THE CUTTING EDGE

Volume V, Number 2, April, 1998

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NEWS AND NOTES

BUSMAN'S HOLIDAY. This issue of The Cutting Edge will be short on news, partly because Manual co-PI Barry Hammel is not presently among us. Not to worry, though; he carries the torch somewhere in the wild interior of Suriname, where he has been for the past month, collecting in the suspect company of MO curator Ramblin' Joe Evans. Initial communiques suggested that the absence of hispanophones was partially compensated for by the ready availability of Indian cuisine. We all anxiously await the final report, to appear in these pages.

HOW YA GONNA KEEP 'EM DOWN ON THE FARM? Barry is not the only one whose absence has been felt during this past quarter. INBio botanist Quírico Jiménez is in Spain for three months, taking courses toward a Master's degree at the Universidad Internacional de Andalucia, Baeza, Jaén. Meanwhile, Manual co-PI Nelson Zamora was in Colombia from 9--21 March, taking a course on "Molecular Genetics for the Inventory and Characterization of Biodiversity," at the Centro Internacional de Agricultura Tropical (CIAT), Palmira, Depto. Valle del Cauca.

TREATMENTS RECENTLY RECEIVED

AMARANTHACEAE. José González (INB). This family of mostly weedy herbs is represented in Costa Rica by 11 genera, of which 32 spp. are here treated in full. Most diverse is Alternanthera, with 9 spp. Three spp. [Alternanthera costaricensis Kuntze, Iresine arrecta Standl., and Pfaffia costaricensis (Standl.) Borsch] are endemic; two formally treated spp. are known mainly or exclusively in cultivation [Alternanthera bettzichiana (Regel) Voss and Celosia argentea L.]. A few nomenclatural adjustments to William Burger's (1983) Flora costaricensis account (Fieldiana, Bot. n. s., 13: 142--180), but the only noteworthy change is the addition of Iresine nigra Uline & Bray [see The Cutting Edge 1(4): 3, Oct. 1994].

CARYOPHYLLACEAE. José González (INB). Treats 10 genera and 19 spp., although six of the genera (Polycarpon, Sagina, Saponaria, Scleranthus, Silene, Spergula) and 8 spp. are non-indigenous. Stellaria, with 5 spp., is the most diverse genus. Two species, Arenaria quirosii Standl. and Stellaria nubigena Standl., are endemic. Just one cultivated sp., Saponaria officinalis L., is accounted for formally; this is the only significant difference from the Flora costaricensis treatment of the family (Fieldiana, Bot. n. s., 13: 227--247. 1983), which includes three additional exotic genera (Dianthus, Gypsophila, Lychnis) and 7 more spp.

LEAPS AND BOUNDS

ARALIACEAE. A specimen collected by Gerardo Herrera from ca. 600 m elevation near Turrialba, Prov. Cartago, appears to represent the first Costa Rican record for Oreopanax echinops (Schldl. & Cham.) Decne. & Planch., otherwise distributed from Mexico (Jalisco, Michoacán) to Honduras. We owe this report to Alfredo Cascante (CR), whose preliminary determination was verified at MO from a scanned image sent as an e-mail attachment.

COMMELINACEAE. Independent collections from Macacona de Esparza, Prov. Puntarenas, by former parataxonomist Alvaro Fernández and former INBio curatorial assistant Eduardo Lépiz represent the first Costa Rican records for Tinantia parviflora Rohweder. This striking sp., previously known from Mexico to Nicaragua and Colombia, is recognized by its elongate inflorescence of two or three whorls of cymes.

POACEAE. Schizachyrium malacostachyum (J. Presl) Nash, widespread from Mexico to Nicaragua, Colombia, and the Antilles, can now be reported from Costa Rica on the basis of a collection from the Península de Santa Elena, Prov. Guanacaste, by co-PI Nelson Zamora. The identification is by agrostologist Lynn C. Clark (ISC), who was in Costa Rica teaching a course in grass taxonomy at UCR. On the same trip, Nelson also found and vouchered new populations of the rare Melocactus ruestii K. Schum. (Cactaceae) and Acacia riparia Kunth (Fabaceae: Mimosoideae), spp. not previously represented in Costa Rican herbaria.

ROSACEAE. Prunus skutchii I. M. Johnst., ostensibly a Guatemalan endemic, turns out to be widespread in the mountains of Costa Rica, where it has been confused with P. brachybotrya Zucc. (formerly P. annularis Koehne). According to INBio botanist Francisco Morales, P. skutchii is distinguished by its larger, pubescent hypanthium.

SAPINDACEAE. A mystery liana collected in 1985 at the Reserva Biológica Carara by co-PI Mike Grayum was then identified tentatively by MO botanist Roy Gereau as belonging to the South American genus Allosanthus. With the benefit of additional material, INBio's Francisco Morales has now confirmed this generic attribution. Going one step further, Chico has determined that the Carara collections represent Allosanthus trifoliolatus Radlk., the only described sp., known previously only from Ecuador and Peru.

SMILACACEAE. A Francisco Morales collection from 1100--1250 m elevation on Fila Bustamante, Cantón de Acosta, Prov. San José, has been identified by its collector (and author of the Manual Smilacaceae treatment) as Smilax aristolochiifolia Mill., otherwise known only from southern Mexico, Belize, and Guatemala.

GERMANE LITERATURE

A self-described "miscellany" of mainly South American novelties, just one of which is of direct relevance to Costa Rican floristics. Dicella aciculifera W. R. Anderson, the first known Central American representative of its genus, is based on two collections made by parataxonomist Luis Angulo from 220--270 m elevation on the Península de Osa. The new sp. differs from all its congeners in the stiff, needle-like hairs densely covering the seed-bearing portion of the fruit. The generic identity of the Osa material was first discerned by the author from a scanned image sent as an e-mail attachment [see The Cutting Edge 3(3): 3, Jul. 1996]. This neotropical genus of 23 spp. ranges primarily from northern Venezuela to central Bolivia, with a single sp., Cinchona pubescens Vahl, extending north to Costa Rica. So it is business as usual for us, although three new spp. and one new name are published here, and eight names are removed from recent synonymy. A lengthy introductory section features a cladistic analysis (based on morphological characters), many drawings and black-and-white photos, range maps, and a bar graph of altitudinal distribution. The taxonomic section includes a key to spp., some illustrations, treatments of hybrids, dubious names, and excluded names, and indices to exsiccatae and scientific names. The authors marshal both morphological and molecular evidence in a discussion that emphasizes the close alliance of the monotypic, neotropical genera Muntingia (traditionally classed in Elaeocarpaceae or Flacourtiaceae) and Dicraspidia (traditionally in Tiliaceae). Although their arguments do not seem persuasive, the authors christen a new family, Muntingiaceae C. Bayer, M. W. Chase & M. F. Fay, to accommodate the aforementioned genera and (probably) the monotypic, Peruvian Neotessmannia (presently in Tiliaceae). The authors assert that any close relationship of Muntingiaceae with either Elaeocarpaceae or Flacourtiaceae can be excluded on both morphological and molecular grounds, while the new family is only remotely allied with the core Malvales (including Tiliaceae), and is probably more closely related to Cistaceae and Dipterocarpaceae. Though most similar superficially to Tiliaceae, Muntingiaceae differ in their absence of mucilage canals, and many other details. No cladogram is published here. This preliminary, ad hoc analysis (for the Generic Flora of the Southeastern United States) yields little of substance relevant to the few and rare Costa Rican members of this world-wide complex of ca. 225 spp. (for which up to 17 segregate genera have been proposed). From their analysis of chloroplast DNA restriction sites, the authors conclude that Satureja s. str., Micromeria, and Piloblephis must be removed to preserve monophyly. The remainder of Satureja s. l. is best treated as three genera, of which only one occurs in North America (north of Mexico). The correct name for this North American entity is apparently Clinopodium. The authors assert that there is "no reliable distinction between Gardoquia and Clinopodium s. l." Satureja discolor (Kunth) Briq., the only sp. in this complex that is definitely indigenous in Costa Rica, is a Gardoquia, hence a Clinopodium, according to this scenario; however, the epithet discolor is preoccupied in the last-mentioned genus. Satureja brownei (Sw.) Briq., conceivably native in Costa Rica, is here referred explicitly to Clinopodium as C. brownei (Sw.) Kuntze. Nothing can be concluded with regard to the Antillean Satureja viminea L., widely cultivated in Costa Rica. The new sp. belongs to the same group as a putatively undescribed Tachia recently collected in Costa Rica (see The Cutting Edge 3(2): 5, Apr. 1996). Includes a revised key to all spp. in the genus, plus a tabularized compilation of pollen data. Cladistic analysis of rbcL gene sequences suggests that Vittaria is polyphyletic and Antrophyum paraphyletic. New generic circumscriptions, designed to be strictly monophyletic, are elaborated. Vittaria is restricted to an almost exclusively neotropical assemblage of spp. with extremely narrow (< 5 mm) leaves, e.g., V. graminifolia Kaulf. and V. lineata (L.) Sm. The remaining neotropical spp. formerly included in Vittaria (those with wider leaves, e.g., V. stipitata Kunze) are here segregated in a new genus, Radiovittaria (Benedict) E. H. Crane (formerly Vittaria subgen. Radiovittaria Benedict), and the necessary new combinations are made. Antrophyum is restricted to Old World spp.; New World spp. that have sometimes been treated under that name are here apportioned into Polytaenium and Scoliosorus, with the combination Polytaenium chlorosporum (Mickel & Beitel) E. H. Crane here established for a sp. occurring in Costa Rica. Ananthacorus, Anetium, and Hecistopteris are retained according to tradition, but given "clarified" circumscriptions. Features a cladogram, a key to the genera, and a brief description of each. Taking the cladogram at face value, there would seem to be few (if any) more sensible (i.e., less nomenclaturally disruptive) classificatory options than the one proposed here. However, this is a single study involving one gene; more seriously, the author provides scant indication of the degree of support for the various clades, and the cladogram features a tetrachotomy in a critical region. Is this too much, too soon? Citing cladistic evidence of a sister-group relationship, the author concurs with Kirkbride's recent (Brittonia 49: 354­p;379. 1997) merger of Lindenia and Augusta (under the latter name). Cladistic analysis of the data casts the study genera in a sister-group relationship. This bears upon the classification of the small, neotropical family Cochlospermaceae, sometimes submerged in Bixaceae. These authors contend that, if Bixaceae is to include Cochlospermaceae, it must be further enlarged to accommodate the monotypic, Madagascan Diegodendron, usually classed in its own family or in Ochnaceae. They recommend that Bixaceae, Cochlospermaceae, and Diegodendraceae continue to be recognized as three separate families, especially in view of the weak internal support for the clade comprising all three. Pseudabutilon, treated most recently by Fryxell as a subgenus of Abutilon, is here restored at generic rank, though with some species excluded and others newly included. According to the new definition, this is a New World genus of 19 spp., ranging widely from the southwestern United States and the Bahamas to Argentina and Paraguay, but apparently absent from Costa Rica and Panama. However, P. umbellatum (L.) Fryxell (comb. nov.), the most wide-ranging sp., is vouchered for Costa Rica by Liesner 4299 (MO) from Parque Nacional Santa Rosa, determined by Fryxell (as Abutilon umbellatum L.) in 1982. Nothing new here for us, with Muhlenbergia flabellata Mez the only Costa Rican representative of this complex of 15 perennial spp., ranging from Montana to Tiquicia. No novelties here, but a key to spp., generous exsiccatae citations, black-and-white line-drawings of most spp. (but not ours!), and full descriptions. A cladistic analysis of Encyclia plus two outgroup genera, using morphological characters, leads to the ostensibly inescapable conclusion that one of the three currently recognized subgenera of Encyclia, E. subgen. Osmophytum (Lindl.) Dressler, must be elevated to generic status under the forgotten name Prosthechea Knowles & Westc. The published cladogram reveals that other options are available. Nonetheless, all of the indicated new combinations are here perpetrated in the author's name, menacing such familiar binomials as Encyclia brassavolae Rchb. f., E. campylostalix (Rchb. f.) Schltr., E. chacaoensis (Rchb. f.) Dressler & G. E. Pollard, E. cochleata (L.) Lemée, and E. fragrans (Sw.) Lemée. And just as we were getting accustomed to Encyclia as distinct from Epidendrum! Features a brief generic description, and a tabularization of the characters distinguishing Prosthechea, Encyclia s. str., and Dinema. More fun with orchids! Here is the easy recipe: find a reasonably well-marked infrageneric group, promote it to generic rank, and stamp your brand on all of the "necessary" new combinations. No license required. Oncidium sect. Cebolletae is distinctive in several respects, most notably in having terete to fusiform leaves. Several spp. have been attributed to Costa Rica, i.e., Oncidium ascendens Lindl., O. cebolleta (Jacq.) Sw., O. stipitatum Lindl., and O. teres Ames & C. Schweinf. Here, the erstwhile section achieves generic status under the name Stilifolium, and the indicated new combinations are made for all eight names accepted by the author (the combinations to be attributed, apparently, to "Königer & D. Pongratz, in Königer"). This is an easy call: because no phylogenetic argument is put forth, nomenclatural stability is the only important consideration, and these names will continue to be found under Oncidium in our data-base. Fully bilingual (German/English). Features an excellent line-drawing of one sp. Cladistic analyses of rbcL sequence data indicate that hydrophily and unisexuality have both evolved repeatedly, even within Alismatidae. Marine angiosperms (unique to Alismatidae) have evolved in three separate lineages. All "seagrasses" are included within the families Cymodoceaceae, Hydrocharitaceae, Posidoniaceae, Ruppiaceae, and Zosteraceae, all here recognized as distinct. The marine Hydrocharitaceae form a strongly supported clade embedded within freshwater genera; thus, their segregation in a separate family (Thalassiaceae) is not supported. The analyses suggest (though not convincingly) that Najadaceae are nested within Hydrocharitaceae (mirroring other recent studies), and Limnocharitaceae within Alismataceae. A cladistic analysis involving 29 morphological characters and 22 taxa portrays the subtribe as monophyletic, and (among other things) offers additional support for a liberal circumscription of Sechium (i.e., to include Frantzia and Polakowskia). Trichosalpinx was created by Luer in 1983, to accommodate those "species with lepanthiform sheaths customarily treated in Pleurothallis." So circumscribed, this exclusively neotropical genus presently includes 108 spp., grouped into four subgenera (two here established). Four new combinations are here validated for Trichosalpinx, and 35 new species are published. The Costa Rican contingent of 19 spp. (representing two of the four subgenera) includes two novelties, Trichosalpinx fruticosa Luer and T. parsonsii Luer & Dod, the latter believed endemic. Two other spp. are apparently endemic to Costa Rica: Trichosalpinx nana (Ames & C. Schweinf.) Luer and T. navarrensis (Ames) D. E. Mora & J. B. García. Trichosalpinx caudata Luer & R. Escobar, from 500 m elevation in Prov. Chiriquí, Panama, is perhaps to be expected in Costa Rica. Features a key to subgenera and nested keys to spp., full descriptions and exsiccatae citations, illustrations of all spp. (grouped by subgenus), and an index to scientific names. Macropharynx comprises 5 spp. of lianas, ranging from Costa Rica to northern Argentina. The genus is here reported for the first time from Mesoamerica, on the basis of several Costa Rican collections of Macropharynx renteriae A. H. Gentry from 300­p;100 m elevation on the Atlantic slopes of the Cordilleras de Guanacaste, Tilarán, and Central. Previously known only by the type collection from Depto. Chocó, Colombia, M. renteriae is also here reported from coastal Ecuador. One new combination; features a key separating Macropharynx and Asketanthera, and a generic description and sp.-key for the former. No illustrations. Only one item relevant to Costa Rican floristics: the widespread sp. known since 1916 as Iresine calea (Ibáñez) Standl. must now be called by the older name I. latifolia (M. Martens & Galeotti) Hook. f. Turns out that an alleged earlier homonym long accepted as precluding the use of I. latifolia never existed in the first place! This reminds us of the case of the widely used name Saurauia veraguasensis Seem. (Actinidiaceae), a superfluous nomen novum (for Saurauia montana Seem.) similarly prompted by an imaginary earlier homonym. The lesson is that, during the course of routine nomenclatural research, the existence and validity of ostensible "blocking names" should always be confirmed. Miscellaneous gleanings relevant to Costa Rican floristics: the familiar sp. known variously to old La Selva hands (and others) as Siparuna nicaraguensis Hemsl., S. andina Tul., and S. macra Standl. is evidently now to be called Siparuna thecaphora (Poepp. & Endl.) A. DC., under which all of the foregoing names (and numerous others) are to be found in synonymy. More surprisingly, Monimia macrophylla J. F. Morales & Q. Jiménez, described only recently (Novon 6: 395--397. 1996) as a Costa Rica endemic, is here reported from Ecuador on the basis of five collections from both slopes of the Andes. It is also attributed to Panama and Colombia, though no specimens are cited. Regarding the novel family circumscriptions, see the following entry. Phylogenetic analyses of morphological and molecular data, the latter comprising sequences for the rbcL gene and the trnL-trnF spacer region of the chloroplast genome, "reveal" that Siparuna is sister to the monotypic West African Glossocalyx, and that neither genus is closely related to the remaining Monimiaceae. This supports a widely held suspicion that the last-mentioned family is polyphyletic. New floral-anatomical data for Siparuna support a close relationship with Glossocalyx. The family Siparunaceae, as here espoused, includes Siparuna, Glossocalyx, plus the "enigmatic," ditypic, Brazilian Bracteanthus, which was not a part of this study. The authors allow that "reconstruction of phylogenetic relationships within Laurales is still in its initial phase." Gentlea is a sharply defined genus of nine spp., restricted to "upper cloud and elfin forests" from Mexico to Peru. This paper presents an updated generic description and synonymy, a key to all spp., and updated sp.-level synonymy (with nine binomials newly synonymized and three newly excluded). The following four spp. are attributed to Costa Rica: Gentlea standleyi Lundell [including G. minor (Standl.) Lundell nom. illeg. and G. costaricensis Lundell], G. venosissima (Ruiz & Pav.) Lundell (including Ardisia meiantha Donn. Sm.), G. austin-smithii (Lundell) Lundell, and G. micranthera (Donn. Sm.) Lundell. The first-mentioned sp. is endemic. The key to genera, mentioned in the title, is particularly useful. "Detailed" Costa Rican distribution maps are provided for 111 (by our count) spp. of Bromeliaceae. The maps are small and cluttered, with symbols for (2--) 4 spp. per map superimposed on Holdridge Life Zone contours. Data are mainly from Costa Rican herbaria (CR, INB, and USJ). A few general trends emerge: Vriesea (s. l.) and Guzmania are most diverse in the highlands, Aechmea in the humid lowlands. Documents first Costa Rican records for 14 fern spp. in 12 disparate genera, with Flora mesoamericana Vol. 1 (1995) as the point of reference. All 12 novelties occur in Costa Rica, and all but one are endemic. Each is duly assigned to a subsection and/or section, and illustrated with an excellent, detailed line-drawing by Silvia Troyo. Niphidium crassifolium (L.) Lellinger, the most widespread and familiar sp. in this Polypodium segregate, is here split in twain with the recognition of N. oblanceolatum A. Rojas. The new sp., which differs in leaf shape and details of the rhizome scales, ranges from Mexico to Peru at 0--1000 (--1400) m. The residual N. crassifolium is similarly distributed geographically, but at somewhat higher elevations [(500--) 800--1400 (--1800) m]. Three new country records are also reported here; none concerns us directly, but the report of Niphidium nidulare (Rosenst.) Lellinger from the Boquete region of western Panama signifies the loss of another Costa Rican endemic. Nine new fern spp. are described from Costa Rica, in as many genera: Blechnum, Diplazium, Hymenophyllum, Lellingeria, Lophosoria, Megalastrum, Polypodium, Saccoloma, and Terpsichore. Just two of the new spp. overlap into Panama. All are illustrated with black-and-white drawings. This pantropical genus of 43 spp. occurs in tropical Africa, from the Himalayan region to China and southeastern Australia, and in Mesoamerica, with the center of sp. diversity in the Indo-Malesian region. No infraspecific taxa are accepted. The (Indonesian) author accepts but a single New World sp., Alstonia longifolia (A. DC.) Pichon, ranging from south-central Mexico to central Costa Rica. Five spp. are here described as new. Based on a cladistic analysis of 68 morphological characters, five sections are maintained in the genus. The monotypic neotropical sect. Tonduzia is well nested among the Old World sections. Features a lengthy consideration of wood anatomy, regional and synoptical keys to all taxa, full synonymy and generous sp. descriptions, range maps, line drawings of most spp., and an index to exsiccatae (but no conventional specimen citations). N.B.: Francisco Morales (INB), who contributed the Manual Apocynaceae treatment, uses the name Alstonia pittieri (Donn. Sm.) A. H. Gentry for Costa Rican material. According to Chico, there are at least two Mesoamerican spp.: the rarely collected A. longifolia, restricted to Mexico and Guatemala, is distinguished by its much larger flowers. This study raises more interesting questions than it answers definitively. Gesneriaceae tribe Episcieae comprises 16 genera, including Columnea and Drymonia, important components of the Central American epiphyte flora. The analysis provides strong support for Hans Wiehler's division of Episcia into various splinter genera (Alsobia, Nautilocalyx, Neomortonia, Oerstedina, Paradrymonia, and Rufodorsia). Nonetheless, Episcia s. str. emerges as paraphyletic, as it includes Neomortonia nummularia (Hanst.) Wiehler. Neomortonia is diphyletic (with N. rosea Wiehler nested among Columnea s