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Volume XI, Number 2, April 2004
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Acevedo, H., J. Bustamante, L. Paniagua & R. Chaves. 2002.
Ecosistemas de la cuenca hidrográfica del Río Savegre,
Costa Rica. Edit. INBio.
Here is a rich source of information on one of the wildest
regions of the Costa Rican Pacific slope, the Río Savegre basin,
draining the northern part of the Cordillera de Talamanca. Two of the
11 Costa Rican conservation areas are represented here: Pacífico
Central and La Amistad-Pacífico. This is the first published product
of the ECOMAPAS project, a joint effort of INBio, the Museo Nacional and
SINAC (Sistemas Nacional de Áreas de Conservación) begun in
1998, with support from the government of Spain [see
The Cutting Edge 8(2): 1, Apr. 2001]. That team applied the methods
of Rapid Ecological Assessment to produce a detailed, folded, color map
of the Río Savegre basin, plus a companion volume chock full of
relevant ecological information. Most of the information in the book is
presented under 44 extended descriptions of the different ecosystems
represented in the area, as defined by these workers. Of special interest
to us are the extensive (though unvouchered) lists of characteristic plant
species, separated according to habit type. The botanical exploration of
the Savegre basin represented a major contribution to Costa Rican floristics,
yielding at least one country record, Sebastiania panamensis G. L.
Webster (Euphorbiaceae) [see
The Cutting Edge 8(2): 5, Apr. 2001]. Two additional spp. are claimed
(falsely, in both cases) as endemic to this river basin: Bartlettina
silvicola (B. L. Rob.) R. M. King & H. Rob. (Asteraceae) and Passiflora
gilbertiana J. M. MacDougal (Passifloraceae). Appendices to the book
include a glossary and a list (also unvouchered) of all the vascular plant
spp. (by family) "observadas" throughout the area. Scattered
through the volume are many color photos, maps, and graphs, and at the end
are several pages of drawings of selected plant spp. by INBio’s Francisco
Quesada. This is Vol. 1 of the series "Ecosistemas de Costa
Rica" (see also under Kappelle et al., below).
Acevedo-Rodríguez, P. 2003. Melicocceae (Sapindaceae):
Melicoccus and Talisia. Fl. Neotrop. Monogr. 87: 1-179.
Both genera mentioned in the title, the only New World members
of tribe Melicocceae, are exclusive to the Neotropics. Melicoccus
comprises 10 spp. of Mesoamerica, South America and Hispaniola; however,
the genus is represented in Costa Rica only by the cultivated M.
bijugatus Jacq. (native to northern South America), esteemed for its
green, litchi-like fruits ("mamón"). The much larger
Talisia (52 spp.) is also widely distributed in the Neotropics
(though absent from the Antilles). The author attributes a surprising total
of four Talisia spp. to Costa Rica, compared to just two, T.
allenii Croat and T. nervosa Radlk., accounted for in our database
and in Francisco Morales’s Manual draft treatment of Sapindaceae
The Cutting Edge 7(1): 3-4, Jan. 2000]. The latter sp. is maintained
here, but the name Talisia allenii, formerly applied to a sp. restricted
to southwestern Costa Rica and adjacent Panama, is synonymized under T.
macrophylla (Mart.) Radlk., extending southward to Peru, Venezuela, the
Guianas, and Brazil. Rounding out the Costa Rican complement of Talisia
spp. are T. cerasina (Benth.) Radlk., of the Pacific lowlands from
Parque Nacional Carara to Golfito (thence south to Bolivia and Brazil), and
T. croatii Acev.-Rodr. sp. nov., from near Palmar Norte and the Golfo
Dulce region (to Peru and Brazil). To be expected in southeastern Costa Rica
is the vegetatively distinctive Talisia princeps Oliv., known by a
sterile collection from Prov. Bocas del Toro, Panama. Features a key to
genera (but no formal description of the tribe), full synonymy and descriptions of
genera and lesser taxa, a key to the two subgenera of Talisia, keys to spp.
(nested under the subgenera in Talisia) and subspp., composite line
drawings and distribution maps, and (for each genus) a section on excluded taxa.
The sp. entries each include paragraphs on phenology, distribution and ecology,
common names and uses (occasionally), a generally brief discussion, and a list
of exsiccatae (representative, even for new spp.). Indices to exsiccatae and
local and common names close out the volume. Fourteen new spp. and three new
subspp. are described in this monograph, and four new combinations are validated
(one subgenus, two spp., and one subsp.); except for the aforementioned
Talisia croatii, none of these taxonomic novelties pertains to Costa Rica
(except insofar as all the Costa Rican Talisia spp. become relegated to
the autonymic subgenus).
Atwood, J. T. 2003. Review of the Maxillaria graminifolia
(Kunth) Rchb. f. (Orchidaceae) suballiance. Selbyana 24: 144-164.
This group comprises six spp., ranging from Costa Rica to Bolivia
and Venezuela, with only Maxillaria appendiculoides C. Schweinf. and
M. linearifolia Ames & C. Schweinf. attributed to Costa Rica (the former
being endemic). This represents no net change from the author’s Manual account
of Maxillaria. The "suballiance" is redefined, and all spp.
are thoroughly described and characterized. Also included are a key to spp.
(at the end), full synonymy and typology, comprehensive specimen citations, and
range maps. Each sp. is illustrated with a photograph (of a herbarium specimen)
and a line drawing showing floral details.
Bauer, R. 2003. A synopsis of the tribe Hylocereeae F. Buxb.
Cactaceae Syst. Init. 17: 3-63.
"Intended to be a precursor to a monographic treatment" of
tribe Hylocereeae (Cactaceae), this work admirably fulfills that function.
Useful features include a key to genera, keys to species and infraspecific taxa
for each genus and, for each species and infraspecific taxon, full bibliographic
and typological information, synonymy, and a summary of distribution. Discussions
are provided as necessary, and these are often extensive and insightful. The
author’s generic concepts are in general accord with the Manual draft treatment
by co-PI Barry Hammel, the major exception being that all the Costa Rican
representatives of Disocactus sensu Hammel (i.e., s. l.) are here
segregated in Pseudorhipsalis; Disocactus sensu Bauer (s. str.) occurs
only from Mexico to northern Nicaragua. All of the remaining genera
(Epiphyllum, Hylocereus, Selenicereus and Weberocereus) are
represented in Costa Rica. Many lectotypes, neotypes, and epitypes are designated
in this paper, and many new combinations are validated (frequently involving a
change in status from var. to subsp.), but just one new taxon (which does not
occur in Costa Rica). Significant innovations relevant to the Costa Rican flora
are as follows: the much-used name Epiphyllum phyllanthus is restricted
to South American plants and replaced, in Costa Rica, with E. hookeri
Haw. subsp. pittieri (F. A. C. Weber) Ralf Bauer comb. nov.; Costa Rican
material of E. thomasianum (K. Schum.) Britton & Rose is distinguished as
E. t. subsp. costaricense (F. A. C. Weber) Ralf Bauer comb. nov.;
Mesoamerican material of Pseudorhipsalis amazonica (K. Schum.) Ralf Bauer
comb. nov. is assigned to subsp. panamensis (Britton & Rose) Ralf Bauer
comb. nov., and that of P. ramulosa (Salm-Dyck) Barthlott to the autonymic
subsp.; Selenicereus wercklei (F. A. C. Weber) Britton & Rose falls into
synonymy under S. inermis (Otto) Britton & Rose; and Weberocereus
biolleyi (F. A. C. Weber) Britton & Rose is demoted as W. tunilla
(F. A. C. Weber) Britton & Rose subsp. biolleyi (F. A. C. Weber) Ralf
Bauer, with W. panamensis Britton & Rose as a synonym. The peculiar
myrmecophile lately known as Selenicereus testudo (Karw. ex Zucc.) Buxb.
is reassigned to the genus Strophocactus-apparently not even a member
of tribe Hylocereeae-with the combination Strophocactus testudo (Karw.
ex Zucc.) Ralf Bauer duly validated. Finally, the mysterious entity known as
Disocactus kimnachii G. D. Rowley or Nopalxochia horichii Kimnach,
based on a Costa Rican type, is dismissed as "probably a hybrid between
E[piphyllum]. crenatum subsp. kimnachii and a plant
of the ‘Heliocereus’ group" (neither of which is known from Costa Rica).
Some of the author’s observations (especially with respect to
Strophocactus and Disocactus kimnachii) suggest (to us) that it
might not be such a bad idea to collapse many of these genera (perhaps
including several entire tribes) back into Cereus. Illustrated with
several composite line drawings, two color photographic plates (including
Barry Hammel’s photo of Weberocereus frohningiorum Ralf Bauer), and
two reproductions of colored drawings.
Chandler, G. T. & G. M. Plunkett. 2004. Evolution in Apiales:
nuclear and chloroplast markers together in (almost) perfect harmony. Bot.
J. Linn. Soc. 144: 123-147.
Phylogenetic analysis of nuclear ribosomal DNA sequences upholds
the distinction of Apiaceae and Araliaceae, virtually in their traditional
senses, except that two tribes (Mackinlayeae and Myodocarpeae) of the latter
family form separate lineages that may or may not merit family ranking. A
complication relevant to Costa Rican floristics involves the ever-problematic
Apiaceae subfam. Hydrocotyloideae, here confirmed as polyphyletic, with
genera scattered among three of the four aforementioned lineages. To address
only those genera occurring in Costa Rica: Hydrocotyle itself is a
solid member of the "core" Araliaceae, while Azorella and '
Spananthe are similarly embedded among the "core" Apiaceae.
However, Centella and Micropleura belong to a third lineage,
the Mackinlaya group, still of uncertain placement; it may warrant
recognition as a distinct family, or perhaps merely as a subfamily of
Christenson, E. A. 2004. The genus Warmingia. Orchid
Rev. 112: 84-88.
Five spp. are tentatively accepted in the obscure genus
Warmingia (Orchidaceae), including W. margaritacea B. Johans.,
a Costa Rican endemic known from a single collection. This is a brief
synopsis, with nice color photos but no key or formal descriptions.
De Queiroz, L. P., R. H. Fortunato & A. M. Giulietti. 2003.
Phylogeny of the Diocleinae (Papilionoideae: Phaseoleae) based on
morphological characters. Pp. 303-324 in, B. B. Klitgaard & A.
Bruneau (eds.), Advances in legume systematics, part 10, Higher
level systematics. Royal Botanic Gardens, Kew.
Parsimony analysis of 82 morphological characters suggests
that both Dioclea and Galactia are polyphyletic. Most of
the Costa Rican representatives of the former genus were included in
the study; of these, only Dioclea virgata (Rich.) Amshoff occurs
in the same clade as the generic type (D. sericea Kunth), and
would thus retain its name in the event of a formal taxonomic split.
The recent merger of the monotypic Cymbosema in Dioclea,
by Manual co-PI Nelson Zamora [see
The Cutting Edge 7(3): 9, Jul. 2000], "had low bootstrap
support" in this study (although the published cladograms suggest
that it is at least a viable alternative). The situation in
Galactia is less clearly resolved. See Maxwell & Taylor (below)
for additional insights on this group.
Drábková, L., J. Kirschner, O. Seberg, G. Petersen &
Č. Vlček. 2003. Phylogeny of the Juncaceae based on
rbcL sequences, with special emphasis on Luzula DC. and
Juncus L. Pl. Syst. Evol. 240: 133-147.
Luzula is monophyletic but, of course, Juncus
is not. What this could portend for the Juncus spp. occurring
in Costa Rica is unclear, because just two (J. bufonius L. and
J. effusus L.) of the six were included in the study; furthermore,
the cladogram is not well resolved in many portions. The authors note
that "if one were to reshape the current taxonomy of the Juncaceae
on the basis of [their cladogram], the taxonomy would undergo radical
changes." Wisely, they leave it at that, in lieu of the usual
Dressler, R. H., W. M. Whitten & N. H. Williams. 2004.
Phylogenetic relationships of Scaphyglottis and related genera
(Laeliinae: Orchidaceae) based on nrDNA ITS sequence data. Brittonia
This is the evidential basis for the merger of Hexisea
and Reichenbachanthus (among other formerly accepted genera) in
Scaphyglottis, a move that was formalized (in anticipation of this
paper), for the purposes of the Manual Orchidaceae treatment, at the
eleventh hour [see
The Cutting Edge 9(2): 4-5, Apr. 2002]. Three new combinations are
validated here, none germane to Costa Rican floristics.
Finot, V. L., P. M. Peterson, R. J. Soreng & F. O. Zuloaga. 2004.
A revision of Trisetum, Peyritschia, and Sphenopholis (Poaceae:
Pooideae: Aveninae) in Mexico and Central America. Ann. Missouri Bot. Gard.
As presaged in this column in our last issue [see
The Cutting Edge 11(1): 11, Jan. 2004], Trisetum has now (rather,
again) been split into smaller genera, of which the first two mentioned in
the title occur in Costa Rica. The Manual Poaceae treatment by Francisco
Morales accounted for five Costa Rican spp. of Trisetum s. l.;
three of these, Trisetum irazuense (Kuntze) Hitchc., T. pringlei
(Scribn. ex Beal) Hitchc., and T. tonduzii Hitchc., remain in that
genus, while the other two are here treated formally under Peyritschia,
in which they aleady have available names: Peyritschia deyeuxioides
(Kunth) Finot [previously Trisetum deyeuxioides (Kunth) Kunth] and
P. pringlei (Scribn.) S. D. Koch (the former T. kochianum Hern.
Torres). The authors provide no phylogenetic rationale for the generic
makeover, and admit that "the systematic relationships of the Mexican
and Central American species of Trisetum are not well understood."
Includes a key to genera, generic descriptions, separated keys to spp.,
synonymy and typology at all levels, comprehensive specimen citations,
composite line drawings of selected spp., and an index to scientific names.
Fuentes-Soriano, S. 2004. A taxonomic revision of Pennellia
(Brassicaceae). Harvard Pap. Bot. 8: 173-202.
Pennellia comprises seven spp., mostly ranging from the
southwestern United States to Guatemala, with one sp. disjunct in Bolivia and
northern Argentina. Pennellia longifolia (Benth.) Rollins, the most
wide-ranging member of the genus, was recently found in Costa Rica [see
The Cutting Edge 5(4): 4, Oct. 1998]; but while Costa Rica is mentioned
in the distribution summaries of both the sp. and genus, no supporting
specimens or references are cited. Includes a key to the four genera of the
"Halimolobine clade," a key to Pennellia spp., synonymy and
descriptions of the genus and all spp., comprehensive specimen citations, and
distribution maps. Most spp. (but not P. longifolia) are illustrated
with composite line drawings. No taxonomic novelties.
Hidalgo, O., T. Garnatje, A. Susanna & J. Mathez. 2004. Phylogeny
of Valerianaceae based on matK and ITS markers, with reference to
matK individual polymorphism. Ann. Bot. (Oxford) 93: 283-293.
The large genus Valeriana appears to be paraphyletic,
"with profound nomenclatural and taxonomic implications"; however,
the authors admit that the "American species are not well represented,
which makes our results somewhat provisional." Indeed, the widespread
Valeriana scandens L. is the only sp. occurring in Costa Rica that was
sampled. While it is removed from the clade that includes the generic type
(Valeriana officinalis L.), V. scandens could yet be retained
in Valeriana (according to these cladograms) were the nested, New
World amphitropical genus Plectritis to be absorbed.
Higgins, W. E., C. van den Berg & W. M. Whitten. 2003. A combined
molecular phylogeny of Encyclia (Orchidaceae) and relationships within
Laeliinae. Selbyana 24: 165-179.
Combined analysis of sequence data from both nuclear and plastid
genes demonstrates that Encyclia s. l. is not monophyletic (though it
could easily be made so with the inclusion of a few mono- or oligospecific
genera, such as Homalopetalum and Meiracyllium). On the other
hand, both Encyclia s. str. and the recently segregated Prosthechea
The Cutting Edge 5(2): 4, Apr. 1998] are strongly supported as monophyletic
clades. As an aside, the phylogenetic distinction of Myrmecophila from
Schomburgkia is upheld.
Hilger, H. H. & N. Diane. 2003. A systematic analysis of
Heliotropiaceae (Boraginales) based on trnL and ITS1 sequence data.
Bot. Jahrb. Syst. 125: 19-51.
The results of this study suggest that (in the understatement of
the year) "traditional generic limits [in Heliotropiaceae] warrant
adjustment" (we’re still groping to understand what became of traditional
familiar limits!). The gist of it is, both Heliotropium and
Tournefortia are polyphyletic, with potentially horrific consequences.
In the worst-case scenario, the name Tournefortia would apparently
cease to be used at all; this is because most of the spp. in the aforementioned
genera (including both generic types) are grouped together in a single, large
clade upon which these authors would bestow generic status under the name
Heliotropium (though why this should be is unclear, because the two
genus names have equal priority, and neither is conserved). The only Costa
Rican Tournefortia to escape this particular holocaust is T.
volubilis L. (and presumably T. maculata Jacq., not included in
the study), isolated in a distinct lineage to which the generic name
Myriopus Small would apply. A fourth lineage, sporting the generic
moniker Euploca Nutt., harbors a fairly sizeable block of
Heliotropium spp., with H. procumbens Mill. being the only
identifiable Costa Rican representative. Pending final resolution of the
relationships within Heliotropium s. l., the authors "advocate
maintaining the genus Tournefortia, which is easily defined,"
while accepting that "under this definition the genus Heliotropium
is paraphyletic." Twenty-two new combinations in the accepted genera
are validated, none pertinent to Costa Rica.
Iwarsson, M. & Y. Harvey. 2003. Monograph of the genus Leonotis
(Pers.) R. Br. (Lamiaceae). Kew Bull. 58: 597-645.
This genus of strikingly orange-flowered herbs and shrubs, best
known for the cultivated ornamental Leonotis leonurus (L.) R. Br., is
here resolved into nine spp. Leonotis is exclusively African, except
for the pantropical, annual weed L. nepetifolia (L.) R. Br. var.
nepetifolia, the only taxon known from Costa Rica (where it is of very
local occurrence and presumably not native). Includes synonymy at all levels,
a genus description, a combined key to spp. and vars., detailed genus
descriptions, selected specimen citations, and distribution maps (though only
African collections of Leonotis nepetifolia are cited and mapped). The
sp. entries provide information on distribution, habitat, conservation, and
(irregularly) local names and notes. The introduction features discussions
of affinities and pollination. Indices to scientific names and exsiccatae are
presented as appendices. Four new taxa (three spp. and one var.) are described
and illustrated, and one new combination (at varietal rank) is validated; none
of these are germane to Costa Rica.
Kadereit, G., T. Borsch, K. Weising & H. Freitag. 2003. Phylogeny of
Amaranthaceae and Chenopodiaceae and the evolution of C4
photosynthesis. Int. J. Pl. Sci. 164: 959-986.
Cladistic analysis of sequence variation of the chlorosplast gene
rbcL provides strong support for the monophyly of Amaranthaceae and
Chenopodiaceae taken together, and also for their sister-group relationship
with the small family Achatocarpaceae. However, the exact relationship between
Amaranthaceae and Chenopodiaceae could not be elucidated, leading the authors
to "follow the traditionally recognized families," at least for the
time being, forestalling their merger as proposed, e.g., by the Angiosperm
Phylogeny Group [see
The Cutting Edge 6(1): 5-6, Jan. 1999]. The evolution of C4
photosynthesis has occurred independently at least 13 times in the two families
(whoever said evolution was parsimonious?).
Kappelle, M., M. Castro, H. Acevedo, L. González & H. Monge. 2003.
Ecosistemas del Área de Conservación Osa (ACOSA)/Ecosystems of the
Osa Conservation Area (ACOSA). Edit. INBio.
This is Vol. 2 of the series "Ecosistemas de Costa Rica,"
with the same genesis and principal features as the first volume (see under
Acevedo et al., above). Unlike its predecessor, this contribution tackles a
single conservation area in its entirety, resolving the Osa Conservation Area
into 38 different ecosystems. It also differs in being fully bilingual
Krapovickas, A. 2003. Revisión de Sida sección
Muticae C. Presl (Malvaceae-Malveae). Bonplandia 12: 123-132.
This section is here regarded as comprising six spp., five of which
(all restricted to northeastern Brazil) are newly integrated. Three of the
latter are newly described. The widespread Sida aggregata C. Presl,
the original member of the section, remains the only Costa Rican representative
(though no Costa Rican material was seen by this author). A synoptic treatment,
including a key to spp., synonymy and typology, specimen citations, and some
illustrations, but omitting descriptions (except for the new spp.).
Landrum, L. R. 2003. A revision of the Psidium salutare
complex (Myrtaceae). Sida 20: 1449-1469.
The Psidium salutare complex, which ranges throughout the
Neotropics, is characterized by a usually subshrubby to shrubby habit, the
ability to resprout from underground stems, leaves with reticulate tertiary
veins and a prominent marginal vein, relatively small flowers, and a preference
for grasslands or frequently burned shrublands. Although "at least 44"
sp. names have been proposed, the author resolves this complex into just two
spp., Psidium laruotteanum Cambess. and P. salutare (Kunth) O.
Berg, the latter with five vars. "that correspond closely with geographic
patterns" (i.e., they meet our main criterion for subspp.). Both spp.
occur in Costa Rica, where P. laruotteanum has traditionally been
called P. savannarum Donn. Sm. (here synonymized); because the latter
has been considered restricted to Costa Rica (P. laruotteanum being
otherwise South American), we lose another endemic. All the Mesoamerican and
Caribbean material of P. salutare belongs to the autonymic var., the
only one of the five occurring outside South America. Features a combined key
to spp. and vars., full synonymy and typology, detailed sp. descriptions and
abbreviated var. descriptions, representative specimen citations, range maps,
and two composite photographic plates of exsiccatae. Four new combinations
(none of which concerns us) are validated at var. rank. Numerous
lectotypifications are proposed.
Levin, R. A., W. L. Wagner, P. C. Hoch, W. J. Hahn, A. Rodríguez,
D. A. Baum, L. Katinas, E. A. Zimmer & K. J. Sytsma. 2004. Paraphyly in
tribe Onagreae: insights into phylogenetic relationships of
Onagraceae based on nuclear and chloroplast sequence data. Syst. Bot. 29:
The only result of significant relevance to Costa Rican floristics
is that Oenothera emerges as paraphyletic with respect to Calylophus,
Gaura and Stenosiphon. The authors betray a mild inclination toward
lumping the last three genera (none of which occur in Costa Rica) into an
inclusive Oenothera, defined by "a unique stigma with four
non-commissural lobes." However, if the splitting option were to be
pursued, those Oenothera spp. in clades remote from the one containing
the generic type (O. biennis L.) could be bounced from the genus. At
least two spp. occurring in Costa Rica, Oenothera rosea L’Hér. ex Aiton
and O. tetraptera Cav., are vulnerable.
Li, J. & J. G. Conran. 2003. Phylogenetic relationships in
Magnoliaceae subfam. Magnolioideae: a morphological cladistic analysis.
Pl. Syst. Evol. 242: 33-47.
This rather weak study pretends to support previous suggestions
that Magnolia be circumscribed as a large, diverse genus (i.e., to
include Talauma and other satellites); however, very few spp. were
included, and the jury should still be considered out on this issue.
Liede, S. & U. Meve. 2004. Revision of Metastelma
(Apocynaceae-Asclepiadoideae) in southwestern North America and Central
America. Ann. Missouri Bot. Gard. 91: 31-86.
Metastelma, with "probably around 70" spp.,
is distributed throughout the Neotropics, but most diverse (with "ca.
30" spp.) in the Caribbean region. Notwithstanding "the
unfortunate inclusion of almost every small American asclepiad with twining
habit and pendulous pollinia in Cynanchum by Woodson," the
distinctness of Metastelma and other genera (with Cynanchum s.
str. largely confined to the Old World) has been confirmed in prior molecular
work involving the first author [see, e.g.,
The Cutting Edge 10(1): 7, Jan. 2003]. Of the 16 Metastelma spp.
recognized from the study region, just two occur in Costa Rica: M.
barbigerum Scheele and M. pedunculare Decne. The only Costa Rica
collection (from near Cañas) of the former sp. here cited is the type of
Cynanchum liesneriana L. O. Williams, the basionym of Metastelma
barbigerum var. liesneriana (L. O. Williams) Liede & Meve comb.
nov., otherwise known only from southern Mexico, Belize and Guatemala (two
other vars. range from southern Texas to southern Mexico). Metastelma
pedunculare, recorded from Guatemala, Nicaragua, and scattered localities
mostly on the Pacific slope in Costa Rica (Monteverde, Valle Central, N
Cordillera de Talamanca), has previously been called Cynanchum sepicola
(Pittier) L. O. Williams, C. woodsonianum L. O. Williams,
Metastelma sepicola Pittier (all now regarded as synonyms), and (in
Flora de Nicaragua) "Metastelma sp. B." We assume
(but cannot be sure) that our database records of a third sp. in Costa Rica,
Metastelma schlechtendalii Decne. [Cynanchum schlechtendalii
(Decne.) Standl. & Steyerm.], correctly pertain to M. barbigerum
(M. schlechtendalii ranging from southern Mexico to Honduras, according
to this revision). Features a genus description, key to spp. and infraspecific
taxa, lengthy sp. descriptions (except where infraspecific taxa are recognized,
when only those are described), synonymy and typology at all levels,
comprehensive specimen citations, distribution maps, composite line drawings
of most taxa, sections on excluded and insufficiently known names, and indices
to exsiccatae and scientific names. An introductory discussion of morphology
is illustrated with superb SEM micrographs of floral details. Under
"Excluded Names," new generic affiliations are indicated (though
the appropriate new combinations are inexplicably not made) for each entry,
including several spp. reported from Costa Rica (mostly as Cynanchum):
Metastelma filisepalum Standl. [C. filisepalum (Standl.) L. O.
Williams], referred to Scyphostelma Baill., and the arguably distinct
M. glaberrimum Woodson [C. glaberrimum (Woodson) L. O. Williams]
and M. sepium (Decne.) W. D. Stevens [C. sepium (Decne.) Standl.],
both members of Orthosia Kunth (the two concepts were combined as
"Orthosia sp. A" in Flora de Nicaragua).
Luckow, M., J. T. Miller, D. J. Murphy & T. Livschultz. 2003.
A phylogenetic analysis of the Mimosoideae (Leguminosae) based on
chloroplast DNA sequence data. Pp. 197-220 in, B. B. Klitgaard &
A. Bruneau (eds.), Advances in legume systematics, part 10, Higher
level systematics. Royal Botanic Gardens, Kew.
Yet another nail in the coffin for the venerable Acacia
s. l., here resoundingly affirmed as non-monophyletic, in general accord
with previous studies. Acacia s. str. (i.e., the A.
farnesiana sp. group plus the ant acacias) is strongly supported as
monophyletic, and is well removed from the other members of Acacia
s. l. Also well supported as monophyletic is the Australasian Acacia
subg. Phyllodineae (DC.) Ser. (Racosperma). However,
Acacia subg. Aculeiferum Vassal (Senegalia), comprising
many scandent spp., here emerges as paraphyletic. In conclusion, the authors
implicitly support generic recognition for three Acacia s. l. lineages
(Acacia s. str., Acacia subg. Phyllodineae, and
"the core" of Acacia subg. Aculeiferum), but stop
short of embracing two other generic segregates promoted by Maslin et al.
The Cutting Edge 10(2): 9-10, Apr. 2003], with the proviso that
"denser sampling of both taxa and characters will be necessary to
further resolve" relationships in the group. Incidentally,
Piptadenia "is still not monophyletic as currently
circumscribed," while the separation of Calliandra and
Zapoteca appears beyond reproach.
Maas, P. J. M., L. Y. T. Westra, L. W. Chatrou & collaborators.
2003. Duguetia (Annonaceae). Fl. Neotrop. Monogr. 88: 1-276.
Duguetia is overwhelmingly neotropical, with just four
West African spp. among its 93 total. The African spp. (formerly segregated
under the name Pachypodanthium) are also treated here, but in a
separate section. In the Neotropics, Duguetia is the third largest
genus of Annonaceae (after Guatteria and Annona). However,
it is largely South American, with just two spp. represented in Costa Rica:
Duguetia confusa Maas, in very wet lowland forests on both slopes
and ranging from southeastern Nicaragua to Colombia; and D. panamensis
Standl., restricted to the Atlantic lowlands from southeastern Nicaragua to
northeastern Panama. Features full synonymy and lengthy descriptions at both
genus and sp. ranks, separate dichotomous keys for the neotropical and African
spp., a synoptical key to the neotropical spp. and (for each sp.) a distribution
summary, a list of specimens examined (sometimes representative), an occasional
section on local names and/or uses, and a discussion. Selected spp. (D.
confusa, but not D. panamensis) are illustrated with composite line
drawings. The well illustrated introductory parts (presented as more or less
independent articles mostly authored by the legion of "collaborators")
deal with a wide range of topics including taxonomic history, morphology,
various aspects of anatomy, karyology, palynology, chemistry, floral biology,
dispersal, distribution, phylogeny, uses, and conservation. Distribution maps
are grouped at the end of the volume, together with sections on insufficiently
known and excluded spp., indices to exsiccatae (with African collectors listed
separately) and local and scientific names, and a list of vouchers (e.g., for
the original anatomical, karylogical, and palynological work presented in the
opening chapters). No taxonomic novelties are proposed in this monograph.
Maxwell, R. H. & D. W. Taylor. 2003. Phylogenetic relationships
of the Diocleinae with particular emphasis on the subgroups of Dioclea.
Pp. 325-353 in, B. B. Klitgaard & A. Bruneau (eds.), Advances in
legume systematics, part 10, Higher level systematics. Royal Botanic
Despite using different outgroups and characters, this cladistic
analysis of mainly morphological characters largely supports an independent
study by a different laboratory (see de Queiroz et al., above): to wit, both
Dioclea and Galactia are polyphyletic. However, in the present
study, the former genus does not resolve quite so nicely into two discrete
groups. Dioclea guianensis Benth., occurring in Costa Rica (and not
studied by de Queiroz et al.), is sister to D. virgata (Rich.) Amshoff
in a clade near to the generic type. From the published cladogram, the merger
of Cymbosema in Dioclea would appear to be at least potentially
tenable, although the authors oppose this on practical grounds.
Moya Roque, R. 2004. Gmelina arborea en Costa Rica.
Bois Forêts Trop. 279(1): 47-57.
Presents details of culture and use in Costa Rica of Gmelina
arborea Roxb. ex Sm. (Verbenaceae), an Asian tree introduced in 1975
which has become "the most commonly used species for timber
production." All sentient beings will recoil at the author’s egregious
misappropriation of the term "reforestación" in reference to
monoculture plantations of this exotic plant (and some may object to the
description of its punky wood as "timber").
Nicolson, D. H. & J. H. Wiersema. 2004. (1618) Proposal to conserve
Sesamum indicum against Sesamum orientale (Pedaliaceae). Taxon
The two names referenced in the title, both validated by Linnaeus
in 1753, have long been regarded as synonyms for commercial sesame, sparingly
cultivated and sometimes escaping in Costa Rica. This is one of those
intractable cases in which the accepted name keeps changing as workers
discover ever older instances of union under a one name or the other. The
present authors propose to put an end to this cycle by conserving the name
that is currently most used (as supported by hard evidence from the Internet).
Pedley, L. 2003. A synopsis of Racosperma C. Mart.
(Leguminosae: Mimosoideae). Austrobaileya 6: 445-496.
A recent proposal to conserve the generic name Acacia so
as to apply to its largest (and overwhelmingly Australian) block of spp.
argued (among other things) that failure to conserve would require over 800
new combinations in Racosperma, the correct genus name for the group
The Cutting Edge 10(4): 10-11, Oct. 2003]. Now, in one swell foop, that
concern is neutralized with the validation of 834 new combinations (at sp.
and infraspecific ranks) in Racosperma, by the Australian worker who
reinstated the name (and first advocated the fragmentation of Acacia)
in 1986. So even the Australians are not united on this issue! The author
notes that he "delayed publishing this paper for a year" to allow
the conservation proposal to be submitted. Should that proposal fail, the
names Racosperma and Pedley will quickly become very familiar to
botanists down under. Looking at the bigger picture, this paper effectively
demolishes the notion of conservation as an expedient to obviate the hardship
of having to make new combinations; indeed, the reverse is true: even
astronomical numbers of new combinations can be published more quickly and
easily than a conservation proposal can be readied for submission (let alone
Pendry, C. A. 2004. Monograph of Ruprechtia (Polygonaceae).
Syst. Bot. Monogr. 67: 1-113.
Ruprechtia, wide-ranging throughout the Neotropics,
comprises (according to this revision) 37 spp. of dioecious trees, shrubs
and lianas. At most two of these occur in Costa Rica, notably the recently
described R. costaricensis Pendry. The description of the
last-mentioned sp. (see under this column in our last issue) disregarded
the issue of whether the name Ruprechtia costata Meisn., previously
applied to all Costa Rican material, should still be used for any of it.
Frustratingly, Pendry’s monograph fails to enlighten on that score: while
"Costa Rica (Puntarenas)" is included in his distribution summary,
no Costa Rican collections are either cited or mapped. Features synonymy,
nomenclatural and typological documentation, and detailed descriptions at
all levels, as well as range maps and composite line drawings of selected
spp. (mainly, those described by Pendry here or elsewhere). The sp. key is
based on pistillate material, but is supplemented (in the back of the volume)
by five regional keys, each a tandem affair serving for both pistillate
specimens and sterile or staminate specimens. The individual sp. entries
include information on phenology, distribution, and vernacular names,
comprehensive exsiccatae citations, and discussions. The introductory part
contains sections (generally brief) on taxonomic history, morphology, generic
delimitation, infrageneric classification, habitats and distribution,
pollination and dispersal, chromosome numbers, and fossil record; a separately
authored chapter presents a phylogenetic analysis of the genus. Concluding
the volume are a list of excluded names and indices to exsiccatae and
scientific names. Two new spp. are described, neither relevant to Costa Rica.
Plowman, T. & N. Hensold. 2004. Names, types, and distribution of
neotropical species of Erythroxylum (Erythroxylaceae). Brittonia
This comprehensive nomenclator, appearing 15 years after the first
author’s untimely death, will be the starting point for any future systematic
work on Erythroxylum. Bibliographic and typological information are
provided for 358 names published through the year 2000, including 187 accepted
spp., eight accepted vars., 27 infraspecific names of uncertain status, and
136 synonyms. For accepted names, a distribution summary is given, often
together with miscellaneous commentary. Reference is made to a URL at which
a searchable specimen database and type-image archive may be accessed.
Powell, E. A. & K. A. Kron. 2003. Molecular systematics of the
northern Andean blueberries (Vaccinieae, Vaccinioideae, Ericaceae). Int.
J. Pl. Sci. 164: 987-995.
Phylogenetic analysis of sequence data from four different genes
shows that six of the 10 genera for which more than one sp. were sampled are
not monophyletic, casting serious doubt on the morphological characters
traditionally used to delimit them. Thibaudia is most seriously
polyphyletic, with spp. occurring in four of the seven major clades identified
in this study. Disterigma and Satyria are also polyphyletic
(with D. trimerum Wilbur & Luteyn excluded a priori, and used as an
outgroup), while Macleania and Psammisia are paraphyletic.
Anthopterus, Cavendishia, Sphyrospermum and Themistoclesia are
all very strongly supported as monophyletic. Classificatory options are
scarcely addressed, pending the usual "increased taxon sampling";
interestingly, however, the Central American Satyria spp. (including
the generic type) "form a strongly supported monophyletic group,"
well characterized morphologically, and sister to Thibaudia
costaricensis Hoerold, the only Central American member of its genus.
Pupulin, F. 2003. Orchideenflora Mittelamerika-Ergänzungen
(Teil 1)/Orchid flora of Central America-Supplements (Part 1)/Additamenta
ad orchideologiam mesoamericanam (pars 1). Orchidee (Hamburg) 54: 467-477.
And what would an issue of The Cutting Edge be without a handful
of new Costa Rican orchid spp. from our pal Franco Pupulin? This is
a peculiar paper in that the abstract ("Summary") describes even
the results of part 2, not appearing in this issue; for example, three new
spp. names are mentioned in the abstract, but just two are addressed in this
paper. Both of the latter are Lepanthes spp.: L. casasae
Pupulin, from Monteverde, is compared to L. ferrelliae Luer, while
L. schugii Pupulin, from near Santa María de Dota, is compared
to L. edwardsii Ames. With these additions (both illustrated with
color photos from life, as well as composite line drawings), Costa Rica now
boasts 95 spp. of Lepanthes, the fourth largest Orchidaceae genus in
the country. New distributional data are provided for four additional orchid
spp. (of the nine enumerated in the abstract). Two of these have been
considered Panamanian endemics, and apparently qualify as country records
for Costa Rica (based on the Manual Orchidaceae account): Pleurothallis
butcheri L. O. Williams [here as Acianthera butcheri (L. O.
Williams) Pridgeon & M. W. Chase], from Fila Tinamastes in the northern Fila
Costeña, and Chondrorhyncha crassa Dressler, allegedly from along
the Río Pacuare in Prov. Cartago; however, the former is vouchered only
by a spirit collection at the "Gaia Botanical Garden" [the same
specimen cited in Pupulin’s checklist; see
The Cutting Edge 9(4): 10-11, Oct. 2002], and the latter by a specimen
(also in liquid) prepared from a cultivated individual. Several collections
(all in spirit) from the Tapantí region, plus one from western Panama,
voucher the natural occurrence of Chondrorhyncha lankesteriana Pupulin
(treated in the Manual, but then known only from cultivated material).
Finally, Kefersteinia orbicularis Pupulin, initially known only from
Cerro Nara, is reported from several other localities in the southern Pacific
region (but these were already accounted for in the Manual). Chondrorhyncha
crassa is redescribed based on the Costa Rican material, and illustrated
with a typically superb composite line drawing and a color photo from life.
Also depicted in color photos are C. lankesteriana and Kefersteinia
--. 2003. Orchideenflora Mittelamerika-Ergänzungen (Teil 2)
/Orchid flora of Central America-Supplements (Part 2)/Additamenta ad
orchideologiam mesoamericanam (pars 2). Orchidee (Hamburg) 54: 561-568.
Here we have the second (and presumably final) installment of this
series, with the third new sp., Maxillaria atwoodiana Pupulin, duly
described. Four collections are cited, but just two have locality data,
documenting their occurrence at 200-400 m elevation in the basin of the
Río Savegre, on the Pacific slope of the northern Cordillera de
Talamanca. The new sp. is compared with both Maxillaria confusa
Ames & C. Schweinf. and M. ramonensis Schltr., and is envisioned as
a counterpart of the latter in relatively xeric habitats. Illustrated with
a fine composite line drawing and a close-up color photo from life. The
three new orchid spp. described in this set of papers raise our running
total of new orchid spp. described since the inception of this newsletter
to 209. The present contribution cites other specimens resulting in
the loss of two Costa Rican endemics, Macroclinium alleniorum
Dressler & Pupulin and M. robustum Pupulin & D. E. Mora, both of
which can now be reported from Panama. However, the converse situation is
presented by Stellilabium fortunae Dressler, a former Panamanian
endemic recently collected in the Monteverde region by Gabriel Barboza
(let’s get his name right!). Both of the Macroclinium spp. are
depicted in color photos from life (and M. robustum also by a
detailed floral drawing). Costa Rica now boasts 108 Maxillaria spp.
(unless we’ve missed something) and 14 spp. of Stellilabium (three
more than were treated in the Manual!). Both this and the first part of
this set are fully bilingual (German/English).
Sánchez-Baracaldo, P. 2004. Phylogenetics and biogeography
of the neotropical fern genera Jamesonia and Eriosorus
(Pteridaceae). Amer. J. Bot. 91: 274-284.
Though familiar and superficially distinctive, the two genera
mentioned in the title are known to hybridize, and are weakly separated by
details of frond architecture that appear (at least in Costa Rica) to be
more or less correlated with elevation. We have long suspected that they
should be lumped (Jamesonia being the older name), and now that
suspicion is supported by solid evidence: cladistic analysis of DNA
sequence data reveals that neither genus is monophyletic. However,
"both genera taken together form a monophyletic group,"
comprising a paraphyletic Eriosorus, from which the polyphyletic
Jamesonia has arisen in several independent lineages. The author
makes no explicit recommendations for taxonomic modifications.
Spooner, D. M., R. G. van den Berg, A. Rodríguez, J. Bamberg,
R. J. Hijmans & S. I. Lara Cabrera. 2004. Wild potatoes
(Solanum section Petota; Solanaceae) of North and Central
America. Syst. Bot. Monogr. 68: 1-209.
Solanum sect. Petota Dumort., to which the
cultivated potato (S. tuberosum L.) belongs, comprises about
200 spp. ranging from the southwestern United States to Chile and
Argentina. South America houses a large majority of the spp., with just
25 (plus four nothospp.) accepted in this contribution for North and
Central America. There the group is best represented in Mexico and
northern Central America, with but a single member, Solanum
longiconicum Bitter, present in Costa Rica
The Cutting Edge 9(2): 10-11, Apr. 2002]; however, it is not
endemic, occurring also in western Panama. Includes full synonymy for
the section and all spp., a genus description and very detailed
sectional description, and a dichotomous key to wild spp. in the study
region, together with a synoptic list of characters. Each sp. entry
presents information on phenology, distribution, a comprehensive list
of exsiccatae (shunted to an appendix if excessively long), and a
discussion. Comprehensive line drawings and distribution maps are
provided for all spp. The introductory part covers various topics
including taxonomic history, exploration, morphology, breeding systems,
interspecific hybridization, relationships (including molecular
phylogeny), sp. concept (intended to be phylogenetic, as is the
classification), sp. groups (11 are distinguished), habitats and
distribution, and uses. The illustrations are supplemented by nine
photographic plates (mostly color). An account of doubtful and excluded
names and indices to exsiccatae and scientific names conclude the volume.
No new spp. are published herein.
Ståhl, B. & M. Källersjö. 2004. Reinstatement of
Bonellia (Theophrastaceae). Novon 14: 115-118.
Pursuant to cladistic analyses revealing Jacquinia s. l.
as paraphyletic [see
The Cutting Edge 10(4): 8, Oct. 2003], the authors fulfill their promise
to validate new combinations for the (mostly) orange-flowered spp. under
the obscure generic name Bonellia. This move affects the only Costa
Rican member of the group, most recently known as Jacquinia nervosa
C. Presl, henceforth to be called Bonellia nervosa (C. Presl) B.
Ståhl & Källersjö.
Steele, K. P. & M. F. Wojciechowski. 2003. Phylogenetic analyses
of tribes Trifolieae and Vicieae, based on sequences of the plastid gene
matK (Papilionoideae: Leguminosae). Pp. 355-370 in, B. B. Klitgaard &
A. Bruneau (eds.), Advances in legume systematics, part 10, Higher level
systematics. Royal Botanic Gardens, Kew.
The only result here with potential consequences for us concerns
Melilotus, rarely adventive in Costa Rica, which is shown to be nested
Steinmann, V. W. 2003. The submersion of Pedilanthus into
Euphorbia (Euphorbiaceae). Acta Bot. Mex. 65: 45-50.
In this sequel to his own molecular work portraying Euphorbia
s. l. as paraphyletic with respect to Pedilanthus and several other
The Cutting Edge 10(1): 10, Jan. 2003], the author formalizes the absorption
of the latter genus nomenclaturally. Pedilanthus nodiflorus Millsp., the
only sp. in the group indigenous to Costa Rica, becomes Euphorbia personata
(Croizat) V. W. Steinm., based on the synonym Pedilanthus personatus Croizat
[the epithet nodiflora being preoccupied by Euphorbia nodiflora Steud.
(1840)]. The name Euphorbia tithymaloides L. is available for the
cultivated Pedilanthus tithymaloides (L.) Poit. (Linnaeus was way ahead of
Tam, S.-M., P. C. Boyce, T. M. Upson, D. Barabé, A. Bruneau, F. Forest & J.
S. Parker. 2004. Intergeneric and infrafamilial phylogeny of subfamily
Monsteroideae (Araceae) revealed by chloroplast trnL-F sequences. Amer.
J. Bot. 91: 490-498.
From the perspective of Costa Rican floristics, just one incidental
result of this study is noteworthy: in agreement with previous studies,
Lemnaceae is strongly supported as nested within Araceae (although it could be
maintained at family rank were the Australian Gymnostachys and the
north-temperate Lysichiton, Orontium, and Symplocarpus to be
removed from Araceae).
Torrecilla, P., J.-Á. López-Rodríguez & P. Catalán.
2004. Phylogenetic relationships of Vulpia and related genera (Poeae,
Poaceae) based on analysis of ITS and trnL-F sequences. Ann. Missouri
Bot. Gard. 91: 124-158.
This study has succeeded in elucidating a large, well supported
monophyletic group, here dubbed the "FEVRE clade," that comprises
Vulpia, most of Festuca (including the type sp.), and numerous
small genera not represented in Costa Rica. Those few Festuca spp.
that occur outside the FEVRE clade cluster with Lolium, and could
conceivably be treated under that name. Within the FEVRE clade, Festuca
appears to be paraphyletic, while Vulpia is polyphyletic; however, the
authors do not attempt or even allude to a formal reclassification of the
clade, presumably because "the phylogenetic relationships among [its] main
groups still remain obscure" (ostensibly due to "events related to
lineage sorting, hybridization, and polyploidization"). Nonetheless,
taking the combined cladograms at face value, several scenarios may be
envisioned: either Vulpia (together with several other genera) could
be sunk (again) into Festuca, or the whole assemblage could be
fragmented in a variety of ways into smaller genera. Considering only those
spp. occurring in Costa Rica and included in the study (or whose affinity we
have been able to establish), Festuca herrerae Davidse, F. ovina
L., F. talamancensis Davidse, and F. tolucensis Kunth all belong
to the autonymic section (strongly supported as monophyletic) and would thus
probably remain in Festuca under any cirscumstances; F. rubra L.
occurs in a different clade (within the FEVRE group), and could thus be
reclassified. The two Vulpia spp. recorded (adventively) from Costa
Rica occur in well separated clades, with nomenclatural consequences inevitable
for V. bromoides (L.) Gray, but less certain for V. myuros (L.)
C. C. Gmel. (the generic type).
Zamora V., N., Q. Jiménez M. & L. J. Poveda A. 2004.
Árboles de Costa Rica/Trees of Costa Rica. Vol. 3. Edit.
INBio, Sto. Domingo de Heredia, Costa Rica.
Diputado Quírico Jiménez M. has called
our attention to the availability of this recently published volume;
however, we have not yet personally beheld a copy, so are unable to
provide an encapsulation at the present time.