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The Cutting Edge
Volume VI, Number 2, April, 1999
News and Notes
Leaps and Bounds
This study of sequence variation reveals that the vast genus Psychotria
"is paraphyletic with respect to all the other genera of the tribe."
From the cladogram, we would have said it was polyphyletic. Quite possibly
it is both.
- Andersson, L. & J. H. E. Rova. 1999. The rps16 intron
and the phylogeny of the Rubioideae (Rubiaceae). Pl. Syst.
Evol. 214: 161--186.
The further breakup of Caprifoliaceae (already shorn of Sambucus
and Viburnum) is proposed, mainly (it seems) to obviate the inclusion
therein of Dipsacaceae, Morinaceae, and Valerianaceae. But lumping is always
preferable to splitting (assuming monophyly), and we have no problem with
an enlarged Caprifoliaceae. Better by far than yet more new, little families!
Features a phylogenetic tree, brief family descriptions, and a ± natural
key to the 12 genera of the three "families" involved.
- Backlund, A. & N. Pyck. 1998. Diervillaceae and Linnaeaceae,
two new families of caprifolioids. Taxon 47: 657--661.
This study supports recent molecular evidence suggesting a division of the
traditional Flacourtiaceae into two smaller families: Flacourtiaceae s.
str. (most genera occurring in Costa Rica), and Kiggelariaceae (including
Lindackeria, Carpotroche, and Mayna, to name only those
genera represented in Costa Rica). Stamens develop centrifugally in the
former group, but centripetally, or in "a sequence that is neither
distinctly centripetal nor centrifugal," in the latter.
- Bernhard, A. & P. K. Endress. 1999. Androecial development and
systematics in Flacourtiaceae s. l. Pl. Syst. Evol. 215: 141--155.
An informal analysis and evaluation of several regional lists and accounts
(including one incarnation of the INBio/Manual data-base), accepting 38
indigenous spp. (plus three infraspecific taxa) for Costa Rica. Of special
interest is Apéndice 1, an annotated and nomenclaturally updated
(but unvouchered) checklist of Mesoamerican Cactaceae. Our count reveals
a surprising total of 16 taxa (13 spp. and three infraspecific taxa) flagged
as endemic to Costa Rica. These are: Disocactus kimnachii G. D.
Rowley (syn. Nopalxochia horichii Kimnach), Hylocereus calcaratus
(F. A. C. Weber) Britton & Rose, Pseudorhipsalis acuminata Cufod.,
P. himantoclada (Rol.-Goss.) Britton & Rose, P. horichii
(Kimnach) Barthlott, P. lankesteri (Kimnach) Barthlott, P. ramulosa
(Salm-Dyck) Barthlott var. angustissima (F. A. C. Weber) Barthlott,
Rhipsalis micrantha (Kunth) DC. f. micrantha and f. kirbergii
(Barthlott) Barthlott, Selenicereus wercklei (F. A. C. Weber) Britton
& Rose, Stenocereus aragonii (F. A. C. Weber) Buxb., Weberocereus
bradei (Britton & Rose) G. D. Rowley, W. imitans (Kimnach
& Hutchison) Buxb., W. tonduzii (F. A. C. Weber) G. D. Rowley,
W. trichophorus H. Johnson & Kimnach, and W. tunilla (F.
A. C. Weber) Britton & Rose.
- Bravo-Hollis, H. & S. Arias. 1999. Sinopsis de la familia Cactaceae
en Mesoamérica. Cact. Suc. Mex. 44: 4--19.
Several names attributed to Costa Rica in this appendix are not accounted
for in our data-base: Epiphyllum columbiense (F. A. C. Weber) Dodson
& A. H. Gentry, Hylocereus polyrhizus (F. A. C. Weber) Britton
& Rose, H. undatus (Haw.) Britton & Rose, Opuntia decumbens
Salm-Dyck, Pereskia bleo (Kunth) DC., Weberocereus panamensis
Britton & Rose, and W. tunilla (F. A. C. Weber) Britton &
Rose. The reverse is true for Disocactus biformis (Lindl.) Lindl.,
Opuntia ficus-indica (L.) Mill., and Rhipsalis teres (Vell.)
This thesis chapter, subtitled "Precursor of Flora Neotropica Monograph,"
is effectively published here, according to the conditions of the Code (Arts.
29--31), whatever the author's intent. The genus Malmea is divided
into four smaller, somewhat technical genera (three newly validated), with
no cladistic analysis and no phylogenetic argument of any kind; indeed,
the author implies that Malmea s. l. is monophyletic. It would appear
that the author has simply managed to recognize some synapomorphies of a
few major clades, and has congratulated himself by elevating these to generic
rank. The only rationale presented (or alluded to) for this re-ranking
is the never-ending quest for genera defined by characters of equal strength:
"These groups can be discerned on the basis of several types of morphological
and anatomical characters, many of which are significant for generic
circumscription within Annonaceae" (emphasis ours). The old table-leg
shortening thing. We would argue that the time has passed for this sort
of pointless reshuffling: if a traditional genus (or other taxon) is monophyletic,
or has not been shown to be otherwise, don't mess with it! Simple as that,
a priori. But now, there is the fait accompli factor to be
considered: because half (13 of 26) of the spp. in Chatrou's new genera
are here described as new, undoing his generic reclassification would wreak
as much nomenclatural havoc as adopting it. In view of this, we would have
to tentatively endorse the new classification.
- Chatrou, L. W. 1998. Revision of the Malmea alliance: Malmea
and three new, neotropical genera: Klarobelia, Mosannona,
and Pseudomalmea. Pp. 103--192 in, L. W. Chatrou, Changing
genera: systematic studies in neotropical and West African Annonaceae.
Utrecht Univ., Utrecht, Netherlands.
The consequences for Costa Rican floristics are, in any case, minor. The
endemic Malmea costaricensis R. E. Fr., a rarely collected lowland
sp. of both slopes, becomes Mosannona costaricensis (R. E. Fr.) Chatrou.
Malmea s. str. is restricted to six, mainly South American spp.,
reaching only as far north as central Panama (as noted by the author, "It
is somewhat paradoxical that the oldest and most well-known name, Malmea,
remains associated with the least known genus of this alliance").
Mosannona, with 14 spp., emerges as the largest genus in the "Malmea
alliance." A second Costa Rican sp. in the group, Klarobelia stipitata
Chatrou sp. nov., is endemic to the Golfo Dulce region, and one of just
two members of Klarobelia (a genus of 10 spp.) to extend beyond South
America. This sp. was treated and illustrated in Paul Allen's Rain
Forests of Golfo Dulce (1956) under the misapplied name Guatteria
lucens Standl. Yet a third Costa Rican sp., from the San Carlos region,
has been listed in the INBio/Manual data-base as Malmea hypoglauca
(Standl.) R. E. Fr. [i.e., Mosannona hypoglauca (Standl.) Chatrou];
however, that name was apparently misapplied, and the Costa Rican material
is now regarded by Chatrou (pers. comm.) as representing a new sp. of Mosannona,
still awaiting description.
The taxonomic section of this work is nicely done, featuring a key to genera
and keys to species in each genus, descriptions of genera and species, synonymy
and typology, complete specimen citations, distribution maps, composite
line drawings of selected spp., and an index to exsiccatae. More generous
discussions would have been welcome.
This new edition of Quírico's seminal work spotlights 40 spp. of
valuable timber trees threatened and in danger of extinction in Costa Rica.
Among these, 18 have already been protected by a ban on cutting, including:
Cordia gerascanthus L. (Boraginaceae), Couratari scottmorii
Prance (Lecythidaceae), Guaiacum sanctum L. (Zygophyllaceae), Myroxylon
balsamum (L.) Harms (Fabaceae: Papilionoideae), and Swietenia macrophylla
- Jiménez, Q. 1999. Arboles maderables en peligro de extinción
en Costa Rica. Ed. II (revisada y ampliada). Inst. Nac. de Biodiversidad,
Sto. Domingo de Heredia, Costa Rica.
Angostura is more narrowly defined, as a genus of just seven spp.
(as opposed to 38, fide The plant-book). The spp. here newly excluded
from Angostura are recognized as members of Conchocarpus,
together with some spp. formerly included in other genera such as Galipea
and Ticorea. Conchocarpus, not even included in the most
recent edition of The plant-book, rises like a phoenix as a genus
of 45 spp., strongly centered in eastern Brazil. It seems from the outset
a suspicious taxon, described as "quite variable in most characters
and...composed of the species that do not fit into the other more easily
definable genera." A key is provided to separate Angostura,
Conchocarpus, and Galipea, three of the 26 genera in tribe
Galipeeae Kallunki subtribe Galipeinae Kallunki (both of the suprageneric
names freshly minted to replace their predecessors, based on the illegitimate
Cusparia). Twenty-eight new combinations are here validated, plus
24 new infrageneric taxa (none of the latter from Costa Rica). Angostura
granulosa (Kallunki) Kallunki comb. nov. (formerly in Galipea),
of Nicaragua and Costa Rica, is the only Mesoamerican member of its genus.
The sp. most recently known as A. nicaraguensis (Standl. & L.
O. Williams) T. S. Elias becomes Conchocarpus nicaraguensis (Standl.
& L. O. Williams) Kallunki & Pirani. The only other Conchocarpus
sp. reaching Costa Rica is C. guyanensis (Pulle) Kallunki & Pirani,
formerly known to us as Ticorea unifoliolata T. S. Elias (see our
last issue). Features comprehensive generic descriptions, keys to all spp.
in both genera, synonymy and typology, distribution summaries, composite
line-drawings of selected (new) taxa, some SEM pollen micrographs, sections
on doubtful and excluded species, an index to accepted and rejected names,
and an index to exsiccatae. Six lectotypes are designated, as well as one
epitype. Absent are sp. descriptions and exsiccatae citations (except for
new spp.; these are synopses, after all), and any kind of cladistic or phylogenetic
analysis or discussion; the reclassification appears to have been accomplished
solely on the basis of character association.
- Kallunki, J. A. & J. R. Pirani. 1998. Synopses of Angostura
Roem. & Schult. and Conchocarpus J. C. Mikan (Rutaceae).
Kew Bull. 53: 257--334.
Only the latter sp. need concern us. The name Stromanthe palustris
is here validated for a sp. discovered and recognized as new more than 20
years ago, at the Estación Biológica La Selva. Originally,
it was identified as belonging to the genus Ctenanthe, a close relative
of Stromanthe; indeed, the classification of the new sp. appears
still to be in some doubt. Apart from La Selva, S. palustris is
known only from the Reserva Biológica Hitoy-Cerere, south of Puerto
Limón. Lengthy description, black-and-white photos of live material.
- Kennedy, H. 1999. Calathea singularis and Stromanthe palustris,
two new species of neotropical Marantaceae. Novon 9: 61--65.
On the basis of both morphological and cladistic analysis, Caesalpinia
(Fabaceae: Caesalpinioideae), a genus of ca. 120--130 spp., has recently
been divided by the author and colleagues into various informal "groups."
The Poincianella-Erythrostemon "group," here revised,
is a strictly New World assemblage of 47 spp., ranging from the southern
United States to southern Chile. It is most diverse in drier regions, and
not well-represented in southern Central America. Both Caesalpinia eriostachys
Benth. and C. exostemma DC., the only spp. reaching Costa Rica, are
essentially restricted to the Guanacaste region. Several new taxa and new
combinations are validated, but the only one affecting us is C. exostemma
subsp. tampicoana (Britton & Rose) G. P. Lewis stat. & comb.
nov.; this name, applied to a little-known Mexican taxon, consigns our material
to the autonymic subsp. A beautifully presented piece of work that provides
nearly all the features one might ask of a taxonomic revision, and then
some: a lengthy introductory section, key to spp., thorough descriptions,
comprehensive specimen citations, distribution maps, excellent line-drawings,
indices to scientific names and exsiccatae, and 15, mostly color, photographic
plates, mainly of live plants. There is also a key to the "groups"
of Caesalpinia and a detailed, formal description of the Poincianella-Erythrostemon
"group"; however, one searches in vain for a concise, diagnostic
characterization of the latter.
- Lewis, G. P. 1998. Caesalpinia: a revision of the Poincianella-Erythrostemon
group. The Royal Botanic Gardens, Kew, UK. 233 pp.
Physalis is characterized as a genus of ca. 90 spp., all but one
indigenous to the New World. The spp. are tentatively grouped in four subgenera,
including subg. Rydbergis Hendrych, characterized by rotate, campanulate,
or urceolate, unlobed, mostly maculate corollas. Physalis sect.
Epeteiorhiza G. Don, one of nine sections of subg. Rydbergis,
is defined by a herbaceous habit, glabrous to pubescent or glandular-pubescent
foliage, and solitary flowers with yellow, rotate corollas and terete to
strongly 5-angled fruiting calyces. The 14 spp. of sect. Epeteiorhiza
range from Canada to South America. Six are recorded from Costa Rica (all
wide-ranging, at least in the Mesoamerican region): Physalis cordata
Mill., P. ignota Britton, P. minuta Griggs, P. porrecta
Waterf., P. pruinosa L., and P. pubescens L. Features introductory
sections on taxonomic history and general morphology, descriptions of the
genus and section, keys to the subgenera, to sections of subg. Rydbergis,
and to spp. of sect. Epeteiorhiza, sp. descriptions, representative
specimen citations, range maps, and some line-drawings.
- Martínez, M. 1998. Revision of Physalis section Epeteiorhiza
(Solanaceae). Anales Inst. Biol. Univ. Nac. Autón. México,
Bot. 69: 71--117.
Macoubea mesoamericana J. F. Morales becomes the third sp. of this
mainly Amazonian genus of trees. The only Central American representative,
discovered in Costa Rica just 10 years ago (but long known from Panama),
has heretofore gone under the name M. sprucei (Müll. Arg.) Markgr.
Nonetheless, it differs significantly from the latter in fruit and seed
morphology. The Costa Rican collections are all from lowland forests on
the Atlantic slope of the Cordillera de Talamanca, while all Panamanian
collections have been made along the El Llano--Cartí road. Includes
drawings of fruits and seeds of M. mesoamericana and M. sprucei,
plus a key to all three spp.
- Morales, J. F. 1999. A new species of Macoubea (Apocynaceae)
from Mesoamerica. Novon 9: 86--88.
According to Art. 62.1 of the Code, "A generic name retains the gender
assigned by botanical tradition, irrespective of classical usage or the
author's original usage." Paclt's own evidence makes it clear that
botanical tradition, classical usage, and the original author's usage converge
in assigning feminine gender to this genus. So is it really necessary to
propose formal conservation, simply because a minority of workers have gotten
it wrong? Paclt's argument for conserving the spelling Zizyphus
over Miller's original Ziziphus is also flawed, as noted in Dan
Nicolson's editorial aside. We wonder why Taxon even publishes such
seemingly needless proposals (only to undermine them with editorial postscripts).
Would it help to add an article to the Code fixing spellings and genders
of generic names according to Index nominum genericorum, or some
other standard work?
- Paclt, J. 1999. (1399) Proposal to conserve the name Zizyphus
(Rhamnaceae) with that spelling and with feminine gender. Taxon
The sundering of Scrophulariaceae, in the traditional sense, marches on.
As we were already aware, "the hemiparasitic herbs historically retained
in the Scrophulariaceae are now associated with the wholly parasitic,
achlorophyllous Orobanchaceae Vent., so that this family is now defined
as a taxon of some 60 genera." These include Agalinis, Buchnera,
Castilleja, Euphrasia, Orobanche, and Pedicularis.
Scrophulariaceae s. str. is "now restricted to approximately 24 genera,"
among which are Scrophularia and Verbascum. This leaves nearly
110 genera that do not belong (according to recent molecular/cladistic analyses)
in either of the aforementioned families, but comprise a family of their
own, for which the oldest available name is Veronicaceae Durande (1782).
In addition to Veronica, this family includes Antirrhinum,
Linaria, Lindernia, and Penstemon. End of story, but
for the fact that "there is considerable evidence" that both Callitrichaceae
and Plantaginaceae also belong in Veronicaceae. This would necessitate
the use of the name Plantaginaceae Juss. (1789) for the whole group, because
it is listed in App. IIB of the Code, while the older Veronicaceae is not.
The authors of this proposal, objecting to "the use of [Plantaginaceae]
for a taxon dominated by genera traditionally associated with the Scrophulariaceae,"
at first thought to conserve Veronicaceae, but were thwarted by a provision
of the Code that fixes the starting date for conserved family names at 4
Aug. 1789. Thus the present strategy, to conserve Antirrhinaceae Pers.
(1806) against Plantaginaceae, in the event the two taxa are united (and
if they are not, then Veronicaceae would prevail).
- Reveal, J. L., W. S. Judd & R. Olmstead. 1999. (1405) Proposal
to conserve the name Antirrhinaceae against Plantaginaceae
(Magnoliophyta). Taxon 48: 182.
But let's back up: if the Veronicaceae really should prove to be more closely
related to Plantaginaceae than to Scrophulariaceae, why would the botanical
community want to take special measures to pay tribute to the "traditional,"
yet discounted, association with the latter family? We disagree with the
authors' rationalization for conservation, and prefer that Plantaginaceae--a
much better known name than either Antirrhinaceae or Veronicaceae--be allowed
to hold sway. But it is also our conviction that personal preferences should
play no role in these matters; this is precisely the kind of disagreement
that the rules of priority were designed to avert, impartially, in the first
Finally, we wonder about the fate of the name Orobanchaceae vis-à-vis
Rhinanthaceae. Why do the authors continue to use the former name, when
they are so anxious to discard Plantaginaceae? The situations appear analogous,
except that, in this case, conservation may not even be necessary, because
the two names were published simultaneously by Ventenat (1799). Perhaps
we can expect another proposal in the near future. And (ultra-finally--as
long as we are giving sway to personal preferences!) what about the possibility,
not discussed here, of putting all the pieces back together again--i.e.,
simply merging Callitrichaceae, Orobanchaceae, and Plantaginaceae into one
big, familiar and comfortable, Scrophulariaceae? Would that be monophyletic?
Poa pratensis L., the familiar Kentucky bluegrass (adventive in Costa
Rica), is treated by some recent authors as comprising two distinct spp.
Turns out that the type does not represent P. pratensis s. str.
but, rather, its circumboreal segregate, P. alpigena (Fr. ex
Blytt) Lindm. This proposal is of no consequence to workers who continue
to treat P. pratensis in the broad sense.
- Soreng, R. J. & F. R. Barrie. 1999. (1391) Proposal to conserve
the name Poa pratensis (Gramineae) with a conserved type.
Taxon 48: 157--159.
Five varieties of this weedy herb are recognized, and the appropriate new
combinations proposed. Four of the varieties are restricted to southern
Brazil and adjacent regions of South America. Costa Rican material would
fall into the autonymic variety, widespread from the United States to south
- Souza, V. C. 1997. Considerações sobre a delimitação
de Mecardonia procumbens (Mill.) Small (Scrophulariaceae). Acta
Bot. Brasil. 11: 181--189.
Separate and combined cladistic analyses of ITS sequence and cpDNA restriction-site
data all identified four major clades within Clerodendrum s. l.,
and showed the genus to be polyphyletic. These results support previous
studies suggesting that one of these four clades be removed from Clerodendrum
s. l. as a separate genus, Rotheca Raf. (see Novon 8: 204--206, 1998),
comprising five Old World spp. Furthermore, the oligotypic, North American
Tetraclea will have be submerged in Clerodendrum s. str.
These changes do not affect any indigenous New World spp. traditionally
assigned to Clerodendrum, nor any Old World spp. commonly cultivated
in Costa Rica.
- Steane, D. A., R. W. Scotland, D. J. Mabberley & R. G. Olmstead.
1999. Molecular systematics of Clerodendrum (Lamiaceae): ITS sequences
and total evidence. Amer. J. Bot. 86: 98--107.
"Embryologically Drypeteae do not fit within [Euphorbiaceae
subfam.] Phyllanthoideae and, as available nucleotide sequence data
from the rbcL gene suggest, are rather placed near Erythroxylaceae,
Rhizophoraceae, Chrysobalanaceae, and Linaceae. Drypeteae
share with these families a combination of the fibrous exotegmen, the endothelium,
and the thick, multiplicative inner integument."
- Tokuoka, T. & H. Tobe. 1999. Embryology of tribe Drypeteae,
an enigmatic taxon of Euphorbiaceae. Pl. Syst. Evol. 215: 189--208.
Rejection of the obscure Adiantum dissimile Schrad. is recommended,
protecting the junior but widely used A. deflectens Mart. [see The
Cutting Edge 4(2):
8, Apr. 1997]. Several other rulings are handed down, none of which
appears to affect the status quo.
- Zimmer, B. 1999. Report of the committee for Pteridophyta: 14.
Taxon 48: 133--134.