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Manual de Plantas de Costa Rica

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The Cutting Edge

Volume XI, Number 3, July 2004

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

Almeda, F. 2004. Novelties and nomenclatural adjustments in the neotropical genus Clidemia (Melastomataceae: Miconieae). Proc. Calif. Acad. Sci. 55: 89–124.

Six of the eleven new Clidemia spp. published here occur in Costa Rica, but only one is endemic. The latter is Clidemia rodriguezii Almeda (dedicated to INBio curator Alexánder Rodríguez), known from both slopes of the northern Cordillera de Talamanca, and most closely related to C. discolor (Triana) Cogn. Shared with Panama are Clidemia allenii Almeda, of the Golfo Dulce region, compared with C. costaricensis Cogn. & Gleason and C. petiolaris (Schltdl. & Cham.) Schltdl. ex Triana; Clidemia coloradensis Almeda, from the Atlantic slope of the northern Cordillera de Talamanca, with “no close relatives in any part of the American tropics”; and Clidemia lanuginosa Almeda, from the Atlantic slope of the eastern Cordillera de Talamanca, “readily recognized” but not compared with any other sp. Clidemia davidsei Almeda, most similar to C. dentata Pav. ex D. Don, is known mainly from the Pacific slope of the Cordillera de Talamanca and the southern Fila Costeña (Fila Cruces), but also by a single, somewhat aberrant collection from northern Nicaragua. Finally, Clidemia evanescens Almeda, most similar to C. densiflora (Standl.) Gleason, is disjunct in northern Costa Rica (Atlantic slope of the Cordilleras de Guanacaste and de Tilarán) with respect to the largest portion of its geographic range, from central Panama to southwestern Colombia. The lone “nomenclatural adjustment” is the validation of the new combination Clidemia quinquenervia (Mill.) Almeda (based on Melastoma quinquenervium Mill.) for a common sp. long known as Ossaea quinquenervia (Mill.) Cogn.; no explicit rationale is provided for this transfer. All of the new spp. are illustrated with excellent, composite line drawings, and even C. quinquenervia gets a full description and representative specimen citations. An appendix enumerates all the spp. of Clidemia known to produce formicaria or domatia. Incidentally, all of the relevant taxonomic and nomenclatural innovations in this paper have already been accounted for in the author’s Manual draft treatment of Melastomataceae [see The Cutting Edge 10(3): 3, Jul. 2003], with only the bibliographic details remaining to be added in.

Barfuss, M., M. R. Samuel & W. Till. 2004. Molecular phylogeny in subfamily Tillandsioideae (Bromeliaceae) based on six cpDNA markers: an update. J. Bromeliad Soc. 54: 9–17.

This is a brief report on a preliminary analysis of sequence variation in chloroplast DNA for 108 taxa of Bromeliaceae subfam. Tillandsioideae; “the final analysis will be published at a later date.” The subfamily as a whole is strongly supported as monophyletic. Alcantarea, most of Vriesea, and Werauhia form a monophyletic group; thus, while the recent segregation of Alcantarea and Werauhia [see The Cutting Edge 2(2): 6–7, Apr. 1995] may be defensible, it was not imperative. Tillandsia insignis (Mez) L. B. Sm. & Pittendr. is nested within Werauhia, while Vriesea monstrum (Mez) L. B. Sm. (the only Costa Rican representative of Vriesea s. str. studied) is basal to Werauhia and remote from the clade harboring the type sp. of Vriesea. Likewise removed from the last-mentioned clade are the Andean members of Vriesea sect. Xiphion (E. Morren) E. Morren ex Mez, among which Tillandsia singularis Mez & Wercklé (occurring in Costa Rica) is nested. Mezobromelia is apparently diphyletic, with the Costa Rican M. pleiosticha (Griseb.) Utley & H. Luther nested within Guzmania; the latter taxon is monophyletic (if M. pleiosticha is included), and apparently sister to the Racinaea/Tillandsia clade. Racinaea is moderately supported as monophyletic, but with Tillandsia venusta Mez & Wercklé (occurring in Costa Rica) at its base. Basal resolution of the Racinaea/Tillandsia clade “is lacking,” however “the authors are optimistic that with the addition of nuclear markers and of more taxa these problems can be solved.” Though it seems a bit premature, Tillandsia insignis is formally transferred to Werauhia as W. insignis (Mez) W. Till, Barfuss & M. R. Samuel; perhaps the reintegration of Werauhia (and possibly Alcantarea) into Vriesea will prove the wiser alternative.

Bauer, R. 2003[‘2002’]. The genus Pseudorhipsalis Britton & Rose. Haseltonia 9: 94–120.

Making a strong bid to become the Franco Pupulin of Cactaceae, the author keeps the flow coming with yet another seminal contribution. According to Bauer, “Pseudorhipsalis belongs—with Epiphyllum, Hylocereus, Selenicereus, and Weberocereus—to the Hylocereeae.” Most Pseudorhipsalis spp. have generally been included in Disocactus (oddly omitted from the foregoing list), and the author takes great pains throughout this paper to distinguish the two genera. The relatively tenuous generic distinction is based on the morphology of seedlings and juvenile stems (though these organs are unknown in several Pseudorhipsalis spp.), together with flowering periodicity (Disocactus flowers open at night and remain so for several days, Pseudorhipsalis flowers open in the morning and close the same day). Disocactus sensu Bauer is absent from Costa Rica, being restricted to northern Mesoamerica (Mexico to Nicaragua). Pseudorhipsalis is widespread from Mexico to Peru and Venezuela, Brazil, and the Greater Antilles, but “the main diversity center is Costa Rica, where all six recognized species occur”; alas, the preceding statement is inaccurate, as Pseudorhipsalis alata (Sw.) Britton & Rose “is known only from Jamaica.” The only taxonomic innovations involve the anomalous Pseudorhipsalis amazonica (K. Schum.) Ralf Bauer comb. nov., the only sp. with flowers that are some color other than white (i.e., blue, violet, or magenta); previously, this sp. was included either in Disocactus or Wittiocactus (a substitute for the illegitimate Wittia K. Schum.). Costa Rican material of P. amazonica may be further qualified as belonging to P. a. subsp. panamensis (Britton & Rose) Ralf Bauer comb. & stat. nov., most recently known as Wittiocactus panamensis (Britton & Rose) Rauschert. Bauer accepts Pseudorhipsalis lankesteri (Kimnach) Barthlott, synonymized under P. himantoclada (Rol.-Goss.) Britton & Rose in Manual co-PI Barry Hammel’s draft treatment of Cactaceae, and cites Disocactus horichii Kimnach as a synonym of Pseudorhipsalis acuminata Cufod., rather than P. ramulosa (Salm-Dyck) Barthlott, as according to Barry. Lacks a formal generic heading and description, but features a key to spp. (with Disocactus excluded in the first couplet) and, for infrageneric taxa, synonymy and (for accepted names) typology, full descriptions, comprehensive specimen citations, generous discussions, opulent color photos of living specimens, and SEM micrographs of pollen and seeds. The distribution of the three subspp. of Pseudorhipsalis amazonica is plotted on a colored map. The sparingly illustrated introductory part includes sections on taxonomic history, definition of the genus, distribution, morphology, and flowering period. Tables are employed to document the flowering phenology of all six Pseudorhipsalis spp., and to compare the genus (sans P. amazonica) with Disocactus and the somewhat transitional P. amazonica. N.B.: the foregoing new combinations in Pseudorhipsalis are actually validated in this paper, not (as we incorrectly indicated) in another Bauer paper reviewed in this column in our last issue.

Catalán, P., P. Torrecilla, J. Á. López Rodríguez & R. G. Olmstead. 2004. Phylogeny of the festucoid grasses of subtribe Loliinae and allies (Poeae, Pooideae) inferred from ITS and trnL–F sequences. Molec. Phylogenet. Evol. 31: 517–541.

Okay, here we have it again: Festuca is paraphyletic with respect to Lolium and a polyphyletic Vulpia (as well as numerous other traditional genera), with past hybridization suspected as a complicating factor. There seems precious little to distinguish this offering from another recent paper by the same group (see this column under Torrecilla et al.,in our last issue).

Cleef, A. M. & M. Kappelle. 2003. Memorias acerca de una científica en el páramo costarricense: Adelaida Chaverri-Polini (21 de mayo de 1947–20 de setiembre del 2003). Brenesia 59-60: 3–5.

This eulogy, by two of her colleagues and co-authors, contains first-hand recollections of field experiences and other interactions with recently deceased Costa Rican phytosociologist Adelaida Chaverri. A photo depicts Adelaida in her beloved páramo.

Dressler, R. L. 2004. A key to the Central American species of Encyclia. Orchid Digest 68: 88–92.

The subject of this contribution is Encyclia (Orchidaceae) s. str., exclusive of Prosthechea and (it would appear) Oestlundia. In all, twenty spp. are included. The key is dichotomous, indented, and annotated to indicate geographic distribution by country. We are relieved to see that there are no major surprises here (e.g., spp. attributed to Costa Rica that were not included in the author’s Manual treatment). The recently described Encyclia davidhuntii Withner & M. Fuente, mentioned in the Manual generic discussion on the basis of an unvouchered report from Costa Rica, is synonymized under E. peraltensis (Ames) Dressler. Among the brief “Notes” that conclude the paper, the possibility is raised that E. peraltensis may prove to be simply a small-flowered form of Encyclia amanda Ames) Dressler (widespread in Central America, but not known from Costa Rica). Flowers of several Encyclia spp. are illustrated in a “centerfold” featuring color photos from life by the author’s wife, Kerry A. Dressler.

Faisthuber, W. 2004. Oerstedella centradenia Rchb. f. und Oerstedella centropetala Rchb. f. [sic] in Costa Rica—Synonyme oder zwei verschiedene Arten? Orchidee (Hamburg) 55: 60–61.

The two names mentioned in the title have been combined, under Oerstedella centropetala (Rchb. f.) Rchb. f., by various authors, including Manual contributor (and genus monographer) Eric Hágsater (AMO). Based on his studies in the Monteverde region of Costa Rica, the author contends that Reichenbach’s names do indeed correspond to two distinct entities, separated locally by habitat type (O. centradenia on the seasonally dryer portions of the Pacific slope, O. centropetala in the relatively wet cloud forests) and details of the labellum. No herbarium vouchers are cited, but both supposed spp. are depicted in color photos of living plants.

Gibbs, P. & J. Semir. 2003. A taxonomic revision of the genus Ceiba Mill. (Bombacaceae). Anales Jard. Bot. Madrid 60: 259–300.

Ceiba (including Chorisia), largely restricted to the New World tropics, is here regarded as comprising 17 spp., of which seven are treated within the “Ceiba insignis aggregate species.” Just two spp. (neither in the C. insignis group) are attributed to Costa Rica: Ceiba aesculifolia (Kunth) Britten & Baker f. and C. pentandra (L.) Gaertn. The latter is widespread in the Neotropics, and is the only member of the genus to occur in the Old World, where it is “probably native, at least in part” in West Africa, but “almost certainly introduced in Asia and the Pacific.” Surprisingly, just one Costa Rican specimen of C. pentandra was studied, though the sp. is common in lowland regions throughout the country. Ceiba aesculifolia, ranging from northern Mexico to northwestern Costa Rica, entrains the only mild surprise for us: most material (including all that from Costa Rica) is consigned to the autonymic subsp. with the recognition of C. a. subsp. parvifolia (Rose) P. E. Gibbs & Semir comb. & stat. nov., a smaller-leaved entity from south-central Mexico. Features synonymy and typology at all levels, a detailed genus description, keys to spp. and (where appropriate) nested keys to subspp., sp. and subsp. descriptions, sometimes extensive discussions, comprehensive specimen citations, distribution maps, excellent composite line drawings, indices to exsiccatae and scientific names, and a section on “Excluded or doubtful names.” One of the excluded names, Ceiba rosea (Seem.) K. Schum., applies to a third Costa Rican sp., denoted in our data-base as “Spirotheca rosea (Seem.) P. E. Gibbs comb. ined.” for at least the past 15 years; exasperatingly, the authors once again perpetuate the eternal “to be transferred to the genus Spirotheca,” inexplicably squandering a golden opportunity to do just that. The rather brief introductory part includes discussions of taxonomic history, materials and methods, morphology, cytology, reproductive biology, and habitat. Citing “some special problems” in the taxonomy of Ceiba (e.g., large forest trees, usually flowering when leafless, often at night, with fleshy flowers exhibiting subtle color variations), the authors doubt that their treatment “will represent the last word on these taxa.”

Henderson, A. J. 2004. A multivariate analysis of Hyospathe (Palmae). Amer. J. Bot. 91: 953–965.

The smallish genus Hyospathe, present (though rarely encountered) in Costa Rica, has proven unusually refractory to taxonomic analysis. Although as many as 18 spp. have been recognized, the most recent revision, by Skov and Balslev (Nordic J. Bot. 9: 189–202. 1989), accepted just two. We followed the last-mentioned authors for the purposes of the Manual Arecaceae treatment, but noted that H. elegans Mart., the sp. occurring in Costa Rica, appeared highly and suspiciously variable in other parts of its geographic range (particularly Panama). In the present paper, NY palm specialist Andrew Henderson applies multivariate statistical analysis and the Phylogenetic Species Concept to delimit a total of six Hyospathe spp., two of which (neither represented in Costa Rica) are described as new. Hyospathe elegans remains the correct sp. name for virtually all Central American material (with H. pittieri Burret “just reaching Panama”); however, Henderson now distinguishes six subspp. of H. elegans, including three described as new and two based on new combinations. All the Costa Rican material is referred to the endemic H. elegans subsp. costaricensis Andrew Hend. subsp. nov., while three of the remaining subspp. are endemic (or very nearly so) to Panama. The bulk of the paper is devoted to the character analyses; the taxonomic part is synoptic in nature, with synonymy and typology at all levels, keys to spp. and subspp., and quantitative descriptions of all infrageneric taxa, but lacking a formal genus description and (except for types) specimen citations.

Kriebel, R. 2004. Two new species and one new name in the Gesneriaceae from Costa Rica. Rhodora 106: 43–51.

All three novelties dealt with in this paper involve the genera Nautilocalyx and Paradrymonia, which (as acknowledged by the author) are extremely similar. The two new spp. are Nautilocalyx purpurascens Kriebel and Paradrymonia alata Kriebel, both endemic to Costa Rica, where they occur at low elevations (below 850 m) on the Atlantic slope of the Cordillera de Talamanca. The former sp. is distinguished by its purplish foliage and flowers with two opposite nectary glands, the latter by its large leaves with winged petioles, lance-ovate calyx lobes distinct almost to the base, and large corollas. The new name, Nautilocalyx biserrulatus Kriebel, is occasioned by the generic reassignment of the Costa Rican endemic sp. heretofore known as Paradrymonia bullata Gómez-Laur. & Chavarría, the original epithet being preoccupied in Nautilocalyx by N. bullatus (Lem.) Sprague (1912). The new generic placement is supported by a tabular comparison. A full description and comprehensive specimen citations are provided for N. biserrulatus, and from the latter feature it may be concluded that a recent report of Paradrymonia ommata L. E. Skog from Costa Rica [see The Cutting Edge 9(3): 3, Jul. 2002] is now considered to have been based on a specimen representing P. bullata (i.e., Nautilocalyx biserrulatus). However, this may not be the end of the story: the author suggests that P. ommata may also prove to be a Nautilocalyx, and perhaps even conspecific with N. biserrulatus (in which case, P. ommata is the oldest name), but prudently withholds judgment pending additional collections of P. ommata (currently known only by its Panamanian type). Both new spp. are illustrated in fine composite line drawings by INBio’s Claudia Aragón. This paper marks the published debut (as far as we are aware) of newest INBio curator Ricardo Kriebel [see The Cutting Edge 10(3): 2, Jul. 2003], from whom we can expect to hear a whole lot more…

--, J. González & E. Alfaro. 2004. Symplocos retusa (Symplocaceae), una nueva especie de Costa Rica. Lankesteriana 4: 47–59.

…and a lot sooner than we expected! Symplocos retusa Kriebel, Gonz. Ram. & Alfaro, one of two provisionally named spp. in the recently submitted Manual draft treatment of Symplocaceae [see The Cutting Edge 11(2): 3, Apr. 2004], is based on three collections from the Pacific slope of the Cordillera de Talamanca at ca. 1800–2650 m elevation; though nominally endemic to Costa Rica, one of the stations is Las Alturas de Cotón, very near the Panamanian border. The new sp. is most similar to another Costa Rican endemic, Symplocos tribracteolata Almeda, from which it differs in its entire, apically retuse leaf-blades (vs. serrate or crenate and apically acuminate) and 5-merous, purple flowers (vs. 6-merous and white to pink). Illustrated with a fine composite line drawing by Claudia Aragón.

Incidentally, beginning with this issue, Lankesteriana is adopting the conventional system of issues nested within volumes; previously, each issue was numbered separately, as though a volume unto itself. The present issue is somewhat arbitrarily designated as issue 1 of Volume 4, and a scheme is presented for assigning issue and volume numbers, retroactively, to the previous eight issues. This will undoubtedly be overlooked by most everyone, and ambiguity will forever dog the usage of these early volume numbers (particularly 4–8). For our part, we will endeavor to tow the company line in future Manual installments. Nonetheless, we feel it would have been much smarter to have simply designated this as the inaugural issue of Volume 9.

Lobo C., S. 2003. Colección tipo del Herbario Nacional de Costa Rica (CR). IV. Revisión y actualización de las monocotiledóneas. Brenesia 59-60: 21–34.

This account completes the author’s revision of monocot type material at CR [see The Cutting Edge 10(2): 9, Apr. 2003], supplementing a previous contribution covering the entire group. The present work accounts for 192 type specimens in the families Orchidaceae, Poaceae, Smilacaceae, Triuridaceae, and Zingiberaceae. Among the categories included are holotypes, isotypes, lectotypes, isolectotypes, and neotypes, as well as clonotypes, paratypes, phototypes, and xerotypes. Best represented are isotypes, comprising 40% of the total. All protologues are referenced, and synonymy is occasionally provided.

Luer, C. A. 2004. Systematics of Pleurothallis subgenus Acianthera (Orchidaceae) and three allied subgenera. Pp. 1–114 in, C. A. Luer, Icones pleurothallidinarum XXVI. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 1–265.

Pleurothallus subg. Acianthera (Scheidw.) Luer, one of about 29 subgenera of Pleurothallis in the author’s classification, is characterized by its sessile leaves and terminal inflorescences of solitary or racemosely arranged, more or less fleshy flowers with connate lateral sepals. The author explicitly refutes generic ranking for this taxon, as recently proposed by Pridgeon and Chase [see The Cutting Edge 9(1): 9–10, Jan. 2002] on the basis of their molecular analyses. Its title glosses over the fact that this revision is not comprehensive: “the species with distributions limited to southern South America” are omitted, to be treated in a forthcoming installment. Of the remainder, 94 spp. are here treated in full, including 13 attributed to Costa Rica. This latter contingent includes no additions to or major changes for the Costa Rican flora, as according to Luer’s Manual treatment of Pleurothallis. We note just one minor change: Pleurothallis cingens Ames & C. Schweinf., indicated (questioningly) as a synonym of P. pantasmi Rchb. f. in the Manual, is here listed (sans query) in the synonymy of P. circumplexa Lindl. Includes a key to spp. (unindented) and specimen citations (representative for some spp.). Four new spp. are described, none relevant to us. In a more or less separate section, three new subgenera are described to accommodate spp. previously assigned to subg. Acianthera, but with “floral aberrations inconsistent with the subgenus.” The new subgenera are briefly characterized, and the seven spp. involved are accorded the same formal treatment as those of subg. Acianthera. Of these seven, only Pleurothallis cachensis Ames and P. lepidota L. O. Williams (together comprising subg. Unguella Luer) are definitely known from Costa Rica; however, P. aberrans Luer (of the monospecific subg. Aberrantia Luer) and P. butcheri Luer (one of four spp. in subg. Didactylus Luer) both occur in western Panama and were attributed to Costa Rica by Pupulin [see The Cutting Edge 9(4): 10–11, Oct. 2002]. A sp. key is provided for subg. Didactylus. An index to scientific names, a bibliography, and a set of 114 composite line drawings (four to a page, representing most spp.) serve for all four subgenera. [N.B.: the title given above is our own amalgam of the two alternative versions confusingly presented in the journal, neither of which is entirely satisfactory.]

--. 2004. Systematics of the genus Zootrophion (Orchidaceae). Pp. 211–232 in, C. A. Luer, Icones pleurothallidinarum XXVI. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 1–265.

Including the three new spp. and three new combinations validated in this work, Zootrophion now comprises a total of 18 spp. However, none of the novelties affects the Costa Rican flora, still with the same four Zootrophion spp. featured in Luer’s Manual treatment. The only significant development for us is that we can now provide more exact locality data for Z. williamsii Luer, as follows: “Bosque muy húmedo, pluvial y nuboso, 600–1550m; vert. Carib. y cerca de la División Continental, Cord. Central.” Because the only two Costa Rican specimens of Z. williamsii flowered in cultivation, we remain ignorant of its flowering phenology in the wild. In terms of its basic features and sp. treatments, this revision is essentially identical to the foregoing one.

--. 2004. New genera and combinations in the Pleurothallidinae. Pp. 253–265 in, C. A. Luer, Icones pleurothallidinarum XXVI. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 1–265.

In a staggering and totally unexpected about-face, the author suddenly embraces a “phylogenetic” classification of Pleurothallis s. l., apparently abandoning the venerable “systematic” classification perpetuated in his Manual treatment and contentiously defended, barely two years ago [see The Cutting Edge 10(1): 8, Jan. 2003], against the sweeping, molecular-based changes proposed by Pridgeon and Chase [see The Cutting Edge 9(1): 9–10, Jan. 2002]. In the quest for “monophyletic” genera, Luer now trumps even Pridgeon and Chase (P&C), atomizing Pleurothallis s. l. into at least 27 segregates (we don’t have his entire classification), of which 13 are represented in Costa Rica (as opposed to just six in the system of P&C). Seven names were already available at generic rank, but 16 names reflect changes of status (i.e., elevations from subgeneric or sectional rank); of the latter, 12 are new combinations and four nomina nova. The remaining four generic names (none applicable to Costa Rican spp.) embody new taxa. At the sp. level, 632 new combinations are validated (by our count), more than half of which (364) are in Specklinia, cryptically characterized as “already polyphyletic.” Luer’s criteria for evaluating monophyly are not indicated, but if he was influenced at all by P&C’s cladograms, he certainly has not adhered to them strictly. For example, Anathallis and Phloeophila, distinct genera according to P&C, “blend within Specklinia,” in the view of Luer, who lumps them accordingly. Also, circumstantial evidence suggests that Luer retains Acostaea, rather than including it within Specklinia as per P&C. And our Costa Rican colleagues will be pleased to learn that the genus Brenesia rides again in the system of Luer, as a result of the removal of its type sp. [now Brenesia johnsonii (Ames) Luer] from Acianthera, where it was assigned by P&C, to the genus called Echinosepala (and before that Echinella) by the latter authors.

At first glance, one is tempted to dismiss most or all of the new names in this paper as invalidly published under Art. 34.1 of the Code—i.e., either because they are “not accepted by the author,” or are “merely proposed in anticipation of the future acceptance of the group concerned, or of a particular circumscription, position, or rank of the group.” Indeed, this very volume features the author’s revision of Pleurothallis subg. Acianthera [see two entries back], in which no inkling of the new classification can be found. On the contrary, the following statement appears in the introductory paragraph: “Because of close relationships, the existence of intermediates, and obvious morphological similarities to other infrageneric taxa of Pleurothallis, Acianthera is logically treated as a subgenus…instead of removal from the genus as recently proposed…” However, upon closer inspection, one notes the following parenthetical caveat at the head of the enumeration of new combinations: “This list is added too late to alter the preceding pages.” Thus, one must assume that Luer will adopt his new “phylogenetic” classification in future publications (particularly his impending Flora mesoamericana contributions), and it may be even said that the validity of his new combinations is contingent upon this.

At more immediate risk are the seven new names (four new spp. and three new subgenera) proposed in the aforementioned Acianthera revision, all rechristened at the end of same volume. This situation is addressed by Art. 34.2: “When, on or after 1 January 1953, two or more different names based on the same type are proposed simultaneously for the same taxon by the same author (so-called alternative names), none of them is validly published.” The subgeneric names (elevated to generic rank at the end of the volume) may or may not be saved by the second sentence of Art. 34.2: “This rule does not apply in those cases where the same combination is simultaneously used at different ranks, either for infraspecific taxa within a species or for subdivisions of a genus within a genus.” However, all four sp. names involved (none of which are applicable to Costa Rican taxa) would appear to be invalid, in both Acianthera and Pleurothallis.

Plunkett, G. M., J. Wen & P. P. Lowry II. 2004. Infrafamilial classifications and characters in Araliaceae: insights from the phylogenetic analysis of nuclear (ITS) and plastid (trnL-trnF) sequence data. Pl. Syst. Evol. 245: 1–39.

The significant findings of this study are considerable, but we are most interested in those involving Schefflera. As noted by the authors, “over the past thirty years, the definition of Schefflera has been broadened to include all araliads having once palmately compound leaves, and lacking both articulated pedicels and prickles or arms.” The result has been that numerous genera traditionally regarded as distinct have been absorbed into Schefflera in recent years. Among these, our readers will be most familiar with Didymopanax, but the list also includes many well known horticultural subjects, e.g., Brassaia, Dizygotheca, and Tupidanthus. Accordingly, Schefflera has swollen to ca. 650 spp., i.e., about half the sp. total for the family Araliaceae. The plant-book (1997) went so far as to state that Schefflera is “prob. congeneric with Hedera.” However, in an abrupt turnabout, the present analyses strongly suggest (in agreement with other recent studies) that Schefflera s. l., even without Hedera, is polyphyletic. The 19 Schefflera spp. represented in this study resolve (congruently in all three data sets) into at least five well separated clades, with the neotropical contingent (represented by two unidentified South American spp.) distant from the clade containing the generic type. Moreover, the same five clades (none of which approaches Hedera) are apparent in the preliminary results of an unpublished molecular study (involving two of these same authors) focusing more intensively on Schefflera. The authors aver that “massive taxonomic revisions may be needed to reflect phylogenetic relationships among the species currently treated in Schefflera.” One of the data sets suggests that Dendropanax may also be polyphyletic along New World/Old World lines; however, this indication is highly preliminary, and the generic type [D. arboreus (L.) Decne. & Planch.] is one of our own.

Ramos García-Serrano, C. & J. P. Del Monte. 2004. The use of tropical forest (agroecosystems and wild plant harvesting) as a source of food in the Bribri and Cabecar cultures in the Caribbean coast of Costa Rica. Econ. Bot. 58: 58–71.

An interesting study, by two Spanish workers, in which 84 spp. of food plants (both native and introduced) were identified as utilized by Bribrí and Cabécar populations in Costa Rica. These plants are enumerated in an appendix (with numerous misspellings and some obvious misidentifactions), which also records vernacular names, life zones, parts used for food, and other uses. Vouchers, though allegedly collected and deposited at CR and MAA, are not cited in the appendix or elsewhere. A fine addition to the rather meagre literature on Costa Rican ethnobotany.

Schneider, H., S. J. Russell, C. J. Cox, F. Bakker, S. Henderson, F. Rumsey, J. Barrett, M. Gibby & J. C. Vogel. 2004. Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its implications for biogeography. Syst. Bot. 29: 260–274.

These analyses confirm previous conclusions that Hymenasplenium is the sister taxon of the remaining asplenioid ferns, with all other satellite genera of Asplenium embedded in the latter clade. As noted by the authors, three broad classificatory options are thus tenable: to classify all asplenioid ferns in a single genus, Asplenium; to recognize only Asplenium and Hymenasplenium; or to segregate a variable number of additional genera, according to the constraints imposed by the cladograms. The final option is rejected as likely to create “ill-defined genera,” to require several new generic names, and to conflict with hybridization among different clades. The authors appear to favor the second option (recognition of Hymenasplenium), citing the absence of hybridization with other clades as an important factor. Costa Rican Asplenium spp. referable to Hymenasplenium include A. delitescens (Maxon) L. D. Gómez, A. laetum Sw., A. riparium Liebm., and A. volubile N. Murak. & R. C. Moran [see The Cutting Edge 3(1): 11, Jan. 1996]. Genera commonly recognized as distinct that would be returned to Asplenium include Loxoscaphe and (presumably) Schaffneria, though the latter was not included in this study. Asplenium s. l. exhibits “a separation of the deeper branches into tropical and temperate clades,” at odds with the usual (in ferns) separation into Old and New World clades.

--, A. R. Smith, R. Cranfill, T. J. Hildebrand, C. H. Haufler & T. A. Ranker. 2004. Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Molec. Phylogenet. Evol. 31: 1041–1063.

This landmark contribution analyzes nucleotide sequences from three plastid loci for 98 spp. in the large, important, and taxonomically intractable group of ferns referenced in the title. Four major monophyletic lineages emerge, of which the smallest, including Loxogramme, is sister to the clade comprising the remaining three lineages. The authors consider the possibility of separating a family Loxogrammaceae, but reject it on morphological grounds. Of the remaining three major lineages, two are exclusively paleotropical and thus of no interest to us. The third comprises the vast majority of neotropical Polypodiaceae (with some circumtemperate members), in which are nested the monophyletic, pantropical Grammitidaceae. Because of the latter circumstance, the authors propose to reinstate Polypodiaceae as a more broadly defined family that includes the grammitids. The cladograms presented herein suggest that, within Grammitidaceae, Terpsichore may be polyphyletic; however, very few spp. of Grammitidaceae were studied, in deference to a pending, more intensive study of that group by Ranker.

One of the major conclusions of this paper is that Polypodium, “as traditionally defined,” is “grossly polyphyletic,” with its spp. segregating into at least four major “recognizable” lineages. Polypodium plesiosorum Kunze and P. rhodopleuron Kunze are among the very few Costa Rican spp. belonging to the same clade as the generic type (the Eurasian Polypodium vulgare L.), and thus the only ones certain to remain in the genus Polypodium. Those Polypodium spp. with persistent scales on the fronds, including P. furfuraceum Schltdl. & Cham., P. myriolepis H. Christ, P. polypodioides (L.) Watt, and P. rosei Maxon (all known from Costa Rica), occur polyphyletically in a clade with Dicranoglossum, Neurodium, and Pleopeltis (the last presumably including Marginariopsis); the authors advocate uniting all of these taxa under the generic name Pleopeltis (while explicitly excluding Pseudocolysis, sister to the rest). A third lineage comprises members of the Polypodium subpetiolatum Hook. group (with no Costa Rican representatives mentioned), together with the genus Pecluma, from which separation “is not supported”; Phlebodium, sister to this lineage, is explicitly excluded. The final major lineage of “traditionally defined” neotropical Polypodium spp., appearing in a sister-group relationship with Grammitidaceae, consists of the P. loriceum L. and P. triseriale Sw. sp.-groups, also including (in Costa Rica) such spp. as P. fraxinifolium Jacq., P. levigatum Cav., P. ptilorhizon H. Christ, and P. wiesbaueri Sodiro. Apparently, no alternative name at generic rank is available for this group.

Campyloneurum, Microgramma (including Solanopteris), and Niphidium are all monophyletic and unimpacted; however, Niphidium is sister to the the very similar Campyloneurum, in which it could thus be justifiably submerged. Hyalotrichopteris (often sunk into Campyloneurum) was not studied.

Szlachetko, D. L. 2004. Matériaux pour la révision des Habenariinae (Orchidaceae, Orchidoideae)—5. Richardiana 4: 103–108.

The author’s ongoing fragmentation of Habenaria (for which no cogent rationale is offered) finally has consequences for Costa Rica with the description of his new genus Platantheroides. Of the 34 new combinations here validated, three apply directly to spp. occurring in Costa Rica: Platantheroides alata (Hook.) Szlach., P. eustachya (Rchb. f.) Szlach., and P. strictissima (Rchb. f.) Szlach. The last-mentioned sp. is illustrated with a composite line drawing.

Taylor, C. M. 2004.The neotropical genus Ronabea (Rubiaceae, Lasiantheae). Syst. Geogr. Pl. 74: 35–42.

The deterioration of Psychotria continues with the segregation of a genus of three spp., denominated by Ronabea Aubl., based on morphological as well as molecular criteria. These data suggest that Ronabea is not even a member of tribe Psychotrieae, but is instead more closely related to Lasianthus and allies. Diagnostic characters of Ronabea include triangular, persistent stipules, axillary, bracteate, congested inflorescences, relatively small, white, usually distylous flowers with valvate corolla lobes, bilocular ovaries with solitary, basal ovules, drupaceous, blue-black fruits, unilocular pyrenes with a flat, smooth adaxial surface and a preformed germination lid, relatively large embryos, and soft, oily endosperm. The genus is exclusively neotropical, with two of its three spp. occurring in Costa Rica: Ronabea emetica (L. f.) A. Rich. (formerly Psychotria emetica L. f.) and R. latifolia Aubl. [previously known as Psychotria erecta (Aubl.) Standl. & Steyerm.]. This is a synoptic treatment that includes synonymy and typology at all ranks, a brief generic description, a key to spp., and short distrubution summaries, but lacks sp. descriptions and specimen citations. Ronabea latifolia is depicted in a composite line drawing. Taxonomic history and generic relationships and characteristics are discussed at length in the first half of the paper.

Torke, B. M. 2004. Two new species of Swartzia (Leguminosae) from the Amazon basin of Brazil, with notes on the genus and a key to the unifoliolate species. Syst. Bot. 29: 358–365.

It is the dichotomous “Key to the commonly or strictly unifoliolate species and varieties of Swartzia” that most interests us; we intend to give it a try on the two Swartzia spp. at La Selva that have long been lumped under the name S. simplex (Sw.) Spreng.

Vaughan, D. A. 2003. Appendix I. Revised key to species in the genus Oryza. Pp. 347–351 in, J. S. Nanda & S. D. Sharma (eds.), Monograph on genus Oryza. Science Publishers, Inc., Enfield, NH.

This dichotomous key, annotated with information on distribution, habitat, morphology, and ploidy, identifies the 22 spp. of Oryza (Poaceae) accepted by this author. Rather less conservative is Chapter 3 of this same volume, “Species of genus Oryza and their interrelationships” (S. D. Sharma, pp. 73–111), in which three additional spp. are segregated, including Oryza glumipatula Steud., comprising the New World material of Oryza rufipogon Griff. sensu Vaughan (and the Manual). Vaughan’s key is revised with respect to his 1994 contribution, The wild relatives of rice (Int. Rice Res. Inst., Los Banos, Philippines), which probably ought to have been cited in the Manual.

Yearsley, G. 2004. Josef Ritter von Rawiez Warszewicz (1812–1866). Orchid Rev. 112: 155–159, 171.

Features the standard photographic portrait of Warszewicz, as well as biographical information of general interest; however, much of this article is a litany of Warszewicz’s numerous orchid discoveries throughout the Neotropics, illustrated with the gaudy plant photos that are de rigueur for these orchid journals.


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