  Biodivers. Res. Conservation 3-4: 200–204.

Cladistic analysis of ITS sequence data suggests that several genera presently or previously accepted as distinct are nested within a paraphyletic Maxillaria s. l., viz.:  Camaridium, Cryptocentrum, Heterotaxis, Ornithidium, Sepalosaccus, and Trigonidium.  Moreover, Ornithidium is itself polyphyletic.  For a more definitive examination of this problem, see under Whitten et al. (this column).

Soto Arenas, M. A., G. Salazar & C. van den Berg.  2007. New combinations in Domingoa, Homalopetalum (Orchidaceae: Laeliinae), and Nemaconia (Orchidaceae: Ponerinae).  Neodiversity 2: 7–9.

The only item here of interest to us concerns the genus Ponera, shown to be non-monophyletic in a recent (unpublished) study by this group.  The situation is remedied by splitting off several spp. under the generic name Nemaconia Knowles & West., entailing the new combination Nemaconia striata (Lindl.) Van den Berg, Salazar & Soto Arenas for the sole sp. treated under Ponera in the Manual.  Neodiversity, by the way, is an on-line, Brazilian journal (with nominal hard-copy distribution) that may be accessed at the following URL:


Szlachetko, D. L. & M. Górniak.  2006. New taxa in the subtribe Oncidiinae (Orchidaceae).  Biodivers. Res. Conservation 1-2: 11–13.

Based on pollinarium details and (unpublished) molecular studies, 13 spp. previously assigned to Ada and Brassia are here commingled in a new genus, Brassiopsis Szlach. & Górniak.  The only Costa Rican sp. affected by this was called Ada chlorops (Endres & Rchb. f.) N. H. Williams in the Manual, henceforth Brassiopsis chlorops (Endres & Rchb. f.) Szlach. & Górniak, for those so inclined.

Vega, A. S. & Z. E. Rúgolo de Agrasar.  2007. Novedades taxonómicas y sinopsis del género Digitaria (Poaceae, Panicoideae, Paniceae) en América Central.  Darwiniana 45: 92–119.

Judging from the specimen citations, this account appears based almost exclusively on a study of material at K and US.  Thus, inevitably, a few new Costa Rican localities can be gleaned to supplement the Manual distribution summaries, which were based mainly on material at CR, INB, and MO.  We will not take the space to enumerate these, but several other items do deserve mention.  Costa Rican specimens of Digitaria argillacea (Hitch. & Chase) Fernald may now be annotated as belonging to the autonymic var., established recently by another var. published by these same authors.  Digitaria pentzii Stent, the name used in Flora mesoamericana (1994) and the Manual for the sp. that had been called D. decumbens in Richard W. Pohl’s Flora costaricensis (1980) treatment, has now itself fallen into synonymy, under Digitaria eriantha Steud.  Digitaria pittieri (Hack.) Henrard, restricted in Flora mesoamericana and the Manual to Nicaragua and Costa Rica, is here ascribed also to Ecuador, Perú, Venezuela, and Brazil.  And finally, Digitaria sanguinalis (L.) Scop., a name with a long history of having been misapplied to Costa Rican material, rears its head yet again in the form of Pittier 4203 (US), from Puerto Limón, ostensibly the first bona fide report for the country.  Features a dichotomous (though non-indented) key to spp., and excellent composite line drawings of selected spp.

Wagner, W. L., P. C. Hoch & P. H. Raven.  2007. Revised classification of the Onagraceae.  Syst. Bot. Monogr. 83: 1–240.

In which the infrafamilial classification of Onagraceae is overhauled according to the results of recent molecular studies, mostly involving these same authors [see, e.g., The Cutting Edge 11(2): 9, Apr. 2004].  We were a bit alarmed when we first learned that this publication was imminent, just as Manual Vol. 6, with Onagraceae in tow, was going to press (see under Hammel et al., this column).  Nothing like instant obsolescence!  But we needn’t have worried:  the impact upon Costa Rican floristics is minimal.  This is partly because many of the changes involve supra- or infrageneric categories (subfam., sect., subsect., etc.), mainly of interest to family specialists, that we rarely if ever deal with in the Manual.  To be sure, there are some radical new developments at the generic level that will have major consequences for certain regions, e.g., western North America:  Camissonia is shattered into nine genera, while Oenothera absorbs Calylophus, Gaura, and Stenosiphon.  However, none of the genera added or lost occurs in Costa Rica.  In the Manual Onagraceae treatment authored by José González (LSCR), only the following minor adjustments are indicated:  the total number of genera in the family should be changed from “Ca. 18” to 22, and the total number of spp. in Oenothera from “Ca. 120” to 145.  Nonetheless, this remains an invaluable compendium of information on Onagraceae, with synonymy, typology, and formal descriptions at all ranks, a dichotomous, indented key to families, tribes, and genera, comprehensive lists of component taxa for all terminal categories, and many excellent illustrations.  All necessary new combinations and names at sp. and subsp. rank are validated in an appendix, and additional appendices account for new synonymies, conservation status of selected spp., and spp. naturalized outside their native range.  There is also an index to scientific names.  The introductory pages address taxonomic history, the rationale for the new classification (including three cladograms), and biogeography.

Whitten, W. M., M. A. Blanco, N. H. Williams, S. Koehler, G. Carnevali, R. B. Singer, L. Endara & K. M. Neubig.  2007. Molecular phylogenetics of Maxillaria and related genera (Orchidaceae: Cymbidieae) based on combined molecular data sets.  Amer. J. Bot. 94: 1860–1889.

Orchidologists have been chipping away at the huge (ca. 580 spp.) genus Maxillaria for several years now [see, e.g., The Cutting Edge 14(2): 11, Apr. 2007], and with this landmark paper, have finally succeeded in bringing it down.  These authors sought to answer two questions:  “Does Maxillaria consist of one megagenus, or is its present circumscription artifactual...?”; and, “Can Maxillaria be subdivided into more manageable clades defined by strong molecular support and putative morphological synapomorphies?”  Of course, both can be easily answered with no research at all:  the circumscription of a genus (as long as it it monophyletic) is purely subjective, dictated mainly by personal preference; and (assuming “more manageable” to mean “smaller”), any monophyletic genus of more than one sp. can be subdivided, at least theoretically, as indicated.  So it boils down to this:  does one prefer one large genus, or lots of smaller ones?  Specialists tend to opt for the latter, we’ve noticed, while generalists are more likely to embrace more inclusive genera.  These authors being specialists (and a most impressive array, it must be noted), Maxillaria is toast.

The downfall for Maxillaria as we knew it is that, according to this research, seven smaller genera (with a total of about 29 spp.) are nested within it (see also Sitko et al., this column).  These seven genera, together with Maxillaria, comprise what is here termed the “core Maxillariinae.”  Two main resolutions to this problem are considered:  to lump all core Maxillariinae into Maxillaria, “with a subgeneric classification that reflects phylogenetic relationships”; or “to divide Maxillaria s. l. into well-supported clades and create several new genera, together with redefined generic concepts of some existing minor genera.”  The splitting option is advocated on the notion that “a single mega-genus...would be morphologically undiagnosable,” while the recognition of smaller “clades as genera will produce a more predictive classification and will be more useful for field identification and floristic treatments.”  We are skeptical that smaller genera would be more useful in the field; as generalists (vis-à-vis orchids), we’ve rarely had any problem recognizing Maxillaria s. l., but despair of ever learning the smaller, more technical genera proposed here, just as we’ve never been able to learn the segregates of Grammitis s. l. (Grammitidaceae).  And as for “morphologically undiagnosable,” the authors themselves provide said diagnosis on p. 1861 of this paper:  “Core Maxillariinae are characterized by the presence of conduplicate leaves..., single-flowered inflorescences..., four pollinia with or without a stipe attached to an often lunate viscidium, and usually a prominent column foot that is typically persistent in fruit.”  But we digress.

Here, then, is what the near future portends for the “core Maxillariinae” (in the sense of this paper):  eight genera will become 17, based on “well-supported clades that [these authors] feel warrant generic recognition.”  Five of the seven original smaller genera survive, though one (Mormolyca) will be substantially enlarged; lost are Anthosiphon and Chrysocycnis (as synonyms of Cryptocentrum and Mormolyca, respectively).  The remaining 12 genera in the new system will result from the fragmentation of Maxillaria s. l.  Seven of these 12 segregates (including Maxillaria s. str.) entrain valid generic names, three of which are of post-Manual coinage (Brasiliorchis R. B. Singer, S. Koehler & Carnevali, Christensonella Szlach., Mytnik, Górniak & Śmiszek, Sauvetrea Szlach.), while three others have been around for a long time, but either largely ignored (Camaridium Lindl., Ornithidium Salisb. ex R. Br.) or only recently resurrected (Heterotaxis Lindl.).  Sauvetrea in the sense of its author is revealed as polyphyletic, but the name survives by a technicality; not so lucky are Adamanthus Szlach. and Laricorchis Szlach., described in the same recent paper [see The Cutting Edge 14(2): 11, Apr. 2007], which fall into synonymy (under Camaridium and Ornithidium, respectively).  These cases are supposed to illustrate “the danger of erecting genera based upon a few, homoplasious morphological characters and/or incomplete molecular sampling.”

The authors of the present paper can certainly not be criticized for “incomplete molecular sampling.”  Indeed, their sample size is nearly unprecedented, comprising 604 accessions representing ca. 354 spp. of “core Maxillariinae,” that is to say, about 58% of the total.  The authors express confidence that this assortment “includes representatives of all major morphological groups...and that the resulting cladograms adequately reflect relationships among them.”  An astounding 88 (82%) of the 107 spp. of Maxillaria treated in the Manual were included, due no doubt to the involvement of Costa Rican botanist Mario Blanco (FLAS).  With respect to this Costa Rican contingent, the prognosis is grim; despite the authors’ assurance that “the majority of species will remain in Maxillaria s. s..., thus keeping the necessary nomenclatural transfers to a minimum,” the opposite is true for Costa Rica:  the large majority of spp. will fall outside Maxillaria s. str.  Of the 88 Costa Rican spp. of Maxillaria s. l. included in these analyses, just 20 (23%) will remain in that genus under the new system, and just one of the nine excluded spp. for which we are able to hazard an educated guess (out of 19 total).  In other words, at least 76 binomials accepted in the Manual will have to be changed (though it bears mentioning that many of the required “new” binomials already exist).  Contrast this with just 10 such changes that would be required under the lumping option, which would entail (in Costa Rica) merging Cryptocentrum, Mormolyca, and Trigonidium into Maxillaria.  Space does not permit us to indicate the disposition of every Costa Rican sp. in the new system, so the following synopsis must suffice:  of the genera with existing names, by far the largest bloc of spp. (at least 43) goes to Camaridium, followed by Maxillaria s. str. (at least 21 spp.), Ornithidium (at least 5), Mormolyca [at least 3, joining M. ringens (Lindl.) Schltr.], Heterotaxis (at least 3), and Christensonella (with only Maxillaria uncata Lindl. in Costa Rica).  Of the remaining 31 spp. treated under Maxillaria in the Manual, 21 are apportioned in blocs (ranging in size from one to at least 14 spp.) among at least four genera still to be named, while the disposition of 10 is (to us) unfathomable.  Cryptocentrum and Trigonidium, the other genera of “core Maxillariinae” in Costa Rica, are unaffected.  All indicated “generic descriptions and nomenclatural tranfers” are promised for “separate publications.”

We sense that there may have been some disagreement among the authors of this paper as to whether the “highly supported” clade A+B+C should be recognized as a single genus or three smaller ones.  Despite the fact that the three smaller clades received only “moderate...support,” all were accepted as genera (Heterotaxis, Ornithidium, and a still nameless genus with only Maxillaria nasuta Rchb. f. in Costa Rica).  Perhaps these ought to have been lumped (Ornithidium being the oldest name), on the grounds that relationships are more important than differences.


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