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Manual de Plantas de Costa Rica

Main | Family List (MO) | Family List (INBio) | Cutting Edge
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The Cutting Edge

Volume XX, Number 3, July 2013

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

Almeda, F.  2013. A new epiphytic species of Blakea (Melastomataceae: Blakeeae) from the Caribbean rain forest of Costa Rica.  Selbyana 31: 1–4.

Blakea hammettiorum Almeda, sp. nov., is known only from the type specimen, collected by the redoubtable Gerardo Herrera at 300 m elevation on the Atlantic slope of the northern Cordillera de Talamanca (Parque Nacional Barbilla).  Judging from the spp. with which it is compared (B. foliacea Gleason, B. gracilis Hemsl., and B. litoralis L. O. Williams), it seems likely that B. hammettiorum would have been assigned to Blakea even before the merger with Topobea (see under “Penneys,” this column, in our last issue).  The epithet honors Benjamin and Ruth Hammett, friends of the author.  Features an excellent composite line drawing.

Archila, F. M. 2013. Note taxinomique sur le genre Tupacamaria (Orchidaceae, Catasetinae).  Richardiana 13: 226–229.

Tupacamaria Archila comprises 12 spp. recently segregated from Galeandra, the new genus name and all the combinations at sp. rank having been published in issues of the author’s obscure, self-published journal that have yet to arrive at MO’s library.  As it so happens, not all the combinations were published validly, due to “basionym errors,” here rectified for the three affected names.  The combinations successfully validated in previous papers are reiterated here, revealing that Galeandra beyrichii Rchb. f., one of the two spp. treated under Galeandra in the Manual, has been swept up in all of this.

——, G. R. Chiron & D. Szlachetko.  2013. Étude de populations pour une meilleure compréhension du genre Restrepia (Orchidaceae) au Guatemala.  Richardiana 13: 230–243.

Five spp. of Restrepia are recognized for Guatemala, of which three (all apparently endemic) are described as new.  Of interest to us is the authors’ usage of the names Restrepia muscifera (Lindl.) Rchb. f. ex Lindl. and R. xanthophthalma Rchb. f. for distinct spp., the latter having been relegated to synonymy under the former in the Manual treatment by Carlyle Luer.  Perhaps significantly, both names are based on Guatemalan material, and the authors present a detailed tabularized comparison of the two entities as they interpret them.  Unfortunately, however, they give no indication of which name ought to be used for Costa Rican material (both could conceivably apply).  Illustrated with detailed line drawings and photos of flowers in life

Batista, J. A. N., K. S. Borges, M. W. F. de Faria, K. Proite, A. J. Ramalho, G. A. Salazar & C. van den Berg.  2013. Molecular phylogenetics of the species-rich genus Habenaria (Orchidaceae) in the New World based on nuclear and plastid DNA sequences.  Molec. Phylogen. Evol. 67: 95–109.

This study, involving 151 neotropical taxa of Habenaria sensu lato and an additional 20 spp. from the Old World, groups the neotropical spp. with members of five African sections in a “strongly supported ‘core Habenaria clade’…that includes the type species of the genus,” H. macroceratitis Willd. (a New World sp. that occurs in Costa Rica).  According to the authors, their results suggest “an African origin for the Neotropical clade” and “a recent radiation of the genus in the New World.”  While allowing that certain recently championed segregate genera [such as Habenella; see The Cutting Edge 13(2): 13, Apr. 2006] “could become monophyletic with some adjustments,” the authors “do not favor a generic fragmentation of the New World Habenaria,” mainly because the group is already monophyletic (contrary to a previous study involving just eight spp.) and “extensive nomenclatural changes” would result.  Instead, they “favor a revision of the current sectional classification of the Neotropical species,…work that is already underway.”

Benítez de Rojas, C. E., M. Nee & P. Rodríguez.  2012[‘2011’]. Estudio taxonómico de representantes de Solanum sección Dulcamara s.l. (Solanaceae) de Sudamérica tropical/Taxonomic study of members of Solanum section Dulcamara s.l. (Solanaceae) in tropical South America.  Acta Bot. Venez. 34: 381–405.

The titular section, a subset of Solanum subgen. Solanum, comprises about 18 spp., three of which have been chosen (for reasons not adequately explained) to be featured in this work (perhaps because they are widespread?).  Two of these three, Solanum seaforthianum Andrews and S. uncinellum Lindl., are members of the Costa Rican flora.  We detect no noteworthy new developments for us, but are mildly puzzled by the non-appearance of the names Solanum granelianum D’Arcy and S. ipomoea Sendtn., both of which we expected to find in synonymy under S. uncinellum.  It may be that the authors were selective in accounting for synonyms.  Features synonymy and typology, a dichotomous key to spp., detailed descriptions for all three spp. (but only a brief, informal one for the section), exhaustive (though mainly Venezuelan) specimen citations, distribution and phenology summaries, and enumerations of common names and uses.  There are no indices.  The single introductory paragraph summarizes taxonomic history.  Each spp. is illustrated with a composite line drawing, supplemented (except for S. seaforthianum) by a detailed depiction of trichome variation.  See under “Knapp,” this column, for the definitive treatment of these taxa.

Blanco, M. A.  2013. New combinations and synonyms in the Maxillariinae (Orchidaceae).  Selbyana 31: 52–59.

Twelve new combinations are validated in Camaridium, Heterotaxis, Inti, Pityphyllum, and Sauvetrea for spp. originallly described in Maxillaria, and 19 new heterotypic synonyms are proposed.  Just one of the new combinations seems relevant to Costa Rican floristics:  Camaridium hematoglossum (A. Rich. & Galeotti) M. A. Blanco, (inlcuding Maxillaria punctostriata Rchb. f. as a new synonym), which we suspect becomes the accepted name for Costa Rican material treated in the Manual as Maxillaria cucullata Lindl.  Two of the new synonymies entail more or less significant changes in Manual usage:  the name Maxillaria attenuata Ames & C. Schweinf., accepted in the Manual for a sp. questionably endemic to Costa Rica, is consigned to synonymy under M. virguncula Rchb. f. (type from Venezuela), together with M. guadalupensis Cogn. (type from the Lesser Antilles); and Maxillaria shepheardii Rolfe is synonymized under M. ringens Rchb. f., extending the range of the latter sp. southward to Colombia.  Additionally, a lectotype is designated for Maxillaria turialbae Schltr., a name that does not appear in the Manual but which has long beeen accepted as a synonym of Maxillaria friedrichsthalii Rchb. f. [i.e., Rhetinantha friedrichsthalii (Rchb. f.) M. A. Blanco].  As for the finely divided generic reclassification of Maxillaria sensu lato recently published by Dariusz Szlachetko and colleagues (see under “Szlachetko” et al., this column, in our last issue):  this author obviously ignores it, in favor of the more conservative system helmed by himself [see The Cutting Edge 15(2): 3–4, Apr. 2008], noting that “many of [Szlachetko’s genera] are polyphyletic or paraphyletic” and that “a detailed critique to that system will be published separately.”  A battle looms!

Dorsey, B. L., T. Haevermans, X. Aubriot, J. J. Morawetz, R. Riina, V. W. Steinmann & P. E. Berry.  2013. Phylogenetics, morphological evolution, and classification of Euphorbia subgenus Euphorbia.  Taxon 62: 291–315.

We do not generally concern ourselves with infrageneric classification, but this paper features some illuminating insights.  Euphorbia in the broad sense, as accepted by these authors (and in the Manual), comprises about 2000 spp., nearly a third of which are included in the autonymic subgenus.  Four subgenera are recognized, the others being subgen. Athymalus Neck. ex Rchb., subgen. Chamaesyce Raf., and subgen. Esula Pers. (the last of which, not represented in Costa Rica, is the subject of a companion paper in this issue).  Those who would elevate Chamaesyce to genus rank should note that to do so would require, on cladistic grounds, that Athymalus and Esula receive similar treatment (they might also wish to consider that genus Chamaesyce would have to include such unlikely spp. as Euphorbia cotinifolia L.).  Those wishing to resurrect the genus Pedilanthus, currently subsumed (and deeply nested) within subgen. Euphorbia as sect. Crepidaria (Haw.) Baill., should consider that to do so would require that at least seven (and more logically, 20) other clades be elevated to generic rank—even assuming prior recognition for Chamaesyce.  Interestingly, all the New World spp. of subgen. Euphorbia form a single, monophyletic clade.  Includes a dichotomous (though non-indented) key to the 21 sections of subgen. Euphorbia (nine of which are described as new), full synonymy and typology, technical descriptions, comprehensive lists of component spp., distribution summaries, and discussions.

Goldenberg, R., F. Almeda, M. K. Caddah, A. B. Martins, J. Meirelles, F. A. Michelangeli & M. Weiss.  2013. Nomenclator botanicus for the neotropical genus Miconia (Melastomataceae: Miconieae).  Phytotaxa 106: 1–171.

Miconia is characterized by these authors as “the largest genus of woody flowering plants with a distribution restricted to tropical America.”  Their nomenclator comprises four “lists,” the principal one enumerating all 1057 accepted names in the genus alphabetically, with full synonymy for each and typology (except for the synonyms), as well as an indication of sectional affinity and a distribution summary.  The genus and its sections are circumscribed traditionally, notwithstanding recent cladistic studies (here acknowledged) suggesting that they may not be monophyletic.  Supplementing the main alphabetical list are:  a list of spp. organized by sections (there are 12 sections in all, though more than half of the spp. belong to just three of them); a vouchered list of spp. organized by country (with diversity highest in the Andean region); and a list of Miconia names synonymized under other genera.  Lacking is an index of Miconia names synonymized under other Miconia names, which would have been useful.  The country list for Costa Rica totes 111 spp., four more than were included in the Manual treatment (2007) by the second author of this paper.  The discrepancy involves seven recently described spp. and three that have been lost.  The missing spp. are Miconia calvescens DC., M. decurrens Cogn., and M. pileata DC.; the last of these has fallen into synonymy under Miconia ciliata (Rich.) DC. (see under “Goldenberg,” this column, in our last issue), already accounted for in the Manual treatment, but the omission of the other two must certainly be an oversight.  The recent additions to the Costa Rican flora are:  Miconia cocoensis Almeda & Kriebel, M. diegogomezii Kriebel & Almeda, M. hildeana Kriebel & Almeda, M. kappellei Almeda & Kriebel, and M. ricardoi Kriebel & Almeda [see under “Kriebel,” this column, in The Cutting Edge 19(3), Jul. 2012]; M. correae Almeda [see under “Leaps and Bounds” in The Cutting Edge 19(3), Jul. 2012]; and M. osaensis Aguilar, Kriebel & Almeda [see The Cutting Edge 16(1): 7, Jan. 2009].  The treatment of infraspecific taxa in this work is ambiguous.  Subspecies or vars. were accepted in the Manual for Miconia impetiolaris (Sw.) D. Don ex DC., M. lateriflora Cogn., M. melanotricha (Triana) Gleason, M. pileata, and M. tonduzii Cogn.  These are all listed as synonyms under their respective spp. (M. ciliata, in the case of M. pileata), but it is unclear whether this means they are no longer accepted or (as we suspect) were simply ignored for procedural purposes.  The lists are briefly analyzed in the introductory section, which features a table documenting the occurrence of the genus and each of its sections in selected countries and regions.  The authors refrained from designating lectotypes or neotypes in this paper.

Grant, J.  2012. A new short-leaved variety of Werauhia insignis.  J. Bromeliad Soc. 62: 332–333.

The name Werauhia insignis (Mez) W. Till, Barfuss & M. R. Samuel [see The Cutting Edge 11(3): 5–6, Jul. 2004] applies to the sp. referred to as Tillandsia insignis (Mez) L. B. Sm. & Pittendr. in the Manual Bromeliaceae treatment (2003) by Francisco Morales (INB).  The subject of this article, Werauhia insignis var. brevifolia H. Luther ex J. R. Grant, var. nov., is Panamanian, and concerns us only insofar as it establishes the autonymic var. to be used for Costa Rican material.  The “Harry E. Luther Memorial Issue” of which this paper is part includes numerous remembrances and photos of the recently deceased bromeliad illuminary [see under “News and Notes” in The Cutting Edge 19(4), Oct. 2012].

Guo, X., R.-J. Wang, M. P. Simmons, P. P.-H. But & J. Yu.  2013. Phylogeny of the Asian Hedyotis-Oldenlandia complex (Spermacoceae, Rubiaceae):  evidence for high levels of polyphyly and the parallel evolution of diplophragmous capsules.  Molec. Phylogen. Evol. 67: 110–122.

The genus name Hedyotis, long a thorn in the side for New World floristics, may now be forgotten by us, as it applies (according to the conclusions of this study) to a strictly Asian assemblage of spp. (see also under “Wikström,” this column).  Eleven other genera belonging to the complex of the title are confirmed as monophyletic, including Arcytophyllum, Bouvardia, Manettia, and Pentodon (to mention only those occurring in or at least very near Costa Rica).  The problem genus, from our limited perspective, is Oldenlandia, already suggested as polyphyletic in previous papers [see, e.g., The Cutting Edge 16(3): 5, Jul. 2009].  In the present study, “Oldenlandia was resolved as a grossly polyphyletic group consisting of 10 separate lineages.”  The authors aver that “Oldenlandia s. str. should include only the species that are resolved in a clade with the type species,” O. corymbosa L., which happens to be one of the two spp. of Oldenlandia sensu lato recorded from Costa Rica.  The other one, Oldenlandia lancifolia (Schumach.) DC., occurs in a clade over which the genus name Scleromitrion (Wight & Arn.) Meisn. holds dominion (though no combination is yet available for O. lancifolia).  The morphological characters adduced to distinguish Oldenlandia s. str. from Scleromitrion are essentially the same ones that have been used in regional floras to separate O. corymbosa from O. lancifolia.  Both genera are virtually restricted to the Old World, with the exception of the two last-mentioned weedy spp., which have presumably achieved their pantropical distributions in recent times.

Hirai, R. Y. & J. Prado.  2012. Monograph of Moranopteris (Polypodiaceae).  Pteridologia 4: 1–113.

The long-dormant serial Pteridologia (which resurfaces, on average, every 10–12 years) is given new life with this handsome and useful monograph of Moranopteris R. Y. Hirai & J. Prado, another segregate of Grammitis sensu latissimo, corresponding in large measure to Micropolypodium sensu Flora mesoamericana (the last-mentioned genus being now restricted to the Old World).  For further information, see our review of the paper in which the name Moranopteris was validated (under “Hirai,” this column, in The Cutting Edge 18(4), Oct. 2011], where new combinations and distribution summaries were provided for 28 spp. (and one hybrid) assigned to the genus.  No new ground is broken along those lines in the present paper, which features no nomenclatural or taxonomic innovations.  This slim, ergonomic volume (would that something similar were available for every genus!) featues full synonymy and typology, technical descriptions for the genus and each of its spp., a dichotomous and indented key to spp., distribution summaries, selected specimen citations, discussions (mostly rather brief), distribution maps, a section on “Names of uncertain application,” and indices to scientific names and exsiccatae.  The well-illustrated introductory portion addresses taxonomic history, ecology, geography, and morphology.  A pair of convenient tables reveals that there are 12 Moranopteris spp. in the Mesoamerican region, 11 of which occur in Costa Rica, and that the Costa Rican endemic M. zurquina (Copel.) R. Y. Hirai & J. Prado is also the only sp. endemic to the region as a whole.  The specimen citations provide at least a partial answer to the question posed in our earlier review as to what had become of Micropolypodium setulosum (Rosenst.) A. R. Sm., used as an accepted name in Flora mesoamericana for a sp. attributed to Costa Rica.  Here, it is one of the three “Names of uncertain application,” banished to that category because the authors have not seen adequate type material.  They speculate that it may correspond to Moranopteris plicata (A. R. Sm.) R. Y. Hirai & J. Prado (for which it would be an older name), although the type locality (656 m, El General) would suggest (to us) M. nana (Fée) R. Y. Hirai & J. Prado; nevertheless, the Costa Rican voucher for Micropolypodium setulosum cited in Flora mesomericana is here identified as Moranopteris basiattenuata (Jenman) R. Y. Hirai & J. Prado—a sp. that was not attributed to Costa Rica in Flora mesoamericana.

The subject of specimen citations brings up what may be perceived as a shortcoming of this contribution:  only a small percentage of the available herbarium material appears to have been revised, at least for southern Central America.  Specimens from the region Nicaragua through Panama are cited from the following rather peculiar selection of herbaria:  B, GOET, K, L, LP, M, NY, RB, SP, and UC (with a few types also from P, S, and US).  Conspicuously missing in action are CR, F, HNMN, INB, JVR, MO, PMA, SCZ, US (except for types), and USJ, which together would house the richest source of material from the aforementioned countries.  It is true that many duplicates from those herbaria have been distributed to the herbaria consulted by the authors, especially (in the case of ferns) NY and UC; however, that would generally require the availability of at least three duplicates, seldom the case for small epiphytes like Moranopteris.  To cite an example that is close to home:  Manual co-PI Mike Grayum, not the most heroic collector of grammitid ferns, has accounted for at least 16 collections of Moranopteris from Costa Rica; yet, just two of these are cited in this monograph (one in the text, the other only in the index to exsiccatae).  If that sample may be taken as representative, then the authors have seen only the tip of the iceberg for southern Central America, acknowledged by them as a “primary center of diversity” for the genus.  Be that as it may, we doubt that the grand, overall picture would be much affected by the examination of additional material, hence can recommend this publication with little reservation.  Oh, and by the way:  each sp. is illustrated with an excellent, full page, composite line drawing.

Hong-Wa, C. & G. Besnard.  2013. Intricate patterns of phylogenetic relationships in the olive family as inferred from multi-locus plastid and nuclear DNA sequence analyses:  a close-up on Chionanthus and Noronhia (Oleaceae).  Molec. Phylogen. Evol. 67: 367–378.

Trouble looms for the moderately large (ca. 100 spp.) genus Chionanthus, with two spp. in Costa Rica.  The authors’ data discriminate four distinct “geographically structured” clades, leading them to “recommend restrictions in the use of the generic epithet ‘Chionanthus’.”  Undeterred by “the possible lack of robust morphological synapomorphies,” the authors transfer all the spp. of the Africa-Indian Ocean clade to Noronhia, validating the necessary new combinations.  They also note that “further extension of Noronhia” to the Asia-Pacific clade of Chionanthus “is a possibility.”  The New World members of Chionanthus account for the two remaining clades, Central America and North America.  The latter harbors the generic type sp., Chionanthus virginicus L., suggesting that the Central American contingent [represented in this study by C. panamensis (Standl.) Stearn] may ultimately wind up in a different genus.

Klopper, R. R., G. F. Smith & A. E. van Wyk.  2013. (2129) Proposal to conserve the family name Asphodelaceae (Spermatophyta: Magnoliidae: Asparagales).  Taxon 62: 402–403.

The family currently accepted by the Angiosperm Phylogeny Group (APG) under the name Xanthorrhoeaceae comprises several genera (e.g., Aloe, Hemerocallis, Kniphofia, and Phormium) that are at least sparingly cultivated in Costa Rica.  In prior APG classifications, this assemblage was apportioned (at least optionally) among three smaller families:  Asphodelaceae, Hemerocallidaceae, and Xanthorrhoeaceae sensu stricto.  These authors, outraged that Xanthorrhoeaceae is “the smallest, youngest, and least-known constituent” of the present conglomerate (it prevails only through conservation), propose to replace it with the largest, oldest, and (arguably) best-known of the three.  What is the point of all this?  Why are proposals that seek to undermine the established consensus even entertained?  Is not nomenclatural stability a principal objective of the IAPT?  We have all dutifully bought into Xanthorrhoeaceae.  Now leave it be!

Knapp, S.  2013. A revision of the Dulcamaroid clade of Solanum L. (Solanaceae).  PhytoKeys 22: 1–432.

The informally named “Dulcamaroid clade” comprises 45 Solanum spp. previously apportioned among several infrageneric taxa that have since been shown to be non-monophyletic.  One of these, sect. Holophylla (G. Don) Walp., was partially revised by the author some time ago [Knapp, 1989; Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 63–112].  The Dulcamaroid clade is best characterized by “a distinctive pedicel ‘sleeve’…that appears to be an extension of the inflorescence axis in which the pedicel base sits.”  Additional distinguishing features include a “mostly vining or weakly scandent” habit, twining petioles, pinnatifid leaf-blades (at least in juvenile plants), terminal, large, branched inflorescences, and brightly colored fruits.  The author discriminates five (according to the Abstract) or nine (according to Table 4) informal sp. groups, though little use seems to be made of these.  The Dulcamaroid clade is circumtemperate and widely distributed throughout the Western Hemisphere, but is best represented in South America.  A handy table (Table 2) quickly reveals that just five spp. are recorded from Costa Rica:  Solanum aligerum Schltdl., S. pubigerum Dunal, S. seaforthianum Andrews (probably introduced, at least in part), S. storkii C. V. Morton & Standl., and S. uncinellum Lindl. (with S. granelianum D’Arcy and S. ipomoea Sendtn. Duly included as synonyms of the last).  None of these is endemic to the country (though S. storkii comes close).  The taxonomic treatment kicks off with formal descriptions of Solanum (including traditional segregates such as Cyphomandra and Lycopersicon) and the Dulcamaroid clade, plus synonymy and  typology of the latter, an artificial, dichotomous (though non-indented) key to its spp. and a synoptic character list.  The sp. entries, ordered alphabetically, each features full synonymy and typology, a lengthy description, a distribution summary and discussion, representative (though often extensive) specimen citations, a composite line drawing, and a distribution map.  Rounding out this deluxe treatment are a section on doubtful, excluded, and invalid names and indices to exsiccatae and scientific names.  The detailed and well-illustrated introductory portion expounds upon taxonomic history, morphology, biology and natural history, and sp. concepts.  Preliminary conservation assessments for each sp. are presented in Table 3 (these are also indicated, with commentary, in the sp. entries).  Just one new (Ecuadorean) sp. is described, but numerous lecto- and neotypifications are effected.

Kriebel, R. & D. Santamaría.  2013. A new species of Blakea (Melastomataceae) from Panama with foliaceous sepal appendages and zygomorphic flowers.  Brittonia 65: 1–4.

Despite the fact that both of its known collections are from Panama (Prov. Bocas del Toro), Blakea bocatorena Kriebel & D. Santam., sp. nov., bears mentioning in these pages because of the proximity of its type locality to the Costa Rican border:  within 10 km, in a habitat suggesting a strong likelihood that it will eventually turn up in Costa Rica.  As in the case of Blakea hammettiorum (see under “Almeda,” this column), the spp. with which it is compared (B. calycosa Gleason, B. tuberculata Donn. Sm., and B. wilburiana Almeda) suggest that B. bocatorena would have been included in Blakea prior to the absorption by that genus of Topobea.  Illustrated with a color plate of (mostly) living material.

Larridon, I., K. Bauters, M. Reynders, W. Huygh, A. M. Muasya, D. A. Simpson & P. Goetghebeur.  2013. Towards a new classification of the giant paraphyletic genus Cyperus (Cyperaceae):  phylogenetic relationships and generic delimitation in C4 Cyperus.  Bot. J. Linn. Soc. 172: 106–126.

The key results of this molecular study are briefly stated, appropriately, in the abstract:  “Species of C4 Cyperus s.s. form a core grade in which the C4 segregate genera are embedded.  Pycreus, the largest segregate genus composed of c. 120 species, is not monophyletic as it includes several C4 species of Cyperus s.s.”  Even though most of the other C4 segregate genera are monophyletic, the authors suggest that all nine “should be included in a more broadly circumscribed Cyperus,” due to the fact that “they are nested in the Cyperus clade, and as a consequence of the multiple origins of the characters used to circumscribe them.”  It is good to learn that Pycreus does not pass muster, as it was not recognized in Jorge Gómez-Laurito’s (USJ) Manual treatment of Cyperaceae (2003).  However, the eight smaller C4 segregate genera include Ascolepis, Lipocarpha, Kyllinga, and Remirea, all of which were accepted by Jorge (with Ascolepis merely mentioned in the family discussion).  No new combinations are proposed here (assuming any would be needed).

Lindström, A. J., M. Calonje, D. Stevenson, C. Husby & A. Taylor.  2013. Clarification of Zamia acuminata and a new Zamia species from Coclé Province, Panama.  Phytotaxa 98: 27–42.

Zamia acuminata Oerst. ex Dyer (Zamiaceae) has long been a perplexing entity, its type ostensibly hailing from a region (along the Río San Juan) from which no modern collections resembling it have been forthcoming.  In recent times, the name has been applied most consistently to a sp. of central Panama.  In the Manual treatment of Zamiaceae (2003), MO colleague Mary Merello restricted her application of the name in Costa Rica to the type specimen and based her description mostly on the Panamanian material, while acknowledging the possibility that the two populations may not be conspecific.  The last-mentioned notion is endorsed by these authors, who recognize the Panamanian population as a different sp., described as new.  More significantly, for us, they appear to have made a major breakthrough in nailing the identity of the real Zamia acuminata:  the (sterile) type specimen is a perfect match for a suite of modern collections from ca. (100–)350–1000 m elevation on the central Pacific slope of Costa Rica (mainly in the Zona Protectora El Rodeo, Parque Nacional Carara, and Parque Nacional La Cangreja).  This revelation involves two major paradigm shifts:  first, the label of the type specimen, specifying an Atlantic lowland locality, is erroneous, or has been misinterpreted; and second, the Pacific-slope collections now assigned to Z. acuminata differ from the well-known Z. fairchildiana L. D. Gómez, to which they have generally been referred.  The authors enumerate several differences in defense of the latter point, perhaps the most significant being the presence of microsporangia only on the abaxial surface of the sporophyllys in the microstrobili of Z. acuminata (vs. on both the abaxial and adaxial surfaces in Z. fairchildiana and Z. pseudomonticola L. D. Gómez ex D. W. Stev. & Sabato, also from the Pacific slope).  We are impressed and persuaded by the authors’ equation of Zamia acuminata with material from the Costa Rican Pacific slope, less so by their separation of yet another entity from the last-mentioned region; in other words, we wonder whether they have simply found an older name for Z. fairchildiana.  For us, the jury is still out on that issue (not to mention the distinctness of Z. pseudomonticola, which was not accepted in the Manual).  Also of interest:  while allowing that it is a somewhat anomalous specimen, the authors question the identity of the sterile Aguilar et al. 2970, cited in the Manual as the voucher and only known Costa Rican collection of Zamia obliqua A. Braun.  They reckon it “more likely represents a juvenile Zamia fairchildiana, as this species is extremely abundant throughout the coastal Golfo Dulce region where this specimen was collected.”  Time will tell (and we have no hardened opinions on this).  Includes a revised description of Zamia acuminata, distribution and phenology summaries, comprehensive specimen citations, an image of the type specimen (with leaflets from modern collections superimposed), distribution maps, and photos of living plants in situ (most of these features being provided as well for the new Panamanian sp.).  An appendix enumerates the known Zamia specimens that have been collected from the vicinity of the supposed type locality (Río San Juan) of Z. acuminata—all of which, tellingly, answer to the description of the very different Z. neurophyllidia D. W. Stev.

[Lodé, J.]  2013. New combinations.  Cact.-Avent. Int. 98(Suppl.): 2–11.

Numerous new combinations are validated “for taxonomical reasons,” with no other rationale presented.  We sense that the author is seizing credit for the conclusions of previously published cladistic studies in which he played no role, e.g., one in which it was suggested that certain spp. ostensibly belonging to Peniocereus were in fact nested in Acanthocereus [see The Cutting Edge 13(2): 7, Apr. 2006].  At any rate, eight of the new combinations are in that particular vein, including Acanthocereus hirschtianus (K. Schum.) Lodé, which would replace Peniocereus hirschtianus (K. Schum.) D. R. Hunt for a sp. occurring in Costa Rica should the aforementioned scenario prove correct.  None of the other new combinations appears to concern us.

We have placed the author’s name in square brackets because no authorship is indicated specifically for this article; however, Lodé is the editor of the journal, so must be presumed responsible.

Meseguer, A. S., J. J. Aldasoro & I. Sanmartín.  2013. Bayesian inference of phylogeny, morphology and range evolution reveals a complex evolutionary history in St. John’s wort (Hypericum).  Molec. Phylogen. Evol. 67: 379–403.

We previously reported on a study [see this column under “Ruhfel” in The Cutting Edge 18(2), Apr. 2011] that suggested the enlargement of Hypericum to accommodate various satellite genera, including both Thornea and Triadenum.  The paper presently under consideration seconds the inclusion of the Triadenum, but not necessarily that of Thornea.

Moura, T. M., A. M. G. A. Tozzi, V. F. Mansano & G. P. Lewis.  2013. Lectotypification of names of Neotropical species of Mucuna (Leguminosae: Papilionoideae: Phaseoleae).  Taxon 62: 391–393.

Three of the seven names lectotypified apply to spp. represented in Costa Rica:  Mucuna andreana Micheli [a synonym of M. holtonii (Kuntze) Moldenke], M. sloanei Fawc. & Rendle, and Negretia mutisiana Kunth [the basionym of Mucuna mutisiana (Kunth) DC.].  As far as we can tell, the application of these names is not affected; however, the authors make the point that Mucuna sloanei is not to be construed as a nomen novum based on Dolichos urens Jacq., non L. (as indicated in the Manual), rather a new sp. name validated by reference to Jacquin’s previously published diagnosis (in which he misapplied the Linnaean name).

——, N. A. Zamora, G. P. Lewis, V. F. Mansano & A. M. G. A. Tozzi.  2013. Mucuna globulifera (Leguminosae: Papilionoideae), a new species from Costa Rica, Panama and Colombia.  Kew Bull. 68: 151–155.

Mucuna globulifera T. M. Moura, N. Zamora & A. M. G. Azevedo corresponds to Mucuna sp. B of co-PI Nelson Zamora’s (INB) Manual Fabaceae treatment (2010).  On the basis of new information presented here, the Manual entry for sp. B should be adjusted for elevation (now correctly “900–1200 m”), phenology ( “jun., jul.”), and overall geographic range (“CR–Col.”).  The new sp. is most similar to the Colombian Mucuna mollis (Kunth) DC., though numerous differences are adduced.  Features a distribution map, composite line drawing, and color photos of living material.

Nelson, E. C.  2013. Archibald Menzies’s visit to Isla del Coco, January 1795.  Arch. Nat. Hist. 40: 149–156.

The title of this article already provides information about Archibald Menzies that was not available in Manual Vol. 1 (p. 3), where he was said to have landed on Isla del Coco “in 1794 and/or 1795.”  Now we have the precise dates of his visit, 20–27 January, 1795, during which time Menzies walked on shore just twice (on 23 and 26 January).  He collected plants only at Bahía Chatham.  His “more than two dozen” surviving herbarium specimens from the island, enumerated by taxon in annotated fashion, comprise 12 spp.:  one bryophyte, seven pteridophytes, and four angiosperms.  Already at that early date, Menzies and his men “saw evident marks of the Axe, & saw,” as well as rats and at least one feral hog.  They even found a note, in a bottle, from a man who boasted of having released hogs and goats on the island (a deed deemed commendable by Menzies).  According to this source, two vessels were involved in the reconnaissance of Isla del Coco, the Chatham and the Discovery (only the latter was named in the Manual), though we do not learn which of these carried Menzies.  Curiously, no mention is made here of William Dampier, who visited Isla del Coco more than a century before Menzies (and is known to have collected plant specimens in “Costa Rica”)—even though the very pages of the Manual on which the details about Dampier appear are cited among the “References” as, indeed, is Dampier’s book itself (though the latter does not seem to be cited anywhere in the text).  Still, this is a most welcome contribution to the history of botanical exploration in Costa Rica.

Pastore, J. F. B.  2013. A review of Vellozo’s names for Polygalaceae in his Flora Fluminensis.  Phytotaxa 108: 41–48.

As far as we can tell, the only conclusion of this paper even tangentially relevant to Costa Rican floristics is the resurrection of Polygala appendiculata Vell. (1829) to become the accepted name for P. leptocaulis Torr. & A. Gray (1838).  However, the last-mentioned name is treated as a synonym of Polygala tenella Willd. (1802) in the Manual draft of Polygalaceae by Francisco Morales (INB), so we are not concerned.

Penneys, D. S. & W. S. Judd.  2013. Combined molecular and morphological phylogenetic analyses of the Blakeeae (Melastomataceae).  Int. J. Pl. Sci. 174: 802–817.

This, “the first phylogenetic analysis of the Blakeeae using combined molecular and morphological characters,” corroborates a prior cladistic analysis of morphological characters only [see under “Penneys,” this column, in The Cutting Edge 18(2), Apr. 2011] in concluding that “Topobea must be transferred to Blakea.”  Good thing, too, because all the consequent new combinations have already been published (see under “Penneys,” this column, in our last issue)!

Pennington, T. D. & J. J. Clarkson.  2013. A revision of Guarea (Meliaceae).  Edinburgh J. Bot. 70: 179–362.

Pursuant to the “considerable number of new species” published during the past three decades, “recent molecular studies [that] have shown that Guarea is now an exclusively American genus” [see under “Koenen,” this column, in The Cutting Edge 19(4), Oct. 2012], and (we wager) widespread repudiation of his treatment in Flora Neotropica Monographs (1981), Terry Pennington (K) takes another crack (with some help) at revising this difficult genus.  The stingy specimen citations and general lack of insightful discussions, in conjunction with our own lack of expertise on some of the issues, make it difficult for us to fully evaluate the results of this study vis-à-vis the Costa Rican repesentatives of Guarea; we are confident that it represents a significant improvement over the initial effort, but put us down as skeptical on several counts.  Chief among these is the authors’ continued use of the name Guarea glabra Vahl in a very broad sense (albeit not so broad as before), to include several morphologically and ecologically distinctive entities, six of which are now distinguished at the rank of subsp.  Four of the latter are attributed to Costa Rica and correspond to the spp. treated (or mentioned) in Alexánder (‘Popeye’) Rodríguez’s Manual treatment (2007) of Meliaceae under the names Guarea ciliata Al. Rodr., G. excelsa Kunth, G. microcarpa C. DC., and G. tuerckheimii C. DC.  Combinations based on those names are validated at subsp. rank in the name of the first author, except in the case of G. ciliata, which is synonymized under Guarea glabra subsp. glabra—together with G. ternifoliola C. DC., a synonym of G. excelsa according to Popeye.  And therein lies a problem, as Costa Rican specimens representing both G. ciliata and G. excelsa (by our reckoning) are cited under G. glabra subsp. glabra in this work; that the authors do not consistently distinguish these two entities, quite different in our opinion, does not augur well for their association of both with G. glabra (based on Antillean material), and calls into question the extension of that name to embrace several other equally distinctive taxa.  Guarea tuerckheimii, by the way, would be an addition to the Costa Rican flora, although at least one of the two relevant vouchers has been determined as G. microcarpa by Popeye.  Likewise, Guarea macrophylla Vahl subsp. pendulispica (C. DC.) T. D. Penn. would be new to Costa Rica, ignoring the fact that Popeye has redeterimined both of its Costa Rican vouchers as G. guidonia (L.) Sleumer.  We are unable to arbitrate these last two cases, as well as the following four, in which spp. recognized as distinct by Popeye are consigned to synonymy in this revision:  Guarea adenophylla Al. Rodr. (now a synonym of G. tonduzii C. DC.), G. montana Al Rodr. (a synonym of G. rhopalocarpa Radlk.), G. pilosa Al. Rodr. (a synonym of G. chiricana Standl.), and G. williamsii C. DC. (presumably a synonym of G. kunthiana A. Juss., although synonymies unchanged from the Flora Neotropica Monographs opus are not repeated here).  Taxonomic concepts remain substantially the same for the 19 other Guarea spp. accepted in the Manual, with the following caveats:  Guarea donnell-smithii C. DC. is now reported from Guatemala and Honduras, signifying the loss of a Costa Rican endemic; the name Guarea kegelii Turcz. was apparently misapplied in the Manual, to the sp. that is here called G. bijuga C. DC. (with G. kegelii now assigned questioningly to synonymy under G. glabra subsp. tuerckheimii); the description of a new, Colombian subsp. of Guarea pterorhachis Harms consigns Costa Rican material to the autonymic subsp. (with essentially the same geographic range as the sp.); and Guarea sp. A of the Manual now has a name, viz., G. grossa T. D. Penn., validated herein (though without an appropriate cross-reference to the provisional Manual name).  Guarea arcuata Coronado, merely mentioned in the Manual (though based on a Costa Rican type), is synonymized under G. glabra subsp. microcarpa, a disposition foreshadowed by Popeye’s comments.  All in all, 69 spp. are accepted in the genus (vs. 75 according to the Manual), of which 25 (plus three subspp.) are attributed to Costa Rica (vs. 30 according to the Manual).  We thus show a net loss of two taxa.  Careful readers will detect innumerable other minor changes, e.g., in geographic distributions both within and outside Costa Rica.  Features technical descriptions at all ranks, a dichotomous (though non-indented) key to spp. (but none for subspp.), synonymy (if different from Flora Neotropica Monographs) and typology, distribution summaries, representative specimen citations, discussions (often too brief), a list of “doubtful” spp., distribution maps (much reduced and grouped at the end), and indices to exsiccatae and scientific names (the latter excluding many names, such as Guarea williamsii, not explicitly cited in the present work).  The introductory portion is mainly devoted to phylogenetic analysis (in a section separately authored by a team headed by the second author of the paper).  Six new spp. and one new subsp. are described, and five new combinations are validated at subsp. rank (the novelties pertinent to Costa Rica having already been mentioned).  The majority of spp. (including all of the new taxa) are illustrated by excellent composite line drawings.

Prado, D. E., L. J. Oakley & J. Prado.  2013. 2133) Proposal to conserve the name Capparis baducca (Capparaceae) with a conserved type.  Taxon 62: 406–407.

The name Capparis baducca L. (1753) was based on mixed material representing two morphologically distinct spp., one endemic to southern India and the other widespread in the Neotropics.  According to its presently accepted lectotypification, C. baducca correctly applies to the Indian sp.; however, citing conflict with the Linnaean diagnosis and current usage (the Indian sp. has become known universally as Capparis rheedei DC.), these authors propose replacement of the reigning lectotype with another representing the neotropical sp.  The neotropical sp. in question has otherwise been known as Capparis frondosa Jacq. (1760), or, more recently, Capparidastrum frondosum (Jacq.) Cornejo & Iltis [see The Cutting Edge 16(2): 4, Apr. 2009]; indeed, C. frondosa is the accepted name used in all the major, regional floras routinely consulted by us, with the exception of Flora of Guatemala, and is the only name we have ourselves used for the sp.  Therefore, from our narrow perspective, the proposed switch would represent a step backward.

Rojas-Alvarado, A. F.  2012. Novedades en Diplazium (Woodsiaceae) con frondas pinnado-lobuladas para el Neotrópico/Novelties in Diplazium (Woodsiaceae) with pinnate-lobulate fronds from the Neotropics.  Actual. Biol. (Medellín) 34: 153–159.

Two new spp. and one new var. are described here, and an existing var. is elevated to sp. rank.  Only one of these novelties directly impacts Costa Rican floristics:  Diplazium salazarianum A. Rojas, endemic to Costa Rica, where it occurs at 750–1300+ m elevation on the Atlantic slope of the Cordilleras de Guanacaste, de Tilarán, and Central.  It has been confused with D. neglectum (H. Karst.) C. Chr. and (as in Flora mesoamericana) D. urticifolium, from both of which it differs in frond architecture and other details that are duly enumerated.  More indirectly, the validation of Diplazium urticifolium Christ var. mesoamericanum A. Rojas (of southern Mexico to Nicaragua) establishes its autonymic counterpart, presumably applicable to Costa Rican material.  Also of passing interest is the author’s allegation that four specimens from the Atlantic lowlands of Costa Rica (including one from the Estación Biológica La Selva) comprise the first report of Diplazium moccennianum (Sodiro) C. Chr. (based on an Ecuadorean type) from the country.  However, F. Araya & Rojas 902 (MO), a specimen from the same region not cited here, was determined by the author as D. moccennianum way back in 1996.  Critical features of all the new taxa (with comparative material of some close relatives) are depicted in a composite line drawing.

——.  2012. Nuevas combinaciones en Alansmia M. Kessler, Moguel, Sundue et Labiak (Polypodiaceae) para Costa Rica, Panamá y Colombia/New combinations in Alansmia M. Kessler, Moguel, Sundue et Labiak (Polypodiaceae) from Costa Rica, Panama, and Colombia.  Actual. Biol. (Medellín) 34: 161–169.

Combinations in the recent Terpsichore (i.e., Grammitis sensu lato) segregate Alansmia that were not validated at the time the last-mentioned genus was established [see under “Kessler,” this column, in The Cutting Edge 18(4), Oct. 2011] are here realized in the name of the author.  The names involved are the former Terpsichore acrosora A. Rojas, T. cocosensis A. Rojas, T. esquiveliana A. Rojas, T. fabispora (Copel.) A. Rojas, and T. lobulata A. Rojas.  It bears mentioning that these names were passed over in the earlier paper because they were consigned to synonymy, as follows:  T. acrosora under Alansmia longa (C. Chr.) Moguel & M. Kessler; T. cocosensis, T. esquiveliana, and T. lobulata under Alansmia elastica (Bory ex Willd.) Moguel & M. Kessler; and T. fabispora under Alansmia stella (Copel.) Moguel & M. Kessler.  The present author seizes this opportunity to reemphasize the distinctions he deems significant, supplementing his narrative with a dichotomous (though non-indented) key to the spp. of Alansmia known from Costa Rica, Panama, and Colombia, as well as a composite line drawing of critical pinna features.  Geographical distributions are specified in a tabular format.

—— & C. G. Herrera Martínez.  2012. Un nuevo híbrido de Adiantum L. (Polypodiales: Pteridaceae) para Costa Rica y Nicaragua.  Brenesia 78: 1–5.

The supposed new hybrid, Adiantum ×janzenianum A. Rojas & C. G. Herrera, restricted to the countries mentioned in the title, is distributed in Costa Rica according to the following Manualese formula:  “bosque seco y húmedo, 0–1000 m; vert. Pac. Cords. de Guanacaste y de Tilarán, llanuras de Guanacaste.”  The hybrid nature of this entity is suggested by its morphological intermediacy (documented in a table and two composite line drawings) with respect to the proposed parent spp., Adiantum concinnum Humb. & Bonpl. ex Willd. and A. deflectens Mart., as well as by the observation, in the type material, of numerous collapsed sporangia and others with empty spores (depicted in photos).  The epithet honors Daniel H. Janzen, a tireless advocate of conservation interests in the Guanacaste region.

Rolleri, C. H., C. Prada, J. M. Gabriel y Galán & L. M. Passarelli.  2013. Especies arborescentes del género Blechnum (Blechnaceae: Pteridophyta).  Revista Biol. Trop. 61: 377–408.

The authors have selected eight arborescent Blechnum spp. “para una caracterización y revisión,” but it is unclear whether this represents a natural group or even a comprehensive sampling of the category in question.  Business as usual for these authors.  For whatever it may be worth, the eight spp. are distinguished by means of a dichotomous (though non-indented) key, with synonymy and typology, technical descriptions, distribution summaries, and representative specimen citations provided for each (as well as “observaciones,” in some cases).  Just two of these spp. are attributed to Costa Rica:  Blechnum schomburgkii (Klotzsch) C. Chr. and B. werckleanum (Christ) C. Chr.  The former name will be unfamiliar to most of our readers, as it is newly applied to the sp. (well-known in Costa Rica and conspicuous on Cerro de La Muerte) long known (e.g., in Flora mesoamericana) as Blechnum buchtienii Rosenst. and, more recently, as B. auratum (Fée) R. M. Tryon & Stolze [see The Cutting Edge 14(3): 12–13, Jul. 2007].  The inclusion of B. werckleanum in this study is a mild surprise for us because, according to our experience with that sp. in Costa Rica (whence the type), it is not notably arborescent, nor is it generally terrestrial, as indicated here; rather, it is a high-growing epiphyte, encountered on the ground (we suspect!) only as a consequence of tree-falls.  The introductory portion of this paper features many illustrations, mostly depicting epidermal features (as is the habit of these authors).

Vázquez-García, J. A., M. A. Muñiz-Castro, E. De Castro-Arce, R. Murguía Araiza, A. T. Nuño Rubio & M. de J. Cházaro-B.  2012. Twenty new Neotropical tree species of Magnolia (Magnoliaceae).  Pp. 91–130 in, E. Salcedo-Pérez, E. Hernández-Álvarez, J. A. Vázquez-García, T. Escoto-García & N. Díaz-Echavarría (eds.), Recursos forestales del occidente de México:  diversidad, manejo, producción, aprovechamiento y conservación, Vol. 4, Tomo I.  Univ. Guadalajara [Mex.].

Most of the new Magnolia spp. described in this book chapter—and all of the six described from Costa Rica—would formerly have been assigned to the genus Talauma, here tentatively demoted to a subsection (or two?) of Magnolia, notwithstanding molecular evidence that generic rank may be tenable for at least the New World contingent [see, e.g., The Cutting Edge 9(1): 3, Jan. 2002].  The six Costa Rican novelties, all endemic, were universally accommodated within Talauma gloriensis Pittier in the Manual treatment of Magnoliaceae (2007) by José González (LSCR), although T. gloriensis was characterized as “bastante variable” and to perhaps harbor two distinct spp.  We have good reason to believe that the second sp. envisioned by José is the one here described as Magnolia costaricensis A. Vázquez, which is fairly common at the Estación Biológica La Selva and ranges up to 1000 m elevation on the Atlantic slope of the Cordillera Central and the northern Cordillera de Talamanca, with an outlying population on the Llanura de Tortuguero.  It is compared with the Ecuadorean M. rimbachii (Lozano) Govaerts.  The other five new Costa Rican spp., unforeseen by anyone connected with the Manual project, are as follows:  Magnolia guanacastensis A. Vázquez, from 700–800+ m elevation on the Atlantic slope of the Cordillera de Guanacaste (and de Tilarán?), compared with M. mexicana DC.; M. inbioana A. Vázquez, from 50–700 m elevation on the Atlantic slope of the Cordillera de Guanacaste and northern Cordillera de Talamanca, and on the Llanura de San Carlos, compared with the Ecuadorean M. equatorialis A. Vázquez (published in this same chapter); M. multinervia A. Vázquez and M. talamancana A. Vázquez, from 450–600 m elevation on the Atlantic slope of the eastern Cordillera de Talamanca, compared with the Mexican M. sinacacolinii A. Vázquez (published in this same chapter) and M. costaricensis, respectively; and M. wetteri A. Vázquez, from 100–650 m elevation on the Península de Osa, compared with a yet-to-be-described Mexican sp. (note that we have corrected the author’s spelling of “wetterii,” according to ICN Art. 60.12).  Most of the new spp. are validated on the basis of brief diagnoses and one or a few collections and illustrated with poorly reproduced (at least in our copy) photos of herbarium specimens.  It goes without saying that Talauma gloriensis is still accepted [as Magnolia gloriensis (Pittier) Govaerts] by these authors in the strict sense of its Costa Rican type; thus the Costa Rican sp. total for Magnoliaceae rockets from four (according to the Manual) to 10 (and recent correspondence with the first author of this paper suggests that at least five more new Costa Rican spp. of Magnolia sensu lato may be in the works).

Vázquez-Sánchez, M., T. Terrazas, S. Arias & H. Ochoterena.  2013. Molecular phylogeny, origin and taxonomic implications of the tribe Cacteae (Cactaceae).  Syst. Biodivers. 11: 103–116.

The only genus of tribe Cacteae represented in Costa Rica is Mammillaria [see The Cutting Edge 15(4): 4, Oct. 2008] which, wouldn’t you know it, “is likely polyphyletic” as currently circumscribed.  At any rate, it would appear to be paraphyletic with respect to the patently dubious generic entity Mammilloydia.  Whatever the case, the authors opine that “recognition of the genus is suggested,” in view of the fact that the generic type, Mammillaria mammillaris (L.) H. Karst., “is deeply nested within the Mammillaria clade” (together, we might add, with M. columbiana Salm-Dyck, conspecific with or at least closely related to the Costa Rican plants).

Vela Díaz, D. M.  2013. Multivariate analysis of morphological and anatomical characters of Calophyllum (Calophyllaceae) in South America.  Bot. J. Linn. Soc. 171: 587–626.

Although it is not made clear in the title, this contribution by MO student Dilys Vela provides a taxonomic revision of Calophyllum for South America, the first since 1893 (according to the author).  Two of the four accepted sp. names, Calophyllum brasiliense Cambess. and C. longifolium Willd., were also used in the Manual treatment of Calophyllum (2010) by co-PI Barry Hammel (wherein the genus was still included in Clusiaceae).  However, the name Calophyllum longifolium is applied in a different sense by the present author, who restricts it to South American material.  She distinguishes Central American material (Costa Rica and Panama) formerly assigned to C. longifolium as a new sp., Calophyllum mesoamericanum Vela Díaz, on the basis of its leaf-blades with inconspicuous secondary veins and flowers with six (vs. eight) tepals and a longer stigma, as well as certain anatomical differences.  The taxonomic treatment (which omits C. mesoamericanum) features a dichotomous key to spp., synonymy and typology, generous descriptions, distribution and phenology summaries, and discussions of variable length.  Includes distribution maps for all spp., and rather basic line drawings of the two spp. described as new (the other of which does not concern us).  Specimens (other than types) are cited skeletally in two appendices.  The introductory portion is concerned mainly with the statistical analysis and morphological and anatomical details.

Wikström, N., S. Neupane, J. Kårehed, T. J. Motley & B. Bremer.  2013. Phylogeny of Hedyotis L. (Rubiaceae: Spermacoceae):  redefining a complex Asian-Pacific assemblage.  Taxon 62: 357–374.

This study, though undertaken independently of a similar one reviewed in this column (see under “Guo”), arrived at grossly similar conclusions:  Hedyotis in its broadest sense (i.e., including the likes of Arcytophyllum, Houstonia, and Oldenlandia) is polyphyletic, while Hedyotis sensu stricto is an excluvisely Asian/Pacific taxon.  Furthermore, Oldenlandia lancifolia (Schumach.) DC. belongs to a clade, distantly removed from that harboring O. corymbosa L. (the generic type), for which no genus name is proffered (but see this column under “Guo”).  It would seem that a consensus is finally being approached for this most difficult group.

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