Bromus catharticus in South America (Poaceae: Bromeae)
Paul Peterson1998, Novon
visibility
…
description
61 pages
link
1 file
AI-generated Abstract
A new combination, Bromus catharticus var. rupestris, is made. Based on morphological evidence this taxon seems best recognized at the varietal level. Detailed illustrations of B. catharticus var. rupestris and B. catharticus var. catharticus are included along with morphological descriptions, distributions, and representative specimens.
Related papers
A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in México and Central America
Phytotaxa, 2014
A taxonomic treatment of Bromus in México and Central America is presented. Twenty-two species and one variety are recognized in the region. Twelve of these species are native, and ten are adventive and introduced. Bromus section Mexibromus is newly described, representing a distinct lineage in molecular studies of the genus. It includes two species endemic to México (B. attenuatus, B. densus) and one endemic to México and Guatemala (B. dolichocarpus), distinguished from species in other Bromus sections by their 3(-5)-nerved lemmas. Bromus pinetorum is a poorly understood species of Bromus sect. Bromopsis endemic to Coahuila. Taxa that have been variously treated in the past are discussed in detail. Bromus mucroglumis and B. thysanoglottis are treated as synonyms of B. richardsonii, and B. meyeri is treated as a synonym of B. anomalus. Two varieties are recognized in B. carinatus, B. carinatus var. carinatus and B. carinatus var. marginatus. Lectotypes are designated for the names B. carinatus var. californicus, B. hookeri var. schlechtendalii, B. trinii, B. trinii var. pallidiflorus and B. trinii var. stricta. A second-step neotype is designated for B. rigidus. Keys for determining the species in English and Spanish, descriptions, synonymies, illustrations, distribution maps, comments and lists of all specimens examined are provided for each species.
<strong>A taxonomic revision of <em>Bromus</em> (Poaceae: Pooideae: Bromeae) in México and Central America</strong>
Phytotaxa, 2014
A taxonomic treatment of Bromus in México and Central America is presented. Twenty-two species and one variety are recognized in the region. Twelve of these species are native, and ten are adventive and introduced. Bromus section Mexibromus is newly described, representing a distinct lineage in molecular studies of the genus. It includes two species endemic to México (B. attenuatus, B. densus) and one endemic to México and Guatemala (B. dolichocarpus), distinguished from species in other Bromus sections by their 3(–5)-nerved lemmas. Bromus pinetorum is a poorly understood species of Bromus sect. Bromopsis endemic to Coahuila. Taxa that have been variously treated in the past are discussed in detail. Bromus mucroglumis and B. thysanoglottis are treated as synonyms of B. richardsonii, and B. meyeri is treated as a synonym of B. anomalus. Two varieties are recognized in B. carinatus, B. carinatus var. carinatus and B. carinatus var. marginatus. Lectotypes are designated for the names B...
Bromus ayacuchensis (Poaceae: Pooideae: Bromeae), a new species from Peru, with a key to Bromus in Peru
2006
Bromus ayacuchensis, a new species from Departamento Ayacucho, Peru, is described and illustrated. The new species is similar to species in Bromus sect. Ceratochloa, particularly B. cebddilla, because of its strongly keeled lemmas. It differs from B. cebddilla by having densely pilose culms near the base, adaxially pilose ligules 3-6.6 mm long, lower glumes 1-veined, upper glumes 3-veined, pilose lemmas (especially along the margins and near the apex), lemma margins that are tightly involute on lower 1/5-1/4, and lemma awns 3-5.5 mm long. Phylogenetic analyses of sequence data from the internal transcribed spacer regions of nuclear ribosomal DNA place B. ayacuchensis in a clade with species of sect. Ceratochloa, whereas analyses of plastid data from the trnL intron and the 3'-end of ndhF indicate that B. ayacuchensis is closely related to this clade. Confirmed records are provided for B. modestus and B. cebadilla in Peru, and B. coloratus and B.flexuosus are reported for the fir...
Typification of eight current and seven related names and a new section in the genus Bromus (Bromeae, Pooideae, Poaceae)
PhytoKeys, 2019
During our nomenclatural revision of the genus Bromus L. (Poaceae) for Flora Iberica, we found that several names were still untypified: nine names in current use or their basionyms and five synonyms. Typifications are still needed since stabilising the names will facilitate their use. We propose lectotypes for Bromus alopecuros Poir., B. contortus Desf. (and the superfluous B. alopecuroides Poir.), B. benekenii (Lange) Trimen, B. intermedius Guss. subsp. divaricatus Bonnier & Layens, B. molliformis J.Lloyd ex Billot, B. lepidus Holmb., B. lepidus f. lasiolepis Holmb., Bromus subg. Stenobromus (Griseb.) Hack. and Bromus sect. Stenobromus Griseb. Neotypes for B. erectus Huds. and B. ramosus Huds. and an epitype for B. intermedius subsp. divaricatus Bonnier & Layens are proposed. In addition, we identify an isoneotype for B. erectus and isolectotypes for B. lepidus and B. lepidus f. lasiolepis. The area inhabited by the typified taxa includes both Africa and Europe. All the selected types are in agreement with the current use of the names and, thus, our selections contribute to stabilising the nomenclature of the genus Bromus. A discussion is provided to justify the selections. In addition, we typified two supraspecific names B. subg. Stenobromus and B. sect. Stenobromus. Finally, a new section, B. sect. Penicillius Llamas & Acedo, is described.
Figure 7 from: Llamas F, Acedo C (2019) Typification of eight current and seven related names and a new section in the genus Bromus (Bromeae, Pooideae, Poaceae). PhytoKeys 121: 53-72. https://doi.org/10.3897/phytokeys.121.30254
During our nomenclatural revision of the genus Bromus L. (Poaceae) for Flora Iberica, we found that several names were still untypified: nine names in current use or their basionyms and five synonyms. Typifications are still needed since stabilising the names will facilitate their use. We propose lectotypes for Bromus alopecuros Poir., B. contortus Desf. (and the superfluous B. alopecuroides Poir.), B. benekenii (Lange) Trimen, B. intermedius Guss. subsp. divaricatus Bonnier & Layens, B. molliformis J.Lloyd ex Billot, B. lepidus Holmb., B. lepidus f. lasiolepis Holmb., Bromus subg. Stenobromus (Griseb.) Hack. and Bromus sect. Stenobromus Griseb. Neotypes for B. erectus Huds. and B. ramosus Huds. and an epitype for B. intermedius subsp. divaricatus Bonnier & Layens are proposed. In addition, we identify an isoneotype for B. erectus and isolectotypes for B. lepidus and B. lepidus f. lasiolepis. The area inhabited by the typified taxa includes both Africa and Europe. All the selected types are in agreement with the current use of the names and, thus, our selections contribute to stabilising the nomenclature of the genus Bromus. A discussion is provided to justify the selections. In addition, we typified two supraspecific names B. subg. Stenobromus and B. sect. Stenobromus. Finally, a new section, B. sect. Penicillius Llamas & Acedo, is described.
Figure 9 from: Llamas F, Acedo C (2019) Typification of eight current and seven related names and a new section in the genus Bromus (Bromeae, Pooideae, Poaceae). PhytoKeys 121: 53-72. https://doi.org/10.3897/phytokeys.121.30254
During our nomenclatural revision of the genus Bromus L. (Poaceae) for Flora Iberica, we found that several names were still untypified: nine names in current use or their basionyms and five synonyms. Typifications are still needed since stabilising the names will facilitate their use. We propose lectotypes for Bromus alopecuros Poir., B. contortus Desf. (and the superfluous B. alopecuroides Poir.), B. benekenii (Lange) Trimen, B. intermedius Guss. subsp. divaricatus Bonnier & Layens, B. molliformis J.Lloyd ex Billot, B. lepidus Holmb., B. lepidus f. lasiolepis Holmb., Bromus subg. Stenobromus (Griseb.) Hack. and Bromus sect. Stenobromus Griseb. Neotypes for B. erectus Huds. and B. ramosus Huds. and an epitype for B. intermedius subsp. divaricatus Bonnier & Layens are proposed. In addition, we identify an isoneotype for B. erectus and isolectotypes for B. lepidus and B. lepidus f. lasiolepis. The area inhabited by the typified taxa includes both Africa and Europe. All the selected types are in agreement with the current use of the names and, thus, our selections contribute to stabilising the nomenclature of the genus Bromus. A discussion is provided to justify the selections. In addition, we typified two supraspecific names B. subg. Stenobromus and B. sect. Stenobromus. Finally, a new section, B. sect. Penicillius Llamas & Acedo, is described.
Figure 4 from: Llamas F, Acedo C (2019) Typification of eight current and seven related names and a new section in the genus Bromus (Bromeae, Pooideae, Poaceae). PhytoKeys 121: 53-72. https://doi.org/10.3897/phytokeys.121.30254
During our nomenclatural revision of the genus Bromus L. (Poaceae) for Flora Iberica, we found that several names were still untypified: nine names in current use or their basionyms and five synonyms. Typifications are still needed since stabilising the names will facilitate their use. We propose lectotypes for Bromus alopecuros Poir., B. contortus Desf. (and the superfluous B. alopecuroides Poir.), B. benekenii (Lange) Trimen, B. intermedius Guss. subsp. divaricatus Bonnier & Layens, B. molliformis J.Lloyd ex Billot, B. lepidus Holmb., B. lepidus f. lasiolepis Holmb., Bromus subg. Stenobromus (Griseb.) Hack. and Bromus sect. Stenobromus Griseb. Neotypes for B. erectus Huds. and B. ramosus Huds. and an epitype for B. intermedius subsp. divaricatus Bonnier & Layens are proposed. In addition, we identify an isoneotype for B. erectus and isolectotypes for B. lepidus and B. lepidus f. lasiolepis. The area inhabited by the typified taxa includes both Africa and Europe. All the selected types are in agreement with the current use of the names and, thus, our selections contribute to stabilising the nomenclature of the genus Bromus. A discussion is provided to justify the selections. In addition, we typified two supraspecific names B. subg. Stenobromus and B. sect. Stenobromus. Finally, a new section, B. sect. Penicillius Llamas & Acedo, is described.
Figure 5 from: Llamas F, Acedo C (2019) Typification of eight current and seven related names and a new section in the genus Bromus (Bromeae, Pooideae, Poaceae). PhytoKeys 121: 53-72. https://doi.org/10.3897/phytokeys.121.30254
During our nomenclatural revision of the genus Bromus L. (Poaceae) for Flora Iberica, we found that several names were still untypified: nine names in current use or their basionyms and five synonyms. Typifications are still needed since stabilising the names will facilitate their use. We propose lectotypes for Bromus alopecuros Poir., B. contortus Desf. (and the superfluous B. alopecuroides Poir.), B. benekenii (Lange) Trimen, B. intermedius Guss. subsp. divaricatus Bonnier & Layens, B. molliformis J.Lloyd ex Billot, B. lepidus Holmb., B. lepidus f. lasiolepis Holmb., Bromus subg. Stenobromus (Griseb.) Hack. and Bromus sect. Stenobromus Griseb. Neotypes for B. erectus Huds. and B. ramosus Huds. and an epitype for B. intermedius subsp. divaricatus Bonnier & Layens are proposed. In addition, we identify an isoneotype for B. erectus and isolectotypes for B. lepidus and B. lepidus f. lasiolepis. The area inhabited by the typified taxa includes both Africa and Europe. All the selected types are in agreement with the current use of the names and, thus, our selections contribute to stabilising the nomenclature of the genus Bromus. A discussion is provided to justify the selections. In addition, we typified two supraspecific names B. subg. Stenobromus and B. sect. Stenobromus. Finally, a new section, B. sect. Penicillius Llamas & Acedo, is described.
A new species in the Genus Bromus
Edinburgh Journal of Botany, 1990
The material here named Bromus lusitanicus Sales & P. M. Smith, sp. nov. (Gramineae) was discovered and collected during 1982-83 at Souzelas near Coimbra (central Portugal). It shows marked affinities to B. racemosus L. and B. pseudosecalinus P. M. Smith. These species form a complex of annuals which have 3-veined lower glumes, 5-veined upper glumes, subterete spikelets, margins of lemma more or less inrolled in fruit, and the awn terete, straight or curved, arising just below the apex of the lemma, although sometimes absent.
Bromus
catharticus
in
South
America
(Poaceae:
Bromeae)
Paul M. Peterson
Department of Botany NHB-166, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
Ana Maria Planchuelo
CONICET: Facultad de Ciencias Agropecuarias, Universidad Nacional Cordoba,
Casilla de Correo 509, 5000 Cordoba, Argentina
ABSTRACT. A new combination, Bromus catharti¬
cusvar.rupestris,ismade.Basedonmorphological
evidence this taxon seems best recognized at the
varietal level. Detailed illustrations of B. catharti¬
cusvar.rupestrisandB.catharticusvar.catharticus
areincludedalongwithmorphologicaldescriptions,
distributions,andrepresentativespecimens.
Resumen. Se realiza una nueva combinacion,
Bromuscatharticusvar.rupestris.Sobrelabasede
lasevidenciasmorfologicas,parecemejorreeonocer este taxon a nivel de variedad. Se incluyen illustracionesdetalladasdeB.catharticusvar.rupes¬
tris y B. catharticus var. catharticus junto con
descripcionesmorfologicas,distribueionesyespecfmenes
representativos.
BromusL.comprisesapproximately150species
found mainly in the temperate regions of both
hemispheres (Clayton & Renvoize, 1986). The ge¬
nus has been divided into five to seven groups
(Stebbins, 1981; Tsvelev, 1976), depending upon
the recognition of Anisantha C. Koch, Boissiera
Steudel,BromopsisFourreau,CeratochloaP.Beauvois, and Nevskiella Krecz & Vvedensky as sepa¬
rategenera.InarecenttreatmentofBromusinthe
United States and Canada, Pavlick (1995) recog¬
nized five sections: Bromopsis, Bromus, Cerato¬
chloa,Genea,andNeobromus.
In North and South America, Bromus sect. Cer¬
atochloa consists of approximately 18 species of
which B. catharticus Vahl is a member (Matthei,
1986; Nicora, 1978; Pavlick, 1995; Pillay & Hilu,
1995; Soderstrom & Beaman, 1968; Tovar, 1993;
Zuloaga et al., 1994). This section is entirely native
to the New World and contains an extensive poly¬
ploid complex, base of x = 7. All taxa thus far
examined are either hexaploid (2 n = 42), octoploid
(2 n = 56), or duodecaploid (2n = 84) (Armstrong,
1991; Pavlick, 1995; Stebbins, 1956, 1981; Steb¬
bins & Tobgy, 1944). Bromus catharticus and im¬
mediaterelativesareallhexaploid(Naranjo,1992).
TheBromuscatharticuscomplexpresentlycon¬
tains six taxa (Planchuelo, 1991; this paper): B.
bonariensis Parodi & J. A. Camara, B. brevis Nees
ex Steudel subsp. brevis, B. brevis subsp. festucanoides Covas & Millot, B. catharticus, B. parodii
Covas & Itria, and B. striatus Hitchcock [= B. ca¬
tharticusvar.striatus(Hitchcock)Pinto],Wechoose
to recognize B. catharticus, as proposed by PintoEscobar (1976), as the true name for rescue grass,
rather than that proposed earlier by Raven (1960)
as B. willdenowii Kunth. The Andean species, B.
unioloides Kunth, and B. willdenowii are treated
hereastaxonomicsynoymsoftheolder,B.cathar¬
ticus.
StudiesoftheBromuscatharticuscomplex(ex¬
cluding B. striatus) indicate that the crossability
among four species (B. bonariensis, B. brevis, B.
catharticus, B. parodii ) is low (Naranjo, 1992). In
recentyearstheuseofhybridizationstudiesinsystematics has declined since exceptions to the cor¬
relation between pairing and genome homology
havebeenfound(Seberg,1989;Doyleetal.,1990a,
1990b).Severalauthors(Ragonese&Marco,1941,
1943; Perez Lopez, 1975; Cladera, 1979; Pahlen et
al., 1980; Wolff et al., 1996) described the mor¬
phologicalfeaturesofB.catharticusundervarying
environmentalconditions.Morerecently,Abbottet
al. (1996), Aulfcino and Arguri (1996), and Gu¬
tierrez et al. (1996) pointed out phenotypic varia¬
tioninthevegetativefeaturesofB.catharticus.The
polymorphic nature of B. catharticus has led to the
description of numerous taxa at the specific and
varietallevels,andweofferourtreatmenthere.Pri¬
orresults(Planchuelo,1991)ofapheneticanalysis
of16morphologicalcharacterssuggestthatB.bon¬
ariensis,B.brevis,andB.catharticusbeconsidered
conspecific.
At the present time we choose to recognize B.
bonariensis,anendemicofProvinciaBuenosAires,
Argentina, as a separate species since there ap¬
pears to be no overlap with other taxa in the comNovon 8: 53-60. 1998.
54
Novon
plex in the length of the lemmatal awns (5-9 mm).
We recognize B. striatus, an endemic of Peru, as a
separate species, as opposed to a variety (PintoEscobar, 1986), since there is no overlap in the
lengthofthelemmatalawns(12—18(—25)mm).The
status of B. bonariensis and B. .striatus is currently
beinginvestigatedforDNAmarkersbytheauthors.
Inthisstudy,acomparisonofmorphologyamong
members of the Bromus catharticus complex has
been made, and the recognition of B. brevis at the
varietallevelseemswarranted.Inthisrankvariety
rupestris,basedonB.unioloidesvar.rupestrisSpegazzini,ataxonomicsynonym,haspriority.Wein¬
cludedescriptions,synonymy,representativespec¬
imens, a key to the varieties, and an illustration of
B.catharticusvar.rupestris(Spegazzini)Planchuelo
& P. M. Peterson and variety catharticus.
pression, then the spikelets fusiform. Glumes
broadly ovate; tin* lower 5-11 mm long, 1.6-3 mm
wide, 5—7-nerved; the upper 6-12 mm long, 9-11nerved. Lemmas 7-15 mm long, 2—4 mm wide, 911-nerved,ovate,glabroustopubescent;themar¬
ginalnervesbifurcatingfromthebaseofthesecond
nerve from the keel; apex acute or cuspidate, the
awn 0.3-0.5(1) mm long. Paleas 5-9 mm long,
stronglykeeledandadherenttothecaryopsis.An¬
thers0.4—0.7mmlong.Caryopsiswithadeep,nar¬
row
furrow.
KKY
'll)
TUK
V
AUIKTIKS
OK
BliOMl
'.S'
CATHAKTICLS
la.Lemmasmueronateorwithashortawn0.3—0.5(—1)
mmlong;youngbladespredominantlyfolded.
.
B.
catharticus
var.
rupestris
lh.Lemmaswithanawn0.5—4(—5)mmlong;young
blades
predominantly
convolute.
.
B.
catharticus
var.
catharticus
Bromus catharticus Vahl var. rupestris (Spegaz¬
zini) Planchuelo & P. M. Peterson, comb. nov.
Basionym:BromusunioloidesKunthvar.ru¬
pestris Spegazzini, Contr. FI. Sierra Vent. 76:
1896. TYPE: Argentina. Prov. Buenos Aires:
Sierra Ventana, Spegazzini 12624 (lectotype,
heredesignated:specimenwasmarkedas“TypusdeBromusunioloidesvar.rupestrisSpe¬
gazzini,” LP!). Figure 1.
BromusbrevisNeesexSteudel,Syn.PI.Glumac.1:326.
1854.CeratochloabrevisNeesexJackson,Index
kew.1:487.1895.Bromusunioloideskunthvar.
brevis(NeesexSteudel)llackel.AnalesMus.Nae.
Hist.Nat.BuenosAiresII:144.1904.Bromusuni¬
oloideskunthf.brevis(NeesexSteudel)kloos,Ned.
kruidk.Arch.1917.175.1918.TYPE:Argentina.
Mendoza,Gilliess.n,(isotype,k,fragment,US!).
Bromus
brevis
Nees
ex
Steudel
subsp.
festucarioides
Covas
&Millot,ApuntesLI.Pampa62:247.1981.TVPE:
Argentina.
Prov.
La
Pampa:
Departamento
Caleu,
La
Adela,G.Covas2163(isotype,CORD!).
Plants annual or biennial. Culms (5)30-60(70)
cm tall. Ligules 2—3 mm long. Sheaths pubescent,
the hairs up to 2 mm long. Blades 2.5-5 mm wide,
flatorfolded(youngbladespredominantlyfolded),
pubescent.Panicles(4)10-15cmlong,erect,semicontractedinflorescencewith2-45spikelets;ped¬
icels3—25mmlong,rigid.Spikelets(10)13—20(25)
mm long, 5-10-flowered, imbricate, commonly
compressedlaterally,occasionallywithlittlecom¬
Distribution. Known from central and south¬
west Argentina from Provincias Buenos Aires and
Cordobainthenorth,southtoSantaCruz,whereit
occurs on sandy river banks and slopes from ele¬
vations of near sea level to 2600 m. This endemic
variety appears to be good forage for cows (Bagonese, 1967) and is infrequently found in open
pampas/punahabitatsandmarginsofforests.The
common name for this species is “Cebadilla pampeana.”
Representative
specimens.
ARGENTINA.
Buenos
Ai¬
res;Campana,1Nov.1936,/..Parodi12244(BAA.US);
Partido de Junfn, 21 Dec. 1928. E. G. Glos 3976 (US);
Estancia“liesGhanaves,”E.Gibsons.n.(US);Depto.Necoehea,PuntaNegra.12Dec.1938.W.J.Eyerdam.A.A.
Beetle&E.Grondona23729(US);Depto.Villarino,60
km E of Rio Colorado, 16 Dec. 1938, IE J. Eyerdam. A.
A.BeetleAE.Grondona23744(US).Cluihut:Depto.
Rawson,2kmSofTrelewonSbankoftheChubutRiver,
22Dec.1938,IE./.Eyerdam.A.A.BeetleAE.Grondona
23582(US).Cordoba:Tupungato,EstanciaLaCarrera,
27Dec.1949,0.MetisA0.Pad44(US):Depto.General
Roca,caucesecodelRfoQuinto,enlosalrededoresdel
cruceconlaRutaNacional35,entreHuincaRenancoy
VicunaMackenna,3Nov.1971.A.T.HunzikerAI).Hil¬
da21451(CORD);Depto.Ischilm,DeanEunes,20Dec.
1946.M.Villafane129(US);Depto.RoqueSaenzIVila.
CercadePachecodeMelo,yendoaLaboulaye,1Nov.
1956.A./.'Ilunziker12772(CORD):Deptos.lerceroAr¬
riba/SanMartin,EstanciaPatria,entreLucayDalmacio
Velez Sarsfield. 18 Jan. 1956, A. T. Hunziker 11566
(CORD);Depto.RfoSegundo,ColoniaRuiz,pedanfaMatorrales,cercadeVilladelRosario,12Nov.1902,Stuckert
12045(CORD).I,aPampa:GrabAcha,29Nov.1959,
V..S’.Troncoso20495(SI,LIS)..Mendoza:Depto.I.aslleras,CuestadeLasMinas,18/22Jan.1897.Kurtz9369
(CORD);PuentadelInca,3Jan.1950,().Pan161(US);
Depto.Lujan,Ugarteche,19Nov.1955,F.A.Roig7396
(CORD,MERL,US);UsAguaditas,18Dec.1985,Dalmasso429(MERL);Depto.Malargiie,Coihueconorte,15
Nov. 1959, F A. Roig 3446 (CORD, MERL, US); Los
MollesinmediacionesrielhotelLahuen-co,20Dec.1981,
DelVittos.n.(MERL);Depto.Maipu,MedanosdelosMorritos(ElTapon),27Nov.1949.A.R.D'<d&0.Pan487
(US);Depto.SanCarlos,EntreYagualitoyLasPenas,5
Dec.1887.Kurtz5409(CORD);Depto.SanRafael,Entre
SantaMariayEortinNuevo,12/13Jan.1892.Kurtz7042
(CORD);CuadroBenegasentreRutaNae.144yRio
Atuel,12Nov.1981,DelVittos.n.(MERL);Cercaniadel
emhalseAguadelToro,22Nov.1996,A.M.Planchuelo
Volume 8, Number 1
1998
Peterson & Planchuelo
Bromus catharticus in South America
55
FigureI.BromuscatharticusVahlvar.rupestris(Spegazzini)Planchuelo&P.M.Peterson.—A.Habit.—B.Ligule.
—C.Spikelet.—D.Glumes.—E.Lemma.—F.Palea.—G.Gynoeciumandandroecium.—H.Caryopsis.Basedon
Fischer51(US).
56
Novon
1068(ACOR);Depto.Tupungato,RodeoSeguro,12Nov.
1953, F. A. 139 ((]()K1), MERI., US). Neuquen: Pino
Hachado,Feb.1920,L.R.Parodi3139(BAA,US);Sierra
AucaMahuda,Nov.1953,11.A.Fabris802(US);Depto.
ChosMala,EntreChosmalalyChorriaea,orillasdelRfo
I’ichi-Neuquen,1<>Nov.1969,F.A.Roig6300(CORD,
MERE,US);Depto.Picunches,UasLajas,IBNov.1969,
F.A.Roig6247(CORD,MERL,US).SantaCruz:Puerto
SantaCruz.22Dee.1895,P.Duseils.n.(US);Depto.Deseado,PuertoDeseado,29Dec.1938,HiJ.Eyerdam,A.
A.BeetleA'F.Grondona23870(US);Depto.GuarAiken,
RioGallegos,3Jan.1939,W.J.Eyerdam,A..4.Beetle&
E.Grondona24072(US).SanJuan:Depto.Iglesias,Si¬
erradeColangUil,1887/1888,Kurtz3354(CORD).Kio
Negro: Los Juneos. 27 Feb. 1934, /.. R. Parodi 11372
(BAA,US).SanLuis:AlrededoresdeSanLuis,5Nov.
1960,G.Goras1085(US);GeneralRoeaandvicinity.21
Jan.1915.1LFischer51(US);Depto.GeneralPedernera,
Ruta148entreLavaisseyElDurazno,23Nov.1962,A.
T. Hunziker & A. Cocucci 15978 (CORD); al borde de
medanoalnortedeLagunaSayape,7Nov.1969,I).An¬
derson1613(CORD);Depto.Pringles,CereadeFragaa
linos40dmdeVillaMercedes,Ruta7,14Nov.1956,A.
T.Hunziker13139(CORD).Tucuman:VillaNougues,4
Nov.1930,G.LFawcetts.n.(US).
BromusstrictusBrongniart,inDuperrey,Voy.Monde2:
45. 1829. TYPE: Brazil. Santa Catarina: 1825,
D'Urvilles.n.(holotype,P,fragment.US!,fragment
&photo,BAA!).
Althoughthevenationcharacteristicofthelem¬
maappearstoheusefulinseparatingB.catharticus
var. rupestris from variety catharticus it is often
(usually) very difficult to see even under 20X mag¬
nification. Other specimens, such as Kurtz 10034
(CORD) from Puesto Lima, Mendoza, have a few
spikelets with additional bracts below the glumes
that are 1—3-nerved. Another specimen (Kurtz
5668, CORD), from the Rio Atuel, Mendoza, has a
few spikelets with an additional 5-nerved bract at
thebase.Besidesthesefewaberrantcharacteris¬
tics,allothercharacteristicsseemtobeconsistent
withthosedescribedforthisvariety.
Bromus catharticus Vahl, Symb. Bot. 2: 22.
1791. var. catharticus. Ceratochloa cathartica (Vahl) llerter. Rev. Sudamer. Bot. 6: 144.
1940. TYPE: Peru. Lima: J. Dombey s.n. (lectotype,selectedbyPinto-Escobar(1976)P-JU,
microfiche, US!; isolectotype, P). Figure 2.
FestueaunioloulesWilldenow,Hort.Berol.1;3.pi.3.
1803.Ceratochloaunioloides(Willdenow)P.Beauvois,Ess.Agrostogr.75,t.15,f.7.1812.Bromus
unioloides(Willdenow)Raspail,Ann.Sci.Nat.Bot.
5:439.1825,horn,illeg.,notkunth1816.Bromus
willdenowiikunth,Revis.Cramin.1;134.1829.
Tragusunioloides(Willdenow)PanzexB.1).Jackson, Ind. kew. 2: 1099. 1895. TYPE: (holotype,
grownatBerlinfromseerlfromCarolinas,USA,BW.
microfiche,
US!).
Bromusunioloideskunth,Nov.(fen.Sp.1:151.1816.
Schedonorus
unioloides
(kunth)
Roemer
&
Schultes,
Syst.Veg.2:709.1817./ernaunioloides(Kunth)
Lindmnn.Sv.Fanerogamfl.101.1918.TYPE:Ec¬
uador.Piehincha:Humboldt&Bonpland2286(ho¬
lotype,P!,fragment,US!:isotype,P).
CeratochloahaenkeanaJ.S.Presl,Reliq.Haenk.1:285.
1830.Bromushaenkeanus(J.S.Presl)kunth,Enum.
PI.1:416.1833.Bromusunioloidesvar.haenkeanus
(J.S.Presl)Shear,U.S.Dept.Agr.Div.Agrost.Bull.
23:52.PXM).TYPE:Chile.T.Haenkes.n.(holotype,
PR,
fragment,
US!).
CeratochloasecundaJ.S.Presl,Reliq.Haenk.1:285.
1830.Bromuspresliikunth,Enum.PI.I:416,545.
1833.TYPE:Peru.T.Haenkes.n.(holotype,PR,
fragment,
US!).
CeratochloabreviaristataHooker,F.Bor.Amer.2:253.
1840.
Bromus
breviaristatus
(Hooker)
Thurber,
U.
S.
Expl.Exped.Bot.17:493.1874,notBuckley,1862.
Forasaccus
breviaristatus
(Hooker)
Lunell,
Amer.
Midi. Naturalist 4: 225. 1915. TYPE: Lewis and
ClarksRiverandnearsourcesoftheColumbia,
1826,Douglass.n.(holotype,K,fragment,US!).
Bromus
unioloides
kunth
var.
sanjuaninus
Hieronymus,
Bol.Acad.Nac.Ci.4:69.1881.TYPE:Argentina.
Prov.SanJuan,Echegarays.n.(isotypes,BAA!,
CORD!).
BromusangustatusPilger,Bot.Jahrb.Syst.25:719.1898.
TYPE:Bolivia.LaPaz,Steubel60c(holotypefrag¬
ment
US!).
Bromusunioloideskunthvar.montanusHackelexStuekert,AnalesMus.Nac.BuenosAires11:144.1904.
Syn.nov.TYPE:Argentina.Prov.Cordoba:Cueva
delArroyodelosTabaquillos,SierradeAchala,
Stuckert10855(holotypefragmentUS!;isotype,
CORD!).
Bromus
unioloides
kunthf.chasmogamaHackel,Anales
Mus.Nac.Hist.Nat.BuenosAires13:527.1906.
TYPE:Argentina.Prov.Cordoba:AltosSuddeCbrdoba,
Stuckert
3448a
(isotype,
CORD!).
Bromusunioloideskunthf.chasmogamaHackelsubf.
achalensisHackel&Stuckert,AnalesMus.Nac.
Hist.Nat.BuenosAires21:172.1911.Syn.nov.
TYPE:Argentina.Prov.Cbrdoha:EstanciaPampade
SanLuis.Achala,Stuckert20651(isotype,CORD!).
Caespitoseannual,biennial,occasionallyperen¬
nial. Culms (5)20—100(120) cm tall. Ligules 2-5
mmlong.Sheathsglabrousorsparinglypubescent.
Blades 2—9 mm wide, usually flat (young blades
predominantlyconvolute),glabrousorsparinglypu¬
bescent. Panicles (2.5)10-20(30) cm long, 5-9flowered,erectorlax,semi-contracted;primaryin¬
florescencewith4—58spikelets;lateralbranches
sometimesnoddingatmaturity;pedicels8—35mm
long.Spikelets(7)15-30(40)mmlong,4—11-flowered,veryimbricateandcompressedlaterally,oc¬
casionallywithlittlecompression,thenthespike¬
lets fusiform, often purplish. Glumes ovate,
glabrous;thelower6—12mmlong,1.6—4mmwide,
3—7-nerved; the upper 8-14 mm long, 2.2-5 mm
wide,(5)7—9-nerved.Lemmas10—17mmlong,3—
Volume 8, Number 1
1998
Peterson & Planchuelo
Bromus catharticus in South America
57
Figure 2. Bromus catharticus Vahl var. catharticus. —A. Habit. —B. I.igule. —C. Spikelet. —1). Glumes. —E.
Lemma.—IFalea.
.
—G.Gynoeciumandandroecium.—H.Caryopsis.BasedonPeterson,Amiable,Lcegaard,Soreng
&Rojcts-Ponce12824(AAl,BH,K,LPB,US).
58
Novon
6 mm wide, 7-11-nerved, ovate, glabrous or occa¬
sionally minutely pubescent; the marginal nerves
originatingindependentlyfromthebaseofthesec¬
ond nerve from the keel; apex acute, aristate, the
awn 0.5^4(5) mm long. Paleas 8-14 mm long,
stronglykeeledandadherenttotheearyopsis.An¬
thers0.5—1.3mmlong.Caryopsiswithadeep,nar¬
row
furrow.
Depto.LaCapital,IslaTimbo,17Oct.1996,A.M.Plan¬
chuelo1085A*1061(ACOR).Tueumau:Depto.laff.
CunibredeMalamala,M.Lillo8510(LIL).BOLIVIA.Co¬
chabamba:(Juillaeollo,CaminoSipe—Lipichi.22Mar.
1990,/.Hanson651(US).LaPaz:I.aFlorida,21Dec.
1923,A.S.Hitchcock22611(US):LikeTiticaca.Omasuyas,22Feb.1987,.S’.,4.Renvoize4577(US);Prov.Ingavi.Titieani-Tacaca,I1May1989,V.Villaviceneia1008
(US);Prov.Lareeaja.Sorata,1886.II.II.Rushy225(US);
Prov.Aroma,Huaraco,10Jan.1981.M.Liberman274
(US): Prov. Loayza. 12.1 mi. NW of Villa Liza, 4 Mar.
1993,P.47.Peterson.C.R.Amiable.S.heguardAR.J.
Soreng12647(US);Prov.NorYungas,Unduavi,3Apr.
1981, .S. A. Renvoize A 71 A. Cope 4178 (US). Oruro:
Prov.Cerado,14mi.SofOruro,6Mar.1993,I'.,47.Pe¬
terson.C.R.Amiable,.S',hrguardAR.J.Soreng12697
(US):Prov.Poopa,4.5mi.NofPazna,6Mar.1993,I’.M.
Peterson,C.R.Amiable,S.LcegaardAR.J.Soreng12719
(US):Prov.S.Pagador,5mi.SofChallapata.7Mar.1993,
P.iM.Peterson,C.R.Amiable.S.LcegaardAR.J.Soreng
12784(US).Potosi:AltiplanoCereadeTupiza,Feb.
1948,A.Bridorolli4807(US);Prov.Ouijarro,4mi.S\\
ofVilaeota,28Mar.1993,P.M.Peterson.R.,/.Soreng&
S.hvgaard18127(US);SudChichas,9Jan.1924,A..S'.
Hitchcock22877(US);58mi.SKofUyuni.13Mar.1993,
P.47.Peterson.C.R.Amiable,S.Lcegaard,R.J.Soreng
AF.Rojas-Ponce12852(US).BRAZIL.SantaCatarina:
Caxambu,29Oct.1963,R.47.Kleins.n.(US);Caxiasdo
Sul.13Nov.1986,C.C.Grazziotin1980(US):SanBenito
do Sul. 26 Nov. 1972. R. 47. Klein s.n. (US); Mini. Blumenau,Spitzkopf,20Mar.1952,L.II.SmithAP.R.Reitz
6275(US);Mini.Uruguai,VilaRica,24Oct.1964,L.II.
SmithAP.R.Reitz12919(US).Parana:Curitiba,2Oct.
19(g).Lindeman2666(US).RioGrandedoSul:Irai.8
Jan.1947.L.Raupp142(US);PortoAlegre,5Nov.1936,
lbA.Archer4880(US);Mini.RioPardo,FasendaSoledade,Oct.1922.C.Jurgenss.n.(US);Mini.Vaearia,14
km de Vaearia rumo Bom Jesus, 25 Oct. 1961, C. F. J.
Pabst6884(LIS).SaoPaulo:10kmSofSaoPaulo,20
Oct.1966,71Sendulsky881(US).CHILL.Antofagasta:
Irrigatedpark.CiudadAntofagasta.24Jan.1924.A.S.
Hitchcock22985(US);nearCalama.4Nov.1914.J.I\.
Rose19482(US).Concepcion:Concepcion,JardfnZoologieo,4Feb.1958.F.Junge8092(US).Coquimbo:Coqiiimba I ,a Laguna, 3 Jan. 1945, F. Harms 5427 (US).
Santiago:LosCerrillos,12Oct.1941,II.Gunckel12525
(US);PuenteAltonearSantiago,3Oct.1919.F.14.I).
WilsonA47.47.Wilson88a(US);CerroSanCristobal,5
Sep.1950,II.Gunckel18708(US);SanJosedeMaipo,5
Oct. 1919, F. W. I). Wilson A 47. 47. Wilson 88 (US).
COLOMBIA.Anlioqiiia:SantaKlena,28Dee.1930.14.
A.Archer1199(US).Cauca:MacizoColombiano,7—27
Sep.1958,Idrobo,PintoAHischler8410(US).Cundinamarea:Bogota,May1916,ApollinaireAArthur26
(US).Narino:Tnquerres,May1853,./.Triana901(US).
NortedeSantander:Pamplona,23Mar.1935.141A.
Archer8281(US).KCUADOR.Azuay:ParqueNaeional
Cajas,21Apr.1990,P.47.Peterson.C.R.AmiableA47.
Poston8867(US).Bolivar:6.4kmKofGuaranda,23
MayP>90,I'.47.Peterson.F.J.Judziewicz.R.47.KingA
I’.47.Jorgensen9271(US).Canar:NearFII'ambo,4May
1945,141//.Camp2922(US).Carehi:5mi.SofTulean,
10Aug.1923,A.S.Hitchcock20974(US).Chimborazo—
Cotopaxi: 20 km N of Ambato, 16 Apr. I'XX). P. 47.
Peterson.C.R.AmiableA47.Poston8788(LIS).Imluibura:Pimampiro,21Nov.1949,47.Acosta-Sohs14589
(US).Imbabura—Pincbincha:KntreProantagyPesillo,
Distribution. A widespread taxon ranging from
NorthAmerica,CentralAmerica,toSouthAmerica,
whereitisnative.InArgentina,Hromu.scatharticus
var. catharticus is more common north of the 38th
parallel. An excellent forage grass in natural pas¬
turesintemperate,subhumid,andhumidzones.
Representative
specimens.
ARGKNTINA.
Buenos
Ai¬
res:DistritoFederal,Palermo,21Jan.1941,J.J.Rossi
60(kll.);Part.'landiI.Tandil,2Dee.1948,Meyer14408
(Lib).Catamarea:Depto.Ambato,CumbresdeNarvaez
(Falda 0), Ruta 92 km. 1436/1435 entre el mice eon
Ruta65yLasChaorilas,rumboaSinguil.,10Dec.1965,
A.71Hunziker,CornedA-Subils18585(CORD):Depto.
FIAlto,SierradeAneasti,Ruta64,entreeldesvi'oaFrias
yLosMorteros,2Dee.I960,A.T.HunzikerACocucci
15656(CORD);Depto.Roman.SierradeAmbato(Falda
W),Mutquin.entreColanayRincon,8Dee.1965,A.T.
Hunziker;CocucciA-Subils18482(CORD);Depto.Santa
Maria,Cerrillos,16Dee.1933,Peirauss.n.(US).Chaco:
Depto.PrimerodeMayo,ColoniaMargaritaHelen,30Jan.
1907,T.Stuckert19187(CORD).Cordoba:Depto.Calamuchita,ValledeLisReartes,30Dee.1919,Castellanos
s.n.(Lib);Deptos.Calamuchita/SanJavier,CerroCampaquf,16Dee.1885,Kurtz2976(CORD);Depto.Capital,
CiudadUniversitaria,10Dee.1990,A.M.Planchuelo
546(ACOR);Depto.SanAlberto,entreoiiestadelcorral
deCeballosyCireodelChampaquf,15Dee.1885,Kurtz
2967(CORD);Depto.SanMartin,entreLucayDalmacio
Velez,18Jan.1956,,4.T.Hunziker//564(CORD);Depto.
SantaMarfa,AltaGracia,9Jan.1940.4.T.Hunziker506
(CORD);entreLaSerranitayVillaCiudaddeAmerica,
KstanciaLaPraviana,If)Nov.1996,A.M.Planchuelo
1062(ACOR);Depto.Union,Ballesteros,KstanciaAtahiva,12Oct.1990,A.M.Planchuelo545(ACOR).Corrientes:Depto.Kmpedrado,KstanciaLaVela,20Oct.
1965, 7. 47. Pederson 7478 (US). Entre Rios: Depto.
Concordia,Kstaeionexperimental1NTA.22Dee.1992,
.4.M.Planchuelo598(ACOR).Jnjiiv:Depto.Cochinoea,
LagunaIresCruces,14Feb.1901,Kurtz11672(CORD);
Depto.SantaCatalina,9Jan.1901.Kurtz11415(CORD).
LaRioja:Depto.Capital,cereadelaminaK1Cantadero
(La Ksperanza), 5/6 Mar. 1944, A. 71 Hunziker 5192
(CORD);Depto.Famatina,SierradeFarnatina,LaMesada.20Mar.1906,Kurtz18845(CORD).Salta:Depto.
Guacbipas,Alemania,15Dec.1929,S.Venturi9954
(US):Depto.SanAntoniodeLosCobres,(JuebradaUreum,12Feb.1945,A.L.Cabrera8678(US).SanJuan:
CalingastaoCordilleradelKspinaeito.LosManantiales,
6 Feb. 1897, Kurtz 9594 (CORD); Depto. Rivadavia,
Margnesado,27Nov.1945,A.R.Cuezzo1502(US).San
Luis:Depto.SanMartin,SierradeSanLuis,SanMartin,
enI.aPileta,A.T.HunzikerA'Cocucci14586(CORD).
SantaFe:Depto.Cayasta,enquintaabandonadaallado
delaRuta,If)Oel.1996,A.M.Planchuelo7(//9(ACOR);
Volume 8, Number 1
1998
7Apr.1952,M.Acosta-Solis21126(US).Loja:between
LojaandSanLucas,6Sep.1923,A.S.Hitchcock21488
(US). Pinchincha: 41 km S of Quito, 13 Apr. 1990, P.
M.Peterson,C.R.Annable&M.Poston8780(US).Tungiiruhua:EntreYamboyAmbato,20Oct.1944,M.Acos¬
ta-Solis8561(US).PARAGUAY.Central:Asuncion,Jardfn Botanico, Nov. 1917, Rojas 3069 (US). PERU.
Amazonas:Prov.Bongara,SWofPomacocha,16June
1962,,/.J.Wurdack855(US).Aneash:Prov.Bolognesi,
Pariarraccra, 2 May 1952, E. Cerrate 1473 (US). Arequipa:Arequipa,22Nov.1923,A.S.Hitchcock22430
(US).Cajamarca:Prov.SanMiguel,CerroQuillon,5July
1986, J. Mostacero L., E. Alvitez /., S. Leiva G„ E Mejia
C.AE.PelaezP.1247(US).Cuzco:Prov.Quispicanchis,
near Oropesa, 2 Mar. 1963. I). A V. Ugent 4029 (US),
lliiancavelica:Prov.Huancavelica,Izcuchaca,16Apr.
1955,0.Tovar2484(US).Jum'ii:betweenOroyaandLa
Merced,24Oct.1923,A.S.Hitchcock22159(US).Lima:
Prov.Huarochiri,Infiernillo.10May1984.I).N.Smith,
R. Eerreyra A’ 0. Tovar 7007a (US). Madrr de Dios:
Juliaca. 26 Feb. 1914. //. U. Harlan s.n. (US). Puno:
LikeTiticaca,4June1954.Monheims.n.(US).URU¬
GUAY.
Montevideo:
American
Consulate,
Montevideo,
29Nov.1925.E.M.Marsh17(US).VENEZUELA.Fed¬
eralDistrict:Galipan,Dec.1932,Tamayo9(US).Lara:
Depto.Jimenez,nearAguaNegre,11Mar.1979,C.Burandt Jr.. II. Garofalo A E. Colton V0568 (US). Merida:
betweenMucurubaandParamodeMueucbies,22Jan.
1922.A.John774b(US).
Morphologicallyhighlyvariableandecologically
diverse,thisvarietywasintroducedasaforagecrop
and is now widely adventive in North America. The
commonnameofBromuscatharticusvar.catharti¬
cus in the United States is rescue grass; in Argen¬
tina it is “Cebadilla, Cebadilla criolla, or Cebadilla
australiana.”Thesynonymy,inpart,followsPintoEscobar (1986) and Zuloaga et al. (1994), other¬
wise, “Syn. nov.” is indicated.
Acknowledgments. We are grateful to Robert J.
Soreng for reviewing this manuscript. Special
thanks are given to Susan C. Escher for her skillful
illustrations.
Literature
Cited
Abbott,L.S.,P.PistoraleAB.Wolff.1996.Variabilidad
fenoti'pica
estructura
y
poblacional
en
Bromus
cathar¬
ticusVahl(cebadillacriolla).JornadasArgentinasde
Botanica25(Resum.):329.
Armstrong,K.C.1991.ChromosomeevolutioninBromus.
Pp. 336-377 in T. Tsuchiya & P. k. Gupta (editors).
Chromosome
Engineering
in
Plants:
Genetics,
Breed¬
ing.Evolution.PartB.Elsevier,Amsterdam.
Aulfeino, M. B. A M. E. Arturi. 1996. Clasifieacion de
poblaciones
de
Bromus
catharticus
Vahl
utilizando
caraeteresdealtaconstanciayherabilidad.JornadasArgentinasdeBotanica25(Resum.):331.
Cladera,J.1979.Polimorfismoisoenzimaticoyvariacion
paralalongituddeanterasenlacebadilla(BromusunioloidesHBk).TesisMagisterSoientiae.Escuelade
Graduados
Ciencias
Agropecuarias,
IN
PA
Castelar,
Ar¬
gentina.
Peterson & Planchuelo
Bromus catharticus in South America
59
Clayton,W.I).AS.A.Renvoize.1986.GeneraGraminum.kewBull.Add.ser.13:1-389.
Doyle, J. .).. J. L. Doyle A A. H. 1). Brown. 1990a. A
chloroplast-DNAphylogenyofthewildperennialrela¬
tivesofsoybean(GlycinesubgenusGlycine):Congru¬
ence
with
morphological
crossing
groups.
Evolution
44:
371-389.
-.
A
-.
1990b.
Chloroplast
DNA
phylogeneticaffinitiesofnewlydescribedspeciesin
Glycine
(Leguminosae:
Phaseoleae).
Syst.
Bot.
15:
466—
471.
Gutierrez, H. E, J. E. Pensiero A A. C. Vegetti. 1996.
TipologfadeinflorescenciadelgeneroBromusL.(Poaeeae:Poeae).JornadasArgentinasdeBotanica25(Re¬
sum.):
186.
Matthei, 0. 1986. El genero Bromus L. (Poaceae) en
Chile.Gayana,Bot.43:47—110.
Naranjo,C.A.1992.Estudioshiosystematicosenespecies
deBromus(seccidnCeratochloa,Poaceae).I.Sistemas
reproductivos
barreras
y
de
aislamiento.
Darwiniana
31:
173-183.
Nicora.E.G.1978.Gramineae.InM.N.Correa(editor).
FloraPatagonicaPartIII.ColeecionCientificadelINTA
8:1-563.
Pahlen, A. W. von der, J. V. Crisei. W. Telleria Polo AE.
PerezLopez.1980.Clasifieaciondepoblacionesdece¬
badillacriollaBromus
(
unioloides)ylacebadaboliviano
(Hordeumvulgare).ActasIVCongr.Latinam.Genetica
2:
207-211.
Pavlick,L.1995.BromusL.ofNorthAmerica.RoyalBrit¬
ish
Columbia
Museum.
British
Columbia,
Canada.
PerezLopez,E.1975.StudiodevariabilidaddeBromus
unioloides
de
diferentes
habitats.
Tesis
Magister
Soien¬
tiae.
Escuela
Graduados
Ciencias
Agropecuarias,
INTA
Castelar,
Argentina.
I’i1lay.M.Ak.W.Hilu.1995.Chloroplast-DNArestric¬
tionsiteanalysisinthegenusBromus(Poaceae).Amer.
J.Bot.82:239-249.
Pinto-Escobar,P.1976.Notasobreelejemplartipode
14Bromuscatharticus”Vahl.Caldasia11:9—16.
-. 1986. El genero Bromus en los Andes Centrales
de
Suramerica.
Caldasia
15:
15—34.
Planchuelo,A.M.1991.EstudiossobreeleomplejoBro¬
muscatharticus(Poaceae)I.Evaluacionestadfstieade
los
cardcteres
taxonomicos.
kurtziana
21:
243—257.
Ragonese,S.1967.VegetacidnyGanaderfadelaRepublicaArgentina.ColeecionCientfficadelINTA5.
-A P. R. Marco. 1941. Observaciones sobre la biologfa floral de la cebadilla criolla. Revista Argent.
Agron.8:196-199.
-A-. 1943. Influencia del fotoperiodo sobre
laformaeiondeflorescleistogamasychasmogamasen
cebadillacriolla.RevistaArgent.Agron.10:178—185.
Raven,P.H.1960.Thecorrectnameforrescuegrass.
Brittonia
12:
219-221.
Seberg,0.1989.Genomeanalysis,phylogeny,andclas¬
sification.PI.Syst.Evol.I(>6:159—171.
Soderstrom,T.R.AJ.H.Beaman.1968.ThegenusBro¬
mus(Gramineae)inMexicoandCentralAmerica.Publ.
Mus.MichiganStateUniv.3(5):469—519.
Stebbins,G.L.1956.Cytogeneticsandevolutioninthe
grassfamily.Amer.J.Bot.43:890—905.
-. 1981. Chromosomes and evolution in the genus
Bromus(Gramineae).Bot.Jahrh.Syst.102:359-379.
-A II. A. Tobgy. 1944. The cytogenetics of hybrids
inBromus.1.Hybridswithinsection44Ceratochloa."
Amer.J.Bot.31:1-11.
60
Novon
Tovar,0.1993.LasGramfneas(Poaceae)delPeru.Ruizia
13:
1-480.
Tsvelev,N.N.1976.(brassesoftheSovietUnion,part1:
298—343.
Nauka
Publishers.
Leningrad.
[English
trans¬
lationpublishedbyAmerindPublishing,NewDelhi,
1983.|
Wolff,R.,L.Abbott&S.Pistorale.1996.Reproductive
behaviorofBromuscatharticusVabl(Cebadillacriolla)
innaturalandcultivatedpopulations.J.Genet.Breed.
50:
121-128.
Zuloaga,F.()..E.G.Nicora,Z.E.RugolodeAgrasar,0.
Morrone.J.Pensiero&A.M.Cialdella.1994.Calalogo
delaFamiliaPoaceaeenlaRepuhlicaArgentina.Monogr.Syst.Bot.MissouriRot.Gard.47:1-178.
Amyris
oblanceolata
(Rutaceae),
a
New
Species
from
Nicaragua
Amy Pool
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Amyris oblanceolata, a new species
fromnorthwesternNicaragua,isdescribed,andits
relationships to other Central American species
withopposite,trifoliolateleavesarediscussed.
The genus Amyris consists of approximately 40
speciesdistributedintheWestIndiesandfromFlor¬
idaandTexasthroughCentralAmericatoPernand
Venezuela.Theyareunarmedtreesorshrubswiththe
leavesopposite,oralternate,and1-11-foliolate;the
flowerssmall,generallyperfectandactinomorphic
with3—5petalsand6—10freestamens;andthefruits
one-seededdrupes.SpeciesfoundinSouthAmerica
were revised by Gereau (1991). A new species of
AmyrisfromCentralAmericawasencounteredduring
preparationofatreatmentofRutaceaefortheFlora
deNicaraguaandisheredescribed.
Antyris oblanceolata A. Pool, sp. nov. TYPE:
Nicaragua. Depto. de Estelf: Estelf, elevation
1600 ft., 21 Feb. 1957 (buds and imm. fr), J.
B. Salas & B. W. Taylor 2267 (holotype, EAP).
Frutexvelarborparva.Foliaopposita,trifoliolata,foliolis
oblanceolatis
vel
ellipticis,
apice
acutis
vel
rotundatis,
retusis,
eoriaceis.
Inflorescentia
multiflora,
pseudoterminalis,
paniculata,
rarnis
pubescentibus,
floribus
sessilibus.
Shrub or small tree, 2-7 m tall, the young
branchespuberulent.Leavesoppositetosuboppo¬
site, trifoliolate; leaflets oblanceolate to elliptic,
acute to rounded at the apex, with retuse tip, cuneate at base, 5—9 cm long, 2—3.5 cm wide with
terminal leaflet larger than laterals, entire, coria¬
ceous, glabrous except midrib puberulent above,
withnumerouspellucidpunctations,theveinsre¬
ticulate, conspicuous and raised on both surfaces,
lateral leaflets sessile or on petiolules to 2 mm, the
petioluleofterminalleaflet4—12mm;petiole1.5—
3.5 cm, not winged, puberulent. Inflorescence
many-flowered,pseudo-terminal,paniculate,the
branchesdenselypuberulent,theflowerssessilein
denseclusters,thebudsglobose;calyx4-lobed,the
lobes triangular, acute at apex; petals 4, glabrous;
stamens7or8intwoseries,thefilamentsglabrous;
gynophore present; ovary with a few hairs at base.
SpeciesofAmyrisfoundinCentralAmericasouth
ofMexicowithoppositeorsubopposite,trifoliolate(or
occasionallytrifoliolate)leavesare:A.oblanceolata,
A.guatemalensisLundell,membersoftheA.elemifera
speciescomplex,andA.brenesiiStandley.
Amyris oblanceolata is most similar to A. guate¬
malensisLundell,knownonlyfromthetype,afruit¬
ingcollectionfromAltaVerapaz,Guatemala.Amy¬
ris guatemalensis is similar to A. oblanceolata in
havingsubcoriaceousleaflets,anunusualcondition
in this genus, and short pedicels. Amyris guate¬
malensis differs from A. oblanceolata in having
leavesthataregenerally5-foliolatewiththelateral
leaflets distinctly petiolulate and the tertiary veins
neitherraisednorconspicuousontheadaxialsur¬
face.TheleafletsofAmyrisguatemalensisaregen¬
erallyellipticwithacuminateapicesbutsometimes
are(likethoseofthenewspecies)oblanceolatewith
roundedandretuseapices.
MembersoftheAmyriselemiferaspeciescomplex
(including: A. elemifera L., A. balsamifera L., and A.
vestitaLundell)differfromA.oblanceolatainhaving
membranoustochartaceousleafletsanddistinctly
pedicellateflowers.Inaddition,theirleafletsaregen¬
erallysuborbiculartolanceolatewithacutetoacu¬
minateapices.Theapplicationofnamesandspecies
circumscription within this group is discussed by
Lundell(1960)andGereau(1991).
AmyrisbrenesiiStandley(includingA.costaricensis Standley), endemic to Costa Rica, is easily dis¬
tinguishedfromotherspeciesofAmyrisfromCen¬
tral America by its large lateral leaflets, 14-26 cm
long, and its long petioles, 9-24 cm long.
Paratypes.NICARAGUA.Jinotega:SierraWofJinotega,alongroadtoCerrodelaCruz,elevation1050—
1350m,chieflyindensewetmixedlowforest.27June
1947(sterile),P.C.Standley10177(EAP).Matagalpa:
ElEden,caminoviejoaJinotega,12°58'N,85°58'W,el¬
evation856m.1Feb.1984(buds),P.PMoreno22909
(HNMN,
MO).
Acknowledgments. I thank W. D. Stevens, Roy
Gereau, and Jacquelyn Kallunki for advice and en¬
couragement,andRoyGereauforassistanceinpre¬
paringtheLatindescription.
Literature
Cited
Gereau, R. E. 1991. El genero Amyris (Rutaceae) en
AmericadelSur.condosespeciesnuevasdelaAma¬
zonia
occidental.
Candollea
46:
227—235.
Lundell,C.L.1960.PlantaeMayanae—I.Notesoncol¬
lectionsfromthelowlandsofGuatemala.Wrightia2:
49-63.
Novon 8: 61. 1998.
Notes
on
Central
American
Scutellaria
(Lamiaceae)
Amy Pool
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Scutellaria ebracteata, in section Cardinalis Epling, from Nicaragua, and S. tenuipetiolata in section Uliginosae Epling, from Costa Rica,
are described, and identification keys to section
Cardinalis as found in Honduras, Nicaragua, and
Costa Rica and section Uliginosae as found in Nic¬
aragua and Costa Rica are provided. The identity
ofScutellarianicaraguensisSesse&Mocinoisdis¬
cussed,andthetypelocalityofS.purpurascensvar.
heterophyllaBenthamisclarifiedanditsaffinityto
othertaxaisconsidered.
In the course of preparing treatments of Lami¬
aceae for the floras of Nicaragua and Costa Rica,
specimensrepresentingtwonewspeciesofScutel¬
laria were found, significant range extensions of
otherspecieswereencountered,andtwopoorlyun¬
derstoodnameswereclarified.
Scutellaria ebracteata A. Pool, sp. nov. TYPE:
Nicaragua.Depto.deMatagalpa:Macizosde
PenasBlancas,SEside,drainageofQuebrada
El Quebradon, slopes N and W of Hacienda
San Martin, ca. 13°14-15'N, 85°38-39'W,
1000-1400 m, 18-20 Jan. 1982 (fl), W. D. Ste¬
vens with P. Moreno & T. Elmquist 21115 (holotype, MO; isotype, HNMN not seen). Eigure1.
SpeciesnovaScutellariaeglabraeK.Leonardsimilis
sedabeacorollapurpureamonatc,tubicumgalea2.7—
3.1mmlonga,tuboangusteinfundibular!nonsaccatodiffert.
Weakshrub(orherb),1—1.5m,youngbranches
withdensesmall,curvinghairs.Leaveswide-ellip¬
tic,elliptic-oblong,orslightlypandurate,short-acu¬
minate to obtuse at apex, rounded to cordate and
often slightly asymmetrical at base, 7.7—13.5 cm
long,2.6—7cmwide,marginsub-entiretoserrate,
adaxialsurfaceglabrous,abaxialsurfaceglabrous
or with minute hairs on primary and secondary
veins; medial petioles 1.5—5 cm long, shorter to
longer than adjacent internodes. Inflorescence a
terminalraceme,4.5—10cmlong;flowers15—25,
looselyspiraled,pendulous;bractsdeciduousprior
to anthesis, pedicel 2—3 mm long; calyx 3.5—4.5
mmlong,minutelypuberulentorglabrous,seutelNovon 8: 62-66. 1998.
lum 1.2—2 mm high; corolla reddish-purple, tube
with galea 2.7-3.1 cm long, tube narrow-funnelform, arcuate, noil-saccate, 1-2 mm wide at base
to 5—7 mm wide at apex, lower lip 3—5 mm long.
Fruiting calyx with lower lip to 5 mm long, scutellum to 4 mm high; nutlets (immature?) tan, nearly
smooth.
Paratypes.NICARAGUA.Jinotega:MacizosdePen¬
asBlancas,alongtrailbetweenfincaofSocorroMejiaand
tineaofLuisMan/.anares,ea.13°16—17'N,85°40-41’W,
1350-1650m,cloudforestongentletosteepslopes,14
Jan.1079(flbuds),1LI).Stevens11339(HNMNnotseen,
MO); Fila Piedra Pelona, al S del Cerro kilambe,
I3°34'N,85°41'W.15(X)—1<565ni,bosqueenano,28Mar.
1981(youngfr),l‘.Moreno7793(HNMNnotseen,MO);
San Ramon, lado E de las faldas del Cerro kilambe.
13°34'N.85°40'W,800-900m,24Mar.1981(fl).P.Mo¬
reno7409(HNMNnotseen,MO);Cerrokilambe.falda
l,delPicoPiedraPelona.I3°34'N,85°40'W,1300-1-MX)
m.bosqueshumedos,28Mar.1981(fl),P.Moreno/784
(HNMN not seen. MO); kilambe, Cerro San Pedro,
I3°36'N,85°39'W.600-800m,25Mar.1981(flbuds,
youngfr),P.Moreno7535(HNMNnotseen.MO);Florde
Liz,alVidelCerrokilambe,I3°35'N.85°40’W,700—900
m.24Mar.1981(fl.fr).PMoreno7430(MO).Zelaya:
CerroSaslayaaunos25kmaloestedeSiuna,1200m,
Oct.1977(fl),A../.Ferguson2(MO);CerroSaslaya,20
kmWofSiuna,1100—1400m,alongeasternridgeof
mountain,cloudforest.5May1977(flbuds).I).Neill
1832
(MO).
Epling (1942) treated all Scutellaria species
withlarge(tubewithgalea16—60mm)redflowers,
known at that time from Central America south of
Mexico, as members of section Cardinalis Epling.
Patou (1990) treated the same species as Scutel¬
laria sect. Scutellaria species-groups “speciosa”
(in part) and “costaricana.” These species are all
large herbs or weak shrubs, often straggling, with
leavesglabrousorsparinglyhirsuteabove,andra¬
cemes with flowers spirally arranged in the axils
of the deciduous to long persistent bracts, the co¬
rollas large and showy, orange-red, scarlet or
bright red, and often arcuate. Epling recognized
thefivespeciestreatedhere,except.S’,ebracteata.
However, the species are difficult to separate and
havebeentreatedinvariouswaysinmodernCen¬
tralAmericanfloras.FloraofGuatemala(Standley
& Williams, 1973) treated .S’, isocheila Donnell
Smith and S. glabra E. Leonard as synonyms of
.S’, longifolia Bentham. Flora of Costa Rica (Stand-
Volume 8, Number 1
1998
Pool
Central American Scutellaria
63
Figure1.ScutellariaebracteataA.Pool.—A.Habit.—B.Flower.—C.Fruitingcalyx.
ley, 1938) recognized S. glabra but treated S. isocheila as a synonym of 5. longifolia. To clarify my
concepts, a key to the species of S. sect. Cardinales found in Honduras, Nicaragua, and Costa
Rica is provided. Scutellaria longifolia, as here
circumscribed,isrestrictedtosouthwesternMex¬
ico and western Guatemala. It is recognized by
having lanceolate leaves with euneate (to rarely
roundedandshortlydecurrent)bases,andflowers
that are ascending to spreading and spirally ar¬
rangedintheaxilsofcaducousbracts,withorange
corollas of medium size (tube with galea 25—32
mm) with straight (to slightly sigmoid) tubes that
are not saccate at the base. Scutellaria hookeri
Epling is now known from Costa Rica, based on
L. 0. Williams et al. 28585 (F) collected in the
Cordillera de Talamanca. Scutellaria hookeri,
known then only from Peru, was treated by Epling
as a member of section Coccineae Epling ( Scutel¬
laria sect. Scutellaria species-group “speciosa”
(in part) of Paton, 1990). It differs from members
of section Cardinales only in having pilose to vil-
64
Novon
lous hairs on the abaxial leaf surface; S. hooheri
is treated here as a member of section Cardinalis
and included in the key.
bracteisestipitatis,corollapallidiore,tuboalbido,labiis
azureis
ad
violaceis
differt.
Kky to in k Species ok Sc in: ij aria Sect. Cariuxai.es in
Honduras,Nicaragua,andCostaRica
la.Inflorescencebractsdeciduousbeforeoratanthesis;corollatubessaccateabovebaseornot;
leaf
bases
cordate,
subcordate,
or
rounded.
2a.Corollatubewithgalea46-60mm,orangeredwithyellowororangelobes;flowers
erect,
congested,
anil
corymbiform;
known
from Costa Rica and Panama .
. S. costaricana H. Wendland
(including
.S’,
argentato
E.
Eeonard)
2b.Corollatubewithgalea16-31mm,darkred
orreddishpurple;(lowersspreadingtopen¬
dent,
loosely
spiraling.
3a.Corollareddishpurple,lubewithgalea
27-31mm,tubenotsaccate;endemic
to Nicaragua ..S. ebracteata A. Pool
3b.Corolladarkred,tubewithgalea16-23
mm,tubesaccateononesideat3—4
mmabovebase;knownfromCostaRica
and Panama. 5. glabra E. Leonard
lb.Bractspersistentintoandbeyondfruiting;corol¬
latubesnotsaccateabovebase;leafbasesat¬
tenuate
to
rounded
or
cordate.
4a.Inflorescencewithoutglandular-capitate
hairs;corollatubewithgalea20—34mm,
bright
pink;
leaves
with
attenuate
bases,
gla¬
brous above; endemic to Costa Rica.
.,S.
isocheila
Donnell
Smith
4b.
Inflorescence
with
glandular-capitate
hairs;
corollatubewithgalea19-40mm,roseor
orange-redtobrightred;leaveswithround¬
edtocordatebases,scatteredhirsuteabove
(at
least
along
margin).
5a.Corollatubewithgalea19—26mm,rose
tored;calyxlong-villous;leaveswith
subcordatetocordatebasesandwith
long(toca.1mm)curlyorwavyhairs
onsurfacebelow;knownfromCosta
Rica, Colombia, and Peru .
.
S.
hooheri
Epling
5b.Corollatubewithgalea32—40mm,or¬
ange-redtobrightred;calyxshort-his¬
pid;leaveswithroundedbasesandwith
shortstraighthairsonsurfacebelow;
knownfromGuatemala,Belize,and
Honduras . S. inflata Epling
Scutellaria tenuipetiolata A. Pool, sp. nov.
TYPE: Costa Rica. LimAn Province: CantAn de
Talamanca Amubri, camino entre Amubri y
Soki, siguiendo el Rfo Nabri hacia Alto Soki,
9°29'50"N, 82°59'10"W, 150 m, 1 July 1989
(fl,youngfr),GerardoHerrera3101(holotype,
MO; isotypes, CR not seen, F). Figure 2.
Species
nova
Scutellariae
purpurascenle
Swartz
similis
sedabeapetiolismedianisinternodiislongioribus,foliis
basireveracordata,sinuusqueadca.3mmprofundo.
Small perennial herb with thickened fascicled
roots, usually with several stems arising from a
small caudex, stems densely puberulent. Leaves
ovate, bluntly acute at apex, shallowly cordate at
base with sinus to 3 mm deep, 3.5—5.6 cm long,
2-4 cm wide, margin crenulate (sinuate), adaxial
surface nearly glabrous with scattered, coarse,
jointed hairs, abaxial surface puberulent on pri¬
maryandsecondaryveins;medialpetioleslonger
than adjacent internodes, 2.5-4 cm long, less than
1mmwide.Inflorescenceaterminalraceme,3.5—
5.5cmlong,4—12-flowered,flowersspiraledtosub¬
opposite at a node; rachis, bracts, pedicels, and
calyces puberulent; bract persistent, elliptic, 2-3
mm long, exstipitate; pedicel 1-2.5 mm long; calyx
2.5—3mmlong,scutellum1.5—2mmhigh;corolla
with white tube and blue to purple lips, tube with
galea 14—16 mm long, tube 1 mm wide at base to
3 mm wide at apex, lower lip 3-4 mm long. Fruiting
calyx with lower lip to 4 mm long, scutellum to 5
mmhigh;nutletsbrown,smoothtosub-papillate.
Scutellariatenuipetiolataisfoundinwet,weedy
areas.
Paratypes.COSTARICA.Linion:CordilleradeTal¬
amanca,CantondeMatina,200maguasabajodelaconfluenciadeQuebradaCafiabralconRfoBarbilla,margen
derecha,siguiendoelcursodelaQuebradaCamagre,
10°00'10"N,83°25'30"W,l(M)m,5Nov.1988(fl).G.Her¬
rera2283(CRnotseen,F,MO);neartheRioCatarata
(RfoSandBox)inthehillsbetweenBriBriontheRfo
SixaolaandtheCaribbeancoastalplain,9°37’N.82°49'W,
50—1(M)m.28—29Nov.1975(flbud,fr).R.Baker&IL
Burger39(F.MO);arribadelacataratadelRfoSandBox,
Talamanca,200m,17July1982(fr),J.Gomez-Laurito
8732(F);entreLinionetMarfa,Sep.1899(frandfl),//.
Pillier16013(knotseen.US).
All the species of Scutellaria with small (tube
with galea 11-18 mm) bluish or purplish flowers
found in Nicaragua and Costa Rica are treated by
Epling (1942) as members of section Uliginosae
Epling(Scutellariasect.Scutellaria“species-group
uliginosa” of Paton, 1990). They are all small herbs
(generally less than 50 cm tall), with fascicledthiekened roots and several stems arising from a
small caudex or slightly elongated rhizome. The
leaves are thinly hirsute above, with stout-jointed
hairs, and puberulent at least on the veins below.
The flowers are spiraled to sub-verticillate in the
axils of bracts on a short raceme with the corollas
small, purple or blue, or white with blue or purple
lips.
EplingrecognizedtwospeciesofScutellariasect.
Uliginosae in Costa Rica. He included the newly
described5.tenuipetiolata,asrepresentedbyPit-
Volume 8, Number 1
1998
Pool
Central American Scutellaria
65
5mm
Figure2.ScutellariatenuipetiolataA.Pool.—A.Habit.—B.Adaxialleafsurface.—C.Abaxialleafsurface.—I).
Flower,frontview—E.Flower,lateralview.—F.Fruitingcalyx,lateralview.—G.Fruitingcalyx,backview.—H.
Fruitingcalyxwithnutlets.
66
Novon
tier16013,inhisconceptofS.purpurascensSwartz.
This group was not known from Nicaragua. Four
species are now known from Costa Rica, while one,
S. galerita Epling, has been found in Nicaragua.
EplingtreatedScutellariapurpurascensvar.heterophyllaBenthamasasynonymof.S',guatemalensis E. Leonard, but it is here maintained in S. pur¬
purascens. However, insufficient material of S.
purpurascensoveritsentirerangehasbeenexam¬
ined to determine whether a varietal distinction is
warranted.ScutellariaguatemalensisdiffersfromS.
purpurascensinitsexstipitatebractsanddense,rel¬
atively long hairs on stems, leaf abaxial surface,
and calyx. Il is not known south of Guatemala.
Epling may have been misled into believing that
theholotypeof.S.purpurascensvar.heterophylla,
Friedrichsthals.n.(K),wasfrommodern-dayGua¬
temala.MostFriedrichstahlspecimensarelabeled
as from Guatemala; only the original set retained
atWindicateactualcollectionlocalityandnumber.
Examinationoftheholotype,theFriedrichsthalcol¬
lections at W, and a photocopy of Friedrichsthal’s
fieldnotessuggeststhattheholotypeofS.purpur¬
ascensvar.heterophyllaisaduplicateofFriedrichs¬
thal 1299 (W), collected at San Jos£, Costa Rica.
ScutellariaorichalceaDonnellSmithwasplaced
by Epling in Scutellaria sect. Pallidiflorae Epling,
based on the color of the corolla, white or white
with lips yellowish green or bluish to purplish tint¬
ed; it is here treated as a member of section Uliginosae.Specimensof.S’,orichalceafromthenorth
of Nicaragua are found from 150 to 1000 m and
have corollas 19-24 mm long, that are white or
whitewithyellow-greenlips.CollectionsfromCosta
Rica are found at and above 1400 m and have
slightly smaller corollas, 15-19 mm long, that are
white with bluish or purplish lips or tints. If further
investigationconfirmstheseobservations,recogni¬
tion of the Costa Rican entity as a distinct subspe¬
ciesmightberecommended.
2b.
Inflorescence
without
glandular-capitate
hairs;
leaves
shallowly
cordate
(sinus
1.5—3
mm); petioles 0.3—0.75 mm wide.
.
S.
tenuipetiolata
A.
Pool
lb.heaveswithbasescuneatetotruncate;medial
petioles
shorter
than
adjacent
internodes.
3a.
Corolla
red-purple
(blue);
racemes
with
flow¬
ersinmany-floweredverticels(especially
denseatapex);bractsstipitate;leavesovate,
less than two times as long as wide.
.
.S’.
purpurascens
Swartz
3b.Corollawhitewithbluish,purplish,oryel¬
low-greenishlipsortints;racemeswithflow¬
ersspiralingoropposite;bractsexstipitate;
leaveslanceolate,twoormoretimesaslong
as wide.5. orichalcea Donnell Smith
Kky to the Srn.it s ok Set teijaria Sect. Ui.h;i\osae in
NicaraguaandCostaRica
la.Leaveswithbasescordate;medialpetiolesgen¬
erally
longer
than
adjacent
internodes.
2a.
Inflorescence
with
some
glandular-capitate
hairs;leavesdeeplycordate(sinus5—10
mm); petioles 1.2—2 mm wide.
.
S.
galerita
Epling
NeitherLeonard(1927)norEpling(1942)treat¬
ed the name Scutellaria nicaraguensis, described
by Sesse and Moeino in Flora Mexicana (1894), as
from“Legione”(equivalenttoLe6n,Nicaragua,ac¬
cording to McVaugh, 1977). No illustrations or
specimens bearing this name have been located.
The description is more compatible with Ocirnum
campechianum P. Miller than with any species of
ScutellariaknownfromCentralAmericaorexpect¬
ed to be found in the dry forests in the area of Le6n;
Ithereforesuggestthatsynonymy.
Acknowledgments. I thank W. D. Stevens and
Michael Grayum for advice and encouragement,
Roy Gereau for reviewing the Latin diagnosis, and
John Myers for providing the illustrations. I also
thank the curators of BM, F, K, OXF, and US for
specimen loans and the curators at F, MA, and the
Hunt Institute for Botanical Documentation for
searchingformaterialofScutellarianicaraguensis.
Literature
Cited
Epling,C.1942.TheAmericanspeciesofScutellaria.
Univ.Calif.Publ.But.20:1-145.
Leonard,E.1927.TheNorthAmericanspeciesofScutel¬
laria.Contr.U.S.Natl.Herb.22:703-748.
McVaugh,R.1977.BotanicalresultsoftheSesse&Mo¬
eino
expedition
(1787—1803).
I.
Summary
of
excursions
andtravels.Contr.Univ.MichiganHerb.11:97—195.
Raton.A.1990.AglobaltaxonomicinvestigationofScu¬
tellaria(Labiatae).kewBull.45:399—450.
Sesse,M.&J.M.Moeino.1894.FloraMexicana,2nded.
Oficina
tipografiea
de
la
Secretarfa
de
Fomento,
Mexico.
Standley,P.C.1938.Labiatae.In:FloraofCostaRica.
Publ.FieldMils.Nat.Hist..Bot.Ser.18(3):1015—1035.
- & L. O. W illiams. 1973. Labiatae. In: L. 0. W il¬
liamsetal..FloraofGuatemala.Fieldiana,Bot.24(9):
237-317.
Stenopadus
andicola
Sp.
Generic
Nov.
Record
(Asteraceae:
for
Mutisieae),
a
New
Ecuador
John F. Pruski
United States National Herbarium, Department of Botany, MRC-166, Smithsonian Institution,
Washington, D.C. 20560-0166, U.S.A.
ABSTRACT. A new species, Stenopadus andicola
(Asteraceae: Mutisieae), is described from the
Cordillera del Condor, Ecuador. It is the first
member of the genus reported outside the Guayana Highland and is a new generic record for
Ecuador. Stenopadus andicola is placed in section
Stenopadus.
Stenopadus andicola Pruski, sp. nov. TYPE: Ec¬
uador.Zamora-Chinchipe:CantonNangaritza,
Valle del Rfo Nangaritza, Miazi, bosque sobre
pendientes fuertes de roca caliza o de pizarra,
bosqueprimario,muydenso,4°18'S,78°40'W,
1200 m, 10 Dec. 1990 (fr), IE Palacios 6712
(holotype, US; isotypes, MO, QCNE not seen).
Figure1.
The genus Stenopadus S. F. Blake (Asteraceae:
Mutisieae) contains 15 species, 14 of these occur¬
ring in the Guayana region of Brazil, Colombia,
Guyana, and Venezuela (Pruski, 1991 [1993]; Pru¬
ski, 1997), principally on sandstone. No species of
Stenopadus were listed as occurring in the Andes
in treatments of the Mutisieae for Colombia (DfazPiedrahita & Velez-Nauer, 1993), Ecuador (Hai¬
ling, 1991), Peru (Ferreyra, 1995), or Venezuela
(Aristeguieta, 1964). The description of a fifteenth
species, 5. andicola, marks the first report of the
genusfromoutsidetheGuayanaregionandthefirst
reportofthegenusinEcuadorandtheAndes.Sten¬
opadus thus joins Gongylolepis R. H. Schomburgk
asGuayana-centeredMutisieaegenerawithasole
species occurring in the Andes. In Pruski (1997:
365),theoccasionwastakentomention.S’,andicola
in passing. The new species is named prior to its
use in the forthcoming Catalogue of the Vascular
Plants of Ecuador, coordinated by P. M. Jprgensen
andS.Leon.
The Mutisieae contain some of the most primi¬
tive species of Asteraceae, and the “ancestral asteraceous plant” illustrated in Bremer (1994) is
Stenopadus- like. Important primitive features
found in the Guayana-centered Mutisieae include
arborescent habit, thick fleshy leaves (taken as a
mechanical deterrent, in light of the weak protec¬
tive chemistry of the group), large homogamous
capitula with florets that are bird-pollinated,
sometimes paleate receptacles, short rounded
style branches with a single stigmatic surface and
without collecting hairs, anthers without a distinct
apicalappendage,andsmoothprolatepollen(Pru¬
ski, 1991 [1993]).
Arbolusque15inalta:caulessericeivelglabrati;folia
alterna,petiolata;petioli1.5—4cmlongitenuinon-amplexicauli;foliorumlaminacoriaceaoblanceolatavel
obovata8—24cmlonga2—7.5cmlataintegrapinnativenia
sericeavelglabrata;capitulasolitariasessiliavelbrevipedunculataepaleaceahomogama.flosculisca.25hermaphroditis;
involucrum
eylindricum
vel
anguste
campanulatum 3.9—4.5 cm longum ca. 2.2 cm latum 6—
7-seriatum;phyllariaimbricatasericeavelapicalis
glabrata;receptaculumplanumepaleaceumca.1cm
diam.;corollaeignotae;achaeniacylindricaca.10cm
longa
glabra:
pappi
setae
numerosae
multiseriatae
usque
20
mm
longae.
Trees to 15 m tall, to 20 cm diam.; stems subterete,grooved,sericeouswhenyoungtoglabrate,
leafy distally, leafless proximally, internodes to 3
cm long. Leaves simple, alternate, often apically
clustered,petiolate;petiole1.5—4cmlong,thinand
non-clasping;bladerigid-coriaceous,oblanceolate
to obovate, 8—24 cm long, 2—7.5 cm wide, apically
broadlyacutetorounded,narrowlycuneatebasally,
marginsentire,thickened,somewhatrevolute,ve¬
nationpinnate,reticulateorthird-orderveinssome¬
timesobscure,theupperbladesurfacedarkgreen,
midrib commonly sericeous to puberulent, upper
surfaceotherwisepuberulentwhenyoungtogla¬
brous, the lower blade surface pale green, midrib
sericeoustoweaklyso,lowersurfaceotherwisese¬
riceous when young to nearly glabrous. Capitula
solitary, terminal, sessile to shortly pedunculate,
homogamous,ca.25-flowered,floretsbisexual;pe¬
duncle0—0.5cmlongandnotmuchexsertedabove
thebasesofthesubtendingleaves,thesubtending
leavesdeciduousanduppernodeortwowithelon¬
gatingaxillary'branch(es)whencapitulainfruit,the
peduncle then leafless with capitula 5 cm above
axillary'branch(es)anduppermostleaves;involucre
cylindrical to narrowly campanulate, 3.9^4.5 cm
Novon 8: 67-69. 1998.
Novon
68
MISSOURI
BOTANICAL
GARDEN
HERBARIUM
v? ji’npiir.j
(Walter
Palacios
6712
)
ASTERACEAE
Stenopadus
colonbianur:
Cuatrec.
'■
Stayerm.
Det.H.Pobir.sonr,;s)<1991
MISSOURI
BOTANICAL
GARDEN
HERBAPIC"
XO>
ECU/•POP
ASTERACEAE
Stwi^p-Ktks*.<WAI
A
Jr.
i"f>
ZAMOPA-CH
INCH
Hangarltza
IRE
:
Car,tor,
ValleoelFloHangaritza.Piazi.
Eosaue
sobre
pendientes
fuerten
de
roca
calizaooepizarra.Bosquepnrano.
. rubier tos da
78 * 40 'W 04 ‘ 18 'S 1200 r.
Arbolde1iisdealtura.Panta:.jovene
patdas.
10die1890
Walter
Palacios
6712
EEPBAPIG
NACIONAL
DEL
ECUADOR
'CONE)
P.ISSOCPI
BOTANICAL
GARDEN
HERBARIUM
KO)
f
Figure1.Isotype(Palacios6712,MO)ofStenopadusandicolaPruski.
Volume 8, Number 1
1998
Pruski
Stenopadus andicola from Ecuador
long, ca. 2.2 cm wide, 6—7-seriate; phyllaries ca.
40,imbricate,graduated,tightlyappressedandrig¬
idlyerect,coriaceous,sericeousorapexsometimes
glabrate,entire,theouterphyllarieskeeled,trian¬
gular-ovate,0.5—1cmlong,0.4—0.6cmbroad,apex
acuteorobtuse,theinnerphyllariesweaklykeeled,
elliptic-lanceolatetolanceolate,ca.3cmlong,3.5—
5 mm wide, apex narrowly acute; receptacle flat,
epaleaceous, ca. 1 cm diam. Corollas unknown.
Cypselas(achenes)nearlycylindrical,mostly5-angled, ca. 10 mm long, brown, glabrous; pappus se¬
taenumerous,several-seriate,linear,stramineous,
to 20 mm long, about twice as long as the cypselas.
considered to be a member of section Connellia
sensu Maguire et al. (1957).
By thinly petiolate, large, occasionally abaxially
sericeousleaves,Stenopadusandicolaappearsto
be most closely related to S. colombianus Cuatrecasas&Steyermark(aColombianendemic,andthe
nearest geographic relative of 5. andicola) and the
widespread S. talaumifolius S. F. Blake, both of
section Stenopadus sensu Maguire et al. (1957).
However, the leaves of the new species are weakly
(vs. strongly) reticulate and commonly (vs. occa¬
sionally) abaxially sericeous, thus differing from
bothS.colombianusandS.talaumifolius.Thephyl¬
lariesofS.andicolaaresericeous(ortheinnermost
sometimesapieallyglabrate)andkeeled,whichfur¬
ther distinguishes it from S. colombianus and S.
talaumifolius,bothofwhichhavenon-keeled,gla¬
brousphyllaries,orrarelywiththeoutermostphyl¬
laries sericeous. The leaves of S. cucullatus Ma¬
guire (sect. Stenopadus ) resemble those of S.
andicola, but S. cucullatus differs from the new 7
species by its weakly seriate, apieally obtuse to
rounded phyllaries. The new species is presumed
tobeamemberofsectionStenopadus.
Distributionandecology.Stenopadusandicola
is known only from two collections in late fruit in
October and December. The specimens in QCNE
were not seen, but Walter Palacios (pers. comm.)
says that they too are in late fruit. These collections
were made in cloud forests from 1100 to 1200 m
elevation on the Cordillera del Condor in the prov¬
inceofZamora-Chinchipe,Ecuador.TheCordillera
delCondorislargelysandstone,butS.andicolahas
been collected only in quartzite or slate areas. The
CordilleradelCondorisconsideredamongtheold¬
est geologic formations in Ecuador (Peter Jorgen¬
sen,pers.comm.),andisimportantbiogeographically; it is home to several other Guayana disjuncts
[e.g., Everardia montana Ridley ex Thurn (Cyperaeeae),PaepalanthusdichotomiesKlotzsehexKornicke(Eriocaulaceae),PeramaAublet(Rubiaceae),
Pterozonium brevifrons (A. C. Smith) Lellinger
(Pteridaceae),andPterozoniumreniforme(Martius)
Fee(Pteridaceae)](JohnWurdack,pers.comm.).
Thisnewspeciesisknownonlyfromfruitingma¬
terial, but is referred to actinomorphic-flowered
Stenopadusbyitsarborescenthabitwithunarmed
stems,largecoriaceousleaveswiththin,non-clasp¬
ing petioles, non-plumose pappus, large capitula,
and keeled phyllaries. The corollas of species of
Stenopadus are mostly red, and the corolla lobes
are flexuous or coiled and about as long as the co¬
rolla tubes. By its keeled phyllaries, S. andicola
resembles5.chimantensisMaguire,Steyermark&
Wurdack, S. connellii N. E. Brown, and .S', sericeus
Maguire & Aristeguieta. The three latter species
are placed in section Connellia Maguire & Wur¬
dack and are confined to the tepuis of the eastern
GuayanaHighland.Stenopadussericeushasretic¬
ulate, abaxially sericeous leaves, further resem¬
bling 5. andicola. However, the leaves of S. chi¬
mantensis, S. connellii, and 5. sericeus are stoutly
petiolate, thus these three species are readily dis¬
tinguished from S. andicola. The new species is not
69
Paratype.ECUADOR.Zamora-Chinchipe:Canton
Nangaritza,
Detrds
del
Campamento
Militarde
Miazi
(oeste).
bosque
nublado
con
arboles
cubiertos
completamente
dehepaticasymusgos,estratodelbosque15mdealtura,
Dystropept.areniseacuarzosameteorizada,4°16'S,
78°42'W, 1100 rn. 21 Oct. 1991 (fr), W. Palacios et al.
8551(MO.QCNEnotseen).
Acknowledgments.IthankPeterJdrgensen,Har¬
oldRobinson,JohnWurdack,andananonymousre¬
viewerforhelpfuldiscussionandcomments.
Literature
Cited
Aristeguieta,L.1964.Compositae.In:T.Lasser(editor),
El.Venezuela10:1-941.
Bremer,K.1994.Asteraceae:Cladistics&Classification.
Timber
Press,
Portland,
Oregon.
Dfaz-Piedrahita,S.&C.Velez-Nauer.1993.Revisionde
lastribusBarnadesieaeyMutisieae(Asteraceae)para
laFloradeColombia.Monogr.Jard.Bot.Jos6Celestino
VIntis 1: xi + 1-162.
Ferreyra,R.1995.FamilyAsteraceae:PartVI[TribeMu¬
tisieae].In:.1.F.Macbride&Collaborators,Floraof
Peru.Fieldiana,Bot.n.s.,35:v+1—101.
Marling,G.1991.Compositae—Mutisieae.In:G.Marling
&L.Andersson(editors).FloraofEcuador42:1—105.
Maguire,B.,J.J.Wurdack&Collaborators.1957.The
botanyoftheGuayanaHighland—PartII.Mem.New
YorkBot.Card.9:235-392.
Pruski.J.F.1991[1993].CompositaeoftheGuayana
Highland—V.TheMutisieaeoftheLostWorldofBrazil.
Colombia,andGuyana.Bol.MuseuParaense,s6r.Bot.
7:
335-392.
-. 1997. Asteraceae. Pp. 177—393 in J. A. Steyer¬
mark.P.F.Berry&B.k.Holst(editors),Floraofthe
VenezuelanGuayana,Vol.3.MissouriBotanicalGar¬
den,St.Louis.
New
Names
in
Chinese
Apiaceae
PU Fa-ting
Chengdu Institute of Biology, Chinese Academy of Sciences,
Chengdu, Sichuan 610041, People’s Republic of China
ABSTRACT. A new name in Peucedanum, a new
varietyofOstericumscaberulum,andfournewcom¬
binationsinMeeboldia,Hydrocotyle,andOenanthe
areproposed.Theyare:PeucedanumfranchetiiC.
Y.Wu&Pu,Ostericumscaberulum(Franchet)Yuan
& Shan var. longiinvolucellatum C. Y. Wu & Pu,
Meeboldiayunnanensis(H.Wolff)Constance&Pu,
Hydrocotyle burmanica Kurz subsp. craibii (H. Eichler) C. Y. Wu & Pu, Oenanthe javanica (Blume)
DC. subsp. rosthornii (Diels) Pu, and Oenanthe
thomsonii C. B. Clarke subsp. stenophyllum (Boissieu)Pu.
Thenomenclatureofseveraltaxaofvariousgen¬
eraofApiaceae(Umbelliferae)needsadjustment.
Thenomenclaturalchangesarehereinproposedto
makethenamesavailablefortheforthcomingvol¬
ume 14 of the Flora of China.
Ostericum scaberulum (Franchet) Yuan & Shan
var. longiinvolucellatum C. Y. Wu & Pu, var.
nov. TYPE: China. Yunnan: Degen, Baima
Snow Mount, among shrubs, Sep. 1935, C. W.
Wang69408(holotype,KUN).
A
varietate
scaberulo
differ!
involucellorum
phyllis
nmbellulislongioribuselfatioribusea.1mmlatis.
Varietylongiinvolucellatumdiffersfromvariety
scaberuluminhavingbractletsca.1mmbroadand
longerthanthefruitingumbellets.Varietyscaber¬
ulum has bractlets ca. 0.5 mm broad and shorter
thanthepedicelsandfruitingumbellets.Thename
longiinvolucellatumwasfirstproposedasanomen
nuduminWang(1993).
Paratypes.CHINA.Yunnan:Zongdian,2780m,with
grassesandotherherbs,3Aug.1962,LikiangHot.Card.
100752(KUN).Degongmeadows,3300m,inshrubs,3
Sep.1959,K.M.Peng23510(KUN).
Peucedanum franchetii C. Y. Wu & Pu, nom.
nov.Replacedname:Peucedanumheterophyllum Franchet, Bull. Soc. Philom. Paris, ser. 8,
6: 141. 1894, not Visiani, Cat. Sem. Hort. Patav. 4. 1836. TYPE: China. Yunnan: Likiang,
3000 m, Oct. 1884, Delavay 192 (holotype, P).
PeucedanumfranchetiiisproposedtoreplaceP.
Novon 8: 70-71. 1998.
heterophyllumFranchet,whichisalaterhomonym
ofP.heterophyllumVisiani.
Meeboldia yunnanensis (H. Wolff) Constance &
Pu,comb.nov.Basionym:Sinodielsiayunna¬
nensis H. Wolff, Notizbl. Bot. Cart. BerlinDahlem 9: 278. 1925. TYPE: China. Yunnan:
Yunnan-fu,Cavalerie42?(holotype,B).
Meeboldia (Wolff, 1924) and Sinodielsia (Wolff,
1925)sharethesameprincipalgenericcharacters:
stronglydevelopedcalyxteeth,narrowlyovoidfruit
attenuate toward style and slightly constricted at
the commissure, 2 or 3 vittae in each furrow and 4
on the commissure, sulcate seed face, 3- or 4-pinnatisect leaves, 5—10 rays that are 4—5 cm long,
andaninvolucelof5—7linear-lanceolatebractlets.
Onthebasisoftheirmorphologicalsimilaritiesand
geographical distribution in the Himalayas and
southwesternChina,itisconcludedthatSinodielsia
shouldbereducedtosynonymyoftheearlierpub¬
lished
Meeboldia.
Hydrocotyle burmanica Kurz subsp. craibii (II.
Eiehler) C. Y. Wu & Pu, comb. nov. Basionym:
HydrocotylecraibiiH.Eiehler,FeddesRepert.
98: 146. 1987. New name for Hydrocotyle chinensis L. 1753, not H. shanii Boufford, Acta
Phytotax. Sin. 28: 331. 1990, superfluous
name. TYPE: China. Yunnan: Mengzi, woods,
8500 ft., A. Henry 10224 (holotype, K).
Subspeciescraibiidiffersfromsubspeciesbur¬
manica in having orbicular-reniform, 5—7-lobed
leaves.Bothsubspeciesshowasouthtonorthtrend
fromshallowlytodeeplylobedleaves.
Hydrocotyle craibii H. Eiehler and H. shanii
Boufford (see Eiehler, 1987; Boufford, 1990) were
bothproposedindependentlyasnewnamesfor//.
chinensis (Dunn ex R. H. Shan & Liou) Craib ex
Tardieu-Blot, which is a later homonym of //. chi¬
nensisL.(1753).
Oenanthe javanica (Blume) DC. subsp. rosthornii (Diels) Pu, stat. nov. Basionym: Oenanthe
rosthornii Diels, Bot. Jahrb. Syst. 29: 498.
1900. TYPE: China. Guizhou: Ping-fa, 21
Aug. 1902, J. Cavalerie 176 (neotype, E).
Volume 8, Number 1
1998
Pu
Chinese Apiaceae
Oenanthejavanicasubsp.rosthorniidiffersfrom
subspeciesjavanicabyitsunequalrays,lanceolate
involut'd braetlets, and ovoid fruit. In subspecies
javanica the rays are subequal, the involucel bractlets are linear, and the fruit is oblong.
has1-or2-pinnatehetermorphicleavesandlinear,
lanceolate,orrhomboid-ovatepinnae.
Oenanthe thomsonii C. B. Clarke subsp. stenophyllum (Boissieu) Pu, stat. nov. Basionym:
Oenanthe diel.sii Boissieu var. stenophylla
Boissieu, Bull. Acad. Int. Geogr. Bot. 16: 185.
1906. TYPE: China. Sichuan: Cheng kou, For¬
ges s.n. (holotype, P; isotype, K).
Because of its homomorphic, finely 3- or 4-pinnate leaves, linear pinnae, and subglobose fruits,
subspecies stenophyllum is more at home in Oe¬
nanthethomsoniithaninO.dielsii.Oenanthedielsii
71
Acknowledgment. I thank Ihsan Al-Shehbaz for
hishelpwiththemanuscript.
Literature
Cited
Boufford,D.E.1990.HydrocolyleshaniiBoufford,anew
nameforH.chinensisofauthors,notL.(Apiaceae).Acta
Phytotax.Sin.28:331-332.
Eichler,H.1987.Nomenclaturalandbibliographicalsur¬
veyofHydrocolyleL.(Apiaceae).PartII.FetIdesRepert.98:145-196.
Viang.W.T.1993.VascularPlantsofHengduanMoun¬
tainsVol.1.SciencePress,Beijing.
Wolff,H.1924.Meeboldia
genus
,
novumumbelliferarum
Himalayicum.Repert.Sp.Nov.RegniVeg.19:313.
-. 1925. Neue Umbelliferen-Gattungen aus Ostasien.Notizbl.Bot.Cart.Berlin-Dahlem9:275—280.
Two
New
Species
of
Larnax
(Solanaceae)
from
Ecuador
Neil W. Sawyer
Department of Ecology and Evolutionary Biology, The University of Connecticut, Box U-43,
75 North Eagleville Road, Storrs, Connecticut 06269, U.S.A.
ABSTRACT.TwonewspeciesofLarnax(Solanaceae)
aredescribedandillustrated.Larnaxandersoniiisa
smallshrubwithunequal,geminateleavesandlong,
bifurcatehairs.Itoccursalongtheeasternslopesof
theEcuadorianAndes.TheflowerstructureofLan¬
dersoniiissimilartoanotherEcuadorianspecies,L
suffruticosa.Larnaxpsilophyta,ahigh-elevationspe¬
ciesendemictosouthernEcuador,isasmall-flow¬
ered,glabrousshrubwithindurate,fleshyleavesthat
isoftenconfusedwithDepreaglabra.
ThegenusLarnax(Miers)Hunziker,firstdescribed
byMiers(1849),has12knownspeciesandisamong
agroupofapproximately12so-called“physaloid”
genera in the large tribe Solaneae (Averett, 1979;
D’Arcy,1991).Thisgroup,whichnotsurprisinglyin¬
cludesthelargegenusPhysalisL.,isunitedbyhav¬
inglongitudinallydehiscentanthers,ovarialnectaries,
andaccrescentcalycesthatsurroundandeitherclose¬
lyinvestorinflatearoundtheberry.Recentmolecular
systematicworksuggestssubtribalstatusofthisphys¬
aloid clade may Ik* justified (Olmstead & Palmer,
1992;Olmstead&Sweere,1994).
Species of Larnax are single-trunked shrubs 30
cm-2 m tall. Although infrequent in most habitats,
theyeasilyarerecognizedbytheirplagiotropicup¬
per stem and leaf growth and by the axillary fas¬
cicles of from 1 to several flowers per node. Corolla
color ranges from cream to yellow to purple and is
variable within species (Sawyer, unpublished).
Fruitsarefleshy,orangeberriesusuallycontaining
from 60 to over 100 small seeds. Larnax is taxonomicallyassociatedwiththegenusDepreaRafinesque (Barboza & Hunziker, 1994; Hunziker,
1977).HeterantheryinLarnaxspeciesisonechar¬
acterthatdelimitsthisgenusfromDeprea.Inspe¬
cies of Deprea , anthers in the same flower are of
equal size, whereas in species of Larnax, the five
anthersaregroupedinarraysofeithertwoorthree
different size classes. Other characters that sepa¬
ratethesegeneraincludethepresenceofthickened
filamentbasesformingastamenpetalum(Barboza
&Hunziker,1991)in Larnaxspecies(filament bas¬
es are never thickened in Deprea), and the degree
of corolla fusion (in Deprea corollas are infundib¬
ular, the limb shorter than the tube; in Larnax co¬
Novon 8: 72-76. 1998.
rollasarealwaysrotate-campanulate,thelimbal¬
ways longer than the tube) (Barboza & Hunziker,
1994;Sawyer,unpublished).
Species of Larnax are tropical, Andean shrubs
oflimiteddistributionoccurringfromColombiato
northern Peru, with eight species in Ecuador, five
of which are endemic. Recently, a new species was
describedextendingtherangeintoVenezuela(Ben¬
itez de Rojas & Martinez, 1995). Species of Larnax
inhabitwet,premontaneormontaneforestedges
andusuallyarefoundonslopesalongstreams.
In addition to the species described herein, the
followingtenspeciescomprisethegenus:
Larnax harlingiana Barboza & Hunziker. 1995.
Kurtziana24:157—160.Distribution:Ecuador.
Ixirnax haivkesii Hunziker. 1977. Kurtziana 10:
7—50.Distribution:ColombiaandEcuador.
Larnax hunzikeriana Benftez & Martinez. 1995.
Phytologia78:353—356.Distribution:Venezuela.
Larnax lutea Leiva. 1996. Amaldoa 4: 15—22.
Distribution:
Peru.
Larnax peruviana (Zahlbruckner) Hunziker.
1977. Basionym: Athenaea peruviana Zahlbruek.
1892. Ann. K. K. Naturhist. Hofmus 7: 7. Distri¬
bution:EcuadorandPeru.
Larnax purpurea Leiva. 1996. Amaldoa 4: 15—
22.Distribution:Peru.
Larnax sachapapa Hunziker. 1977. Kurtziana
10:7—50.Distribution:ColombiaandEcuador.
Larnax steyermarkii Hunziker. 1977. Kurtziana
10:7—50.Distribution:Ecuador.
Larnax subtriflora (Ruiz & Pav6n) Miers. 1849.
Basionym:PhysalissubtrifloraRuiz&Pav6n.1794.
FI. Per. 2: 42. Distribution: Peru.
Larnax suffruticosa (Dammer) Hunziker. 1977.
Basionym:IochromasuffruticosaDammer.1905.
Bot. Jahrb. 36: 386. Distribution: Ecuador.
Larnax andersonii N. W. Sawyer, sp. nov. TYPE:
Ecuador. Napo: km 25 of Hollfn-Loreto road,
fincaentrancenexttobridgeoveraquebrada
in secondary pluvial forest, 950 m, 00°40'S,
77°40'W, 1 July 1995, Sawyer & Tirado 714
(holotype,MO;isotypes,CONN,US).Figure1.
Interquaternispeciesceterisgenerisfructuinvoluto
laxecalycetrichomatibuslongisetinterdumramosis,ra-
Volume 8, Number 1
1998
Sawyer
New Species of Larnax from Ecuador
73
FigureI.LarnaxandersoniiN.W.Sawyer(Sawyer714).—A.Branchapex.Bar=1cm.—B.Matureflower.—C.
Interiorofmaturecorollaandandroecium.—I).Maturefruitingcalyx.—E.Maturegynoeciumwithannularnectary
atbase.—F.Bifurcateleaftrichome.—G.Matureseed.BarsB—F=1mm.
misseeundariisgenieulatis,foliisgeminatisdisparibus,
floribussolitariisetdistalibusinramis,corollisluteolis
autpurpuratis,antherisapiculatissedrecedensabisdem.
Suffnitescent perennial, 0.5—2 m tall. Stems
branched,secondarybranchesgeniculate.Stems
and leaves villous—sericeous, covered with long,
multicellular, simple or occasionally branched trichomes. Leaves membranous, entire, paired, un¬
equal, the larger (major) elliptic, 4.5—11 cm long,
1.6-3.9 cm wide with 5—8 secondary veins, the
smaller (minor) elliptic-ovate—ovate, 0.3-2.5 cm
long, 0.6—1.9 cm wide. Leaves strigose-sericeous
above, villous—sericeous below especially along
veins,bifurcatehairsoccurringoccasionallyalong
leafmarginandalongveins;apexacuminateinma¬
jorleaves,acute—mucronateinminorleaves,base
oblique—equal,attenuateinmajorleaves,some¬
timesattenuate—truncateinminorleaves.Petiole
74
Novon
sericeous, 3—10 mm long on major leaves, 0.5—3
mm long on minor leaves. Inflorescence distal on
thebranch,axillary,typicallysolitary,occasionally
paired.Flowerspendentonsericeouspedicels3—5
mmlong.Floweringcalyxvillous—sericeous,green,
1.5—2 mm long, 2.5—3 mm wide, margin ranging
from broadly pyramidal at the 5 major veins to 5lobed, lobes acute, <1 mm long. Corolla campanulate, 6—8 mm long, tube 2—3 mm long, lobes of
thelimbmembranous,narrowlytriangular,sparsely
villousexternally,denselytomentosealongmargin
andatapex,bifurcatehairsoccurringrarelyexter¬
nally and along margin, glabrous within. In shade
plants,corollapaleyellow—cream—green;insun
plants,corollavioletwithcreambase.Stamensin¬
cluded. Filaments glabrous, filamentose, the free
portion 1.6-2 mm long, adnate to the basal third of
thecorollaandtherebroadenedandthickenedinto
astamenpetalumwithprojectionsattheshoulders,
separated from each other by the corolla vascula¬
ture, fused portion 0.9—1.8 mm long. Anthers
white-pink,ovatewithlongapiculum,occurringin
two size arrays of three large and two small, 1.3—
1.6 mm long, 0.75 mm wide, slightly dorsifixed,
base sagittate. Ovary glabrous, ovate, 1 mm long,
less than I mm wide, ringed basally by a greenish
yellow nectary. Style glabrous, 2—4 mm long, ex¬
tendingbeyondantherslateinanthesis;stigmaclavate—subbilobate,green—purple.Fruitingpedicels
sericeous, 5 mm long. Fruiting calyx accrescent,
looselyenvelopingthefruit,openattheapex,green
withdarkgreenveins,villous,membranous,pyri¬
form, 1.2-1.3 cm long, 0.8—1 cm wide, with scat¬
tered,two-celledglandularhairswithin;lobesdis¬
tinct, triangular, acute, unequal in length, to 2 mm
long. Fruit an orange, fleshy berry containing 60—
80 reniform, faveolate seeds to 2.8 mm diam.
Solanum sect. Basarthrum, notably his ground¬
breaking work on cryptic dioecy, warrants recog¬
nition.Hisexemplary,broad-scaledinvestigations
ranging in context from experimental research to
applied systematic studies, including studies of
pollenandhairtypes,haveprovidedbothinspira¬
tionandguidanceforhisstudentsandcolleagues.
A suite of characteristics are diagnostic for this
species. They include the geniculate younger
stems, a character present in many species of this
group,andthedistinctlyunequal-geminateleaves
on younger branches; the usually solitary flowers
occurring distally on branches; the apieulate an¬
thers also found in several species; and the long,
sometimesbifurcate,multicellularhairsalsofound
inL.subtrifloraandL.suffruticosa(Sawyer,unpub¬
lished).
Distribution.Inwetlowermontaneforest,usu¬
ally on slopes next to streams. Limited to the east¬
ern Andean slopes of north and central Ecuador,
from 900 to 1500 m.
ThespecificepithethonorsGregor)'J.Anderson,
whosededicatedandexhaustiveresearchintothe
evolutionaryandreproductivebiologyofthegenus
Paratypcs.ECUADOR.Napo:Arehidona,faldasal
surdelVolcanSurnaco,CarreteraHollfn—Loreto,km31.
ComunaClialluaYacu.1200m.(H)°43'S,77°36'W,8—17
Jan.1989,Alvarado222(N\);newroadtoLoreto(56km
SofBaeza),28kmKofjunctionwithBaeza-lenaHoad,
ll(M)m.00°50'S,77°33'W,21Dee.198ft.Hummel&
Wilder17262(NY):LIChacoCanton.ProyectoHidroelectricoCoca,I’untoST3,margenderechadelRfoQuijos,
1500m,00°1I'S.77°39'W,3-5Oct.1990,Palacios5805
(QCNE);LIChacoCanton,ProyectoHidroelectrieoCoca,
PuntoST4,margenderechadelRfoQuijos,ca10kmal
surtieReventador,1450m,00°08'S,77°3()'W,C>—10Oct.
1990,Palacios6050(QCNE).Pastaza:HaciendaSan
AntoniadelBaronvonHumboldt,2kmalNLdeMera,
13(H)m,01°27’S.78°(XVW,27Feb.-19Mar.1985,Baker■
Xeill.PalaciosA’Zaruma5662(MO.NY.QAME);along
roadfromPuyotoMacas,ca.33kmSofPuyo,24.9km
SofVeracruz,10kmSofLseuelaFiscalCotopaxi,dis¬
turbedprimaryforest,'HK>m.01°38'S,77°52'W,3May
1984,Croat58046(NY).
Larnax psilophyta IN. W. Sawyer, sp. nov. TYPE:
Ecuador.Zamora-Chinchipe:NudodeSabanilla, pass on road from Yangana to Valladolid,
2800—2900 m, elfin forest and clearings, 5
Apr. 1985, Hurling & Andersson 23724 (holotype, NY). Figure 2.
Speciesrarissimaalocounicocognitointerquaterni
speciesceterisgenerisfruclibusinvolutisarteealyce.Ah
caulibus,ramis,etfoliisglaberisomnino,foliisinduratis
amboapiceetbaseacutato,floribusparvulisusque0mm
longis,corollaviridia-alba,antherisexsertis.ealycefructiferorotundotantum7mmindiametroaspeciebusdescriptisIructibusinvolutisarteealycenotisbenedistincta.
Suffrutescent perennial, 1-2 m tall. Stems
branched,secondarybranchesgeniculate.Stems
andleavesentirelyglabrous.Leavessomewhatin¬
durate-fleshy, entire, elliptic, 4-5 cm long, 1—2 cm
widewith3^4secondaryveinsprominentabaxially,
apex acute, base cuneate, oblique; minute, stalk¬
less,unicellularredglandsoccurringabaxiallyto¬
ward the leaf base. Petiole glabrous, 5-7 mm long.
Inflorescenceaxillary,1—3flowerspernodeit)fas¬
ciclesfromamuchreducedpeduncle,pendenton
glabrous pedicels 6—7 mm long. Flowering calyx
glabrousexceptattheapexwhereshorthairsmay
bepresent,green,1.5mmlong,2.5mmwide,mar¬
gin broadly pyramidal at the 5 major veins. Corolla
campanulate-rotate,4.6—6.4mmlong,tube1.5—2.8
mm long, lobes of the limb coriaceous, ovate-tri¬
angular, reflexed, margins and apex puberulent.
Volume 8, Number 1
1998
Sawyer
New Species of Larnax from Ecuador
75
Figure2.LarnaxpsilophytaN.W.Sawyer{litirlmp&Andersson23724).—A.Branchapex.Bar=Icm.B.Mature
flower.—C.Interiorofmaturecorollaandandroeeium.—I).Maturefruitingcalyx.—h.Maturegynoeciumwithannular
nectaryatbase.BarsB-E=1mm.
otherwiseglabrouswithout,nonglandular,pluricellular huger hairs present in an annular ring at an¬
therlevelwithin.Corollapaleyellow—eream-green.
Stamensincluded.Filamentsglabrous-puberulent,
hlamentose-ribbon-like,thefreeportion1—1.5mm
long, adnate to the basal third of the corolla and
therebroadenedandthickenedformingastamen
petalumwithextendedshoulders,separatedfrom
each other by the corolla vasculature, the fused
portion0.8-1.4mmlong.Antherswhite,ovate,mi¬
nutely apiculate or, more commonly, without apiculum, occurring in two size arrays of three large
and two small, 1.2-1.6 mm long, 1 mm wide, slight¬
lydorsihxed,basesomewhatsagittate.Ovarygla¬
brous, ovate, 1 mm long, less than 1 mm wide,
ringed basally by a greenish yellow nectary. Style
glabrous, 3^1 mm long, extended beyond anthers
lateinanthesis;stigmaclavate-subbilobate,green.
76
Novon
Fruiting pedicels glabrous, to 10 min long, raising
thematurefruitabovetheleafplane.Fruitingcalyx
accrescent,tightlyenvelopingthefruit,openatthe
apex,green,glabrous,membranous,globose,7mm
diain.withscattered,2-celled,glandularhairswith¬
in; teeth short yet distinct, triangular, acute, <1
mm long. Fruit an orange, fleshy berry containing
about30,reniform,faveolateseedsto2.6mmdiam.
cional
Podocarpus,
Road
Yangana—Valladolid,
km
21,
vi¬
cinityofsampleplot,2560m,4°28'S.79°09'W.31July
1996,Sawyer770(CONN,LOJA):Provinceboundary,
passoverNudodeSabanilla,elfinforest,2740m,4°27’S.
79°10'W.IIMay1985,Stein&I)Alessandro2733(k.
NY).
Diagnostic characteristics for this species in¬
clude the geniculate younger stems as in L. andersonii,theglabrousnatureoftheplant,theindurate
leaveswithacuteapexandbase,andtheverysmall
greenishwhiteflowers.Larnaxpsilophytaisfound
athigherelevationsthanotherknownLarnaxspe¬
cies and appears to be limited in occurrence to the
Parque Nacional Podocarpus and the Nudo de $abanillaareasofsouthernEcuador.Thespecificep¬
ithet invokes the smooth texture of the foliage re¬
sulting from the combined effects of the indurate
and glabrous conditions found in the foliage. This
specieshasbeenconsistentlyconfusedwithDeprea
glabra,anotherglabrousspeciesfoundathighel¬
evations in Ecuador. However, D. glabra has sev¬
eralfeaturesthatallowiteasilytobedistinguished
from L. psilophyta, viz., falcate leaves with attenu¬
ate base, a denser ring of pubescence within the
muchlargercorollatube,anthocyaninsthatareal¬
wayspresentinthecorolla,antherthecaewithbas¬
es that are connate rather than sagittate, and a
northerlyEcuadoriandistribution.
Distribution. Endemic to the elfin cloud forest
of the Parque Nacional Podocarpus and the Nudo
deSabanillapassregionattheborderofZamora—
ChinchipeandLojaprovincesinsouthernEcuador,
from 2500 to 3000 m.
Paralypes.ECUADOR.Loja:ParqueNacionalPodoearpus,newroadLoja—Zamora,EofCerroYanococha,
montaneforestalongformerIndiantrailtoZamora,slight¬
lydisturbed,2550-2650m,3°59'S,79°07'W,26Nov.
1988,Madsen7.55X1(AAU,LOJA,OCA.OCNK).Loja/
Zamora-Chinchipe:
Parque
Nacional
Podocarpus.
Road
Yangana—Valladolid,km21,vicinityofsampleplot,
2700-2800m,4°28'S,79°09'W,24-25Jan.1989,Modsen8566/(AAU,LOJA,QCA,OCNK):RoadYanganaValladolid,atentranceofParqueNacionalPodocarpus,
2500—3000m.4°28'S,79°10'W,10Dec.1989,Madsen
86688(AAU,LOJA.QCA,OCNK);ParqueNacionalPo¬
docarpus,
pass
on
road
Yangana—Valladolid
(Nudo
de
Sa¬
banilla),2750-2900m.4°27'S.79°08'W,28Feb.1985.
0llgaardelal.58374(AAU,LOJA.OCNK):ParqueNa¬
Acknowledgments. This work would not have
beenpossiblewithoutagrantforfieldworkfromthe
Bamford Fund of the Department of Ecology and
Evolutionary Biology, University of Connecticut. I
thank the folks at the New York Botanical Garden
and the Royal Botanical Gardens at Kew for loans
ofherbariumspecimens;DavidNeillforproviding
me with a vehicle and a place to work in Ecuador;
Milton Tirado for his navigational skills and me¬
chanical assistance; Gregory Anderson, Antoni
Damman,WilliamD’Arcy,KentHolsinger,Arman¬
do Hunziker, and Michael Nee for invaluable ad¬
vice; and, finally, Mary Jane Spring for tier assis¬
tanceinthecreationoftheillustrations.
Literature
Cited
Averett,J.E.1979.Riosystematicsofthephysaloidgen¬
eraoftheSolaucaeinNorthAmerica.Pp.493—503in
J.G.Ilawkes,R.N.LesterA'A.I).Skelding(editors),
he
I BiologyandTaxonomyoftheSolanaeeae.Academ¬
ic
Press,
London.
Barboza,C.F,.AA.T.Hunziker.1991.Flstudiossobre
SolanaeeaeXXXI.Peculiaridadesdelandroceodeinteres
taxonomico
en
Solanum.
Kurtziana
21:
185—194.
- & -. 1994. Estudios sobre Solanaeeae
XXXVII.SinopsistaxonomicadeDeprea.Kurtziana23:
101-124.
Benitez de Rojas. (.. L. & M. Martinez. 1995. Ixirnax
hunzikeriana
(Solanaeeae:
Solanoideae).
Una
nueva
especieylaprimeramenciondelgeneroparaVenezuela.
Phytologia
78:
353—356.
D'Arcy.Vi.C.1991.TheSolanaeeaesince1976,witha
reviewofitsbiogeography.Pp.75-137inJ.C.Hawkes,
R.N.Lester,M.NeeAN.Estrada(editors),Solanaeeae
III:Taxonomy,Chemistry,Evolution.RoyalBotanic
Gardens,Kew,andLinnaeanSocietyofLondon.
Hunziker,A.T.1977.EstudiossobreSolanaeeaeVIII.
Novedadesvariassobretribus,generos,seceionesyespeeiesdeSudAmerica.Kurtziana10:7—50.
Miers,J.1849.ContributionstothebotanyofSouthAmer¬
ica.Ann.Mag.Nat.Hist.Ser.2,4:37-39.
Olmstead,R.G.&J.I).Palmer.1992.Achloroplastl)NA
phylogeny
of
the
Solanaeeae:
Subfamilial
relationships
andcharacterevolution.Ann.MissouriRot.Card.79:
346-360.
-& .1. A. Sweere. 1994. Combining data in phylo¬
genetic
systematies:
An
empirical
approach
using
three
moleculardatasetsIntheSolanaeeae.Syst.Biol.43:
467-481.
Nomenclatural
Changes
in
Leptochloa
(Poaceae,
P.
Beauvois
Sensu
Lato
Chloridoideae)
NeilSnow
Washington University, Department of Biology, P.0. Box 1137,
St. Louis, Missouri 63130, U.S.A. and
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Current Address: Queensland Herbarium, Meiers Road, Indooroopilly, QLD 4068, Australia
Email: [email protected] ,ld .gov.au
Abstract. The following new names are pro¬
posed for Leptochloa P. Beauvois s.l. (including
Diplachne P. Beauvois), based on recent revision¬
ary and cladistic studies: Leptochloa caudata, L.
decipiens subsp. asthenes, L. decipiens subsp.
peacockii, L. eleusine, L. fusca subsp. muelleri, L.
fusca subsp . fascicularis, L. fusca subsp. uninervia, L. gigantea, and L. panicea subsp. brachiata.
Recent revisionary and cladistic studies in
Leptochloa P. Beauvois (Snow, 1997a), along with
preparation of the grass treatment for Flora Zambesiaca (Cope, in press), support nomenclatural
changes for this nearly worldwide grass genus.
The changes are necessary given the consistent
lack of cladistic support for Diplachne P. Beau¬
vois as a taxon distinct from Leptochloa (Snow,
1997a) and the need to reduce in rank several
species (Snow, 1997a, 1997b).
The synonymy of Leptochloa is extensive and
only recently has been assessed from a global
perspective (Snow, 1997a). However, since the
primary purpose of this article is to make these
new names available, complete synonymy will be
withheld for a later publication (or see Snow,
1997a). In addition to these changes, descrip¬
tions of two new species are forthcoming (Snow,
1998; Snow & Simon, 1997).
Leptochloa caudata (K. Schumann) N. Snow,
comb. nov. Basionym: Diplachne caudata K.
Schumann,inEngler,Pflanzenw.Ost.-Afrikas
C: 113. 1895. TYPE: Tanzania. Ukera, Fischer
674(holotype,B).
Leptochloa decipiens (R. Brown) Stapf ex Maid¬
en subsp. asthenes (Roemer & Schultes) IN.
Snow, comb, et stat. nov. Basionym: Foa as¬
thenes Roemer & Schultes, Syst. Veg. 2: 574.
1817. Poa imbecilla R. Brown, Prodr. 181.
1810, nom. horn, illeg., non P. imbecilla Solander ex Sprengel, PI. Nov. Herb. Spreng., 9
no. 14. 1807. Leptochloa asthenes (Roemer &
Schultes) C. E. Hubbard, Bull. Misc. Inform.
Kew: 26. 1941. TYPE: Australia. Queensland,
Upper Head [=Chadron Point], Broad Sound,
R. Broivn 6270 (lectotype, here designated,
BM; isolectotype, K). Blake (1972: 6) correctly
cited the type collection but did not designate
a particular duplicate as the type specimen.
EragrostisciliolataJedwabnick,Bot.Arch.5(3-4):192.
1924.Leptochloaciliolata(Jedwabnick)S.T.Blake,
Contr.QueenslandHerb.14:6.1092.TYPE:Aus¬
tralia.NewSouthWales,Narrabri,Maidens.n.(lec¬
totype(asholotypebyLazarides,1980:262),B;iso¬
lectotype.
Bill).
Mydissertationindicatedthat“Eragrostisimbe¬
cilla Benth. FI. Austral. 7: 643. 1878, non E. im¬
becilla (R. Brown) R. Brown ex Steudel, Syn. PI.
Glumac. 1: 279. 1854” was a taxonomic synonym
of Leptochloa decipiens subsp. asthenes (Snow,
1997a:166).Afewadditionalcommentsareuseful
at this time. Given the format used by Bentham
(1878) in volume 7 of Flora Australiensis, Eragros¬
tis imbecilla appears to be merely a new combina¬
tion, not a new taxon (e.g., see Article 58.3, Greuter
et ah, 1994). This is evident from his citation of
Poa imbecilla and explicit reference to the type
specimencollectedbyForsterinNewZealand(ho¬
lotype, B, Willdenow Herbarium Cat. No. 01896,
microfiche). The correct citations for these names
appear to be Eragrostis imbecilla (Solander ex
Novon 8: 77-80. 1998.
78
Novon
Sprengel) Bentham, which was based on Poa imbecilla Solander ex Sprengel (Veldkamp, pers.
comm.; see also Hiepko, 1969; Garnock-Jones,
1986). The paniculate inflorescence of the type of
P. imbecilla Solander ex Sprengel precludes its in¬
clusion in Leptochloa, as does its origin from New
Zealand,whichliesbeyondthenormalrangeofthe
genus (Snow, 1997a). Although the clarity of the
microfiche was inadequate for me to suggest the
propergenericplacementoftheForstercollection,
it clearly is not Leptochloa, and Blake (1972: 6) has
ascribed the specimen back to Poa. Most impor¬
tantly, names based on Poa imbecilla Solander ex
Sprengel,includingEragrostisimbecillaBentham,
Leptochloa debilis Stapf ex C. E. Hubbard (Hub¬
bard, 1941: 26), and Poa sprengleii Kunth (Kunth,
1833: 363) are to be excluded from Leptochloa
(contraSnow,1997a;Lazarides,1997).Itshouldbe
noted, however, that Hubbard (1941) considered
Eragrostis imbecilla Bentham as a new taxon, not
merelyanewcombination.
Leptochloadecipienssubsp.astheneswasformer¬
ly recognized at the species level as Leptochloa ciliolata (Lazarides, 1980; Staidey & Ross, 1989;
Simon, 1993). Although it can be locally distinct
inthefield,allcharactersintergradetosomeextent
with L. decipiens subsp. decipiens (Snow, 1997a,
1997b).
Leptochloa eleusine (Nees) T. A. Cope & N.
Snow,comb.nov.Basionym:Diplachneeleu¬
sine Nees, FI. Afr. Austr. 255. 1841. Triodia
eleusine (Nees) T. Durrand & Schinz, Consp.
FI. Afr. 5: 877. 1894. Uralepis eleusine (Nees)
Steudel, Syn. PL Glumac. 1: 248. 1854.
TYPE: South Africa. Katrivierspoort, Drege
3906(lectotype,heredesignated,B;isolectotype,P).
Leptochloa decipiens subsp. peaeoekii (Maiden
& Betche) N. Snow, comb, et stat. nov. Basionym:DiplachnepeaeoekiiMaiden&Betche,
Agrie. Gaz. New South Wales 15: 925. 1904.
Leptochloapeaeoekii(Maiden&Betche)Domin, Biblioth. Bot. 85: 379. 1915. TYPE: Aus¬
tralia. New South Wales: Coolabah, 4 Dec.
1904, Maiden & Boorman s.n. (lectotype |as
holotype by Lazarides], 1980: 263], NSW; isolectotypes, BM, BRI, K, W). As correctly not¬
ed by Blake (1972: 9), plate 2 in tlx* original
protologueisanerroneouselement(Maiden&
Betche, 1904), and represents the American
speciesLeptochloadubiaKunth.
This taxon has been recognized as L. peaeoekii
(Stanley & Ross, 1989; Simon, 1993) or synonymizedunderL.decipienssubsp.decipiens(Lazari¬
des, 1980). Like the previous taxon, some popula¬
tionsaredistinctfromL.decipienssubsp.decipiens,
butinothersmorphologicalintergradationiscon¬
tinuous, such that no character or combination
thereof can consistently diagnose it as a distinct
species.
Leptochloa fuse a (L.) Kunth subsp. muelleri
(Bentham) N. Snow, comb, et stat. nov. Bas¬
ionym:DiplachnemuelleriBentham,FI.Aus¬
tral. 7: 619. 1878. Leptochloa muelleri (Ben¬
tham) Stace, Watsonia 18: 413. 1991. TYPE:
Australia. Charlotte waters, Giles s.n., Herb.
Munro(lectotype,heredesignated,K;isolectotype,
K).
DespitepriorityoftheepithetLeptochloamalabarica (L.) Veldkamp over fusca (L.) Kunth (Veld¬
kamp, 1971), Snow and Davidse (1998) have pro¬
posed rejection of Poa malabarica in the spirit of
theTokyoCode,whichencouragesmaintenanceof
namesincurrentuse.Thisseemsappropriate,giv¬
en the nearly global geographic range of the spe¬
cies, the nearly universal historical usage of the
epithet fusca, the restricted usage of the epithet
malabarica, and the considerable confusion that
hassurroundedtheapplicationoftheepithetmal¬
abarica.
The reduction in rank of this taxon and the two
thatfollowisbasedonexaminationofseveralthou¬
sandherbariumspecimens(representingover50
herbaria)ofthisspeciescomplexfromthroughout
itsrange,coupledwithfieldworkinNorthAmerica,
southern Africa, and Australia, as well as multi¬
variate statistical analyses of eleven population
samples(Snow,inprep.).
Leplochloa fusca (L.) Kunth subsp. faseieularis
(Lamarck)N.Snow,comb,etstat.nov.Basio¬
nym: Festuca faseieularis Lamarck, Tabl. Encyd. 1: 189. 1791. Diplachne faseieularis (La¬
marck) P. Beauvois, Ess. Agrostogr. 81, 160,
pi. 16, f. 9. 1812. Cynodon faseieularis (La¬
marck) Raspail, Ann. Sci. Nat., Bot. 5: 303.
1825. Festuca aquatica Bose ex Roemer &
Schultes, Syst. Veg. 2: 615. 1817, nom. inval.,
as syn. of Diplachne faseieularis P. Beauvois.
DiplachneaquaticaBoseexRoemer&Schul¬
tes, Syst. Veg. 2: 615. 1817. TYPE: South
America. D. Richard s.n. (holotype, P).
79
Volume 8, Number 1
1998
Snow
Nomenclatural Changes in Leptochloa
Leptochloa fusca (I..) Kunth subsp. uninervia (J.
Presl) N. Snow, comb, et stat. nov. Basionym:
Megastachya uninervia J. Presl, Reliq. Haenk.
1: 283. 1830. Poa uninervia (J. Presl) Kunth,
Enum. PI. 1: 344. 1833. Eragrostis uninervia
(J. Presl) Steudel, Syn. PI. Glumac. 1: 278.
1854. Brizopyrum uninervium (J. Presl) E.
Fournier, Mex. PI. 2: 121. 1886. Leptochloa
uninervia (J. Presl) Hitchcock & Chase, Contr.
U.S. Natl. Herb. 18(7): 383. 1917. Diplachne
uninervia (J. Presl) Parodi, Revista Centro Estud. Agron. 18: 147. 1925. TYPE: Mexico.
Haenke 101 (lectotype, here designated, PR
not seen; isolectotypes, W, EE not seen).
and lower portions of the Mississippi River drain¬
age and portions of the Ohio River drainage to the
east. The application herein of /,. panicea subsp.
mucronata (and Snow, 1997a) is in a narrower
sense than that of Nowaek (1994, 1995), who in¬
cluded all New World specimens under this name.
Leptochloa gigantea (Launert) T. A. Cope & N.
Snowr,comb.nov.Basionym:Diplachnegigantea Launert, Bol. Soc. Broteriana ser. 2a, 47:
349. 1974. TYPE: Zambia: Mbala (Abercorn),
Vesey-Fitzgerald 1551 (holotype, K; isotypes
BM, SRGH not seen).
Leptochloa panicea (Retzius) Ohwi subsp. hrachiata (Steudel) N. Snow, comb, et stat. nov.
Basionym:LeptochloabrachiataSteudel,Syn.
PI. Glumac., 209. 1854. TYPE: Guadaloupe.
Duchassaings.n.(holotype,P,fragmentUS).
AsrecognizedbySnow(1997a),Leptochloapan¬
iceasensulatoisapolymorphicspeciesthatranges
throughoutmuchofthewarmtemperateandtrop¬
ical regions of the world. It is comprised of three
subspecies.
Leptochloapaniceasubsp.panicea,asrecognized
by Snow (1997a), corresponds closely to the taxon
ofthesamenameasrecognizedbyNowaek(1994).
It is an Old World taxon occurring mostly in Africa
and southern Asia, but which has been verified re¬
cently from several duplicates collected at a site in
northwesternQueensland,Australia(Snow,1997a;
Snow & Simon, in press).
Leptochloapaniceasubsp.brachiata,asrecog¬
nized by Snow (1997a), is by far the more common
of the two New World subspecies in this complex
andhasaconsiderablylargerrange.Itrecentlyhas
been known as L. filiformis (Lamarck) P. Beauvois,
L. mucronata (Snow & Davidse, 1993), and L. pan¬
iceasubsp.mucronata(Miehaux)Nowaek(propar¬
te) (Snow & Davidse, 1993; Nowaek 1994, 1995).
Leptochloa panicea subsp. mucronata , sensu
Snow (1997a), is the correct name for the taxon
formerly known as Leptochloa attenuata (Nuttalll
Steudel (Allen, 1980) or L. filiformis var. attenuata
(Nuttall)Steyermark&Kucera(Steyermark,1963).
This taxon is restricted to the U.S.A. in the central
Acknowledgments. My sincere thanks to Gerrit
Davidse of the Missouri Botanical Garden for as¬
sistance with the numerous and challenging no¬
menclatural puzzles in Leptochloa ; however, any
misinterpretations or oversights are mine. The
prompt and thorough review by J. Veldkamp was
muchappreciated.Thanksalsotothenumerouscu¬
ratorswholoanedherbariumspecimens.Research
was supported in part by the Missouri Botanical
Garden (Mellon Foundation), Evolutionary and
PopulationBiologyProgramatWashingtonUniver¬
sity,andtheNationalGeographicSociety.
I.iterature
Cited
Allen,C.M.1980.GrassesofLouisiana.Univ.South¬
western
Louisiana,
Lafayette.
Bentham,G.1878.FloraAustraliensis:Adescriptionof
thePlantsoftheAustralianTerritory.Vol.7.RoxburghiaoeaetoKilices.L.Reeve,London.
Blake,S.T.1972.PlinthanthesisandDanthoniaanda
review'oftheAustralianspeciesofLeptochloa(Gramineae).Contr.QueenslandHerb.14:1-19.
Cope.T.A.Inpress.InFloraZambesiaca:Floraterraruni
Zarnbesiiaquiseonjunctarum.Vol.10,E.Launert&G.
V.
Pope
(editors).
Garnoek-Jones,PJ.1986.SouthPacificplantsnamedby
k.I*.J.Sprengelin1807.Taxon35:123—171.
Greuter,W.F.R.Barrie,If.M.Burdet,W.G.Chaloner,
V.Demoulin,I).L.Hawksworth,P.M.J0rgensen,I).H.
Nicolson,P.C.Silva,P.Trehane&J.McNeill,Editors.
1994.InternationalCodeofBotanicalNomenclature
(TheTokyoCode),AdoptedbytheXVtliInternational
Botanical
Congress,
Yokohama,
August-September,
1993.RegnumVeg.131.
Hiepko,P.1969.VonJ.R.undG.Forstergesammelte
PflanzeninHerbarWilldenowinBerlin.Willdenowia
5:
279-294.
Hubbard.C.E.1941.GrarnineaeAustralienses:III.Bull.
Misc.
Inform,
(no
volume
number),
25—31.
kunth,C.S.1833.EnumeratioPlantarum.TomusPrimus.
StutgardiaeetTubingae.SumitbusJ.G.Cottae.
Lazarides,M.1980.ThegenusleptochloaBeauv.(Poaeeae.Eragrostideae)inAustraliaandPapuaNew
Guinea.
Brunonia
3:
247—269.
-. 1997. A revision of Eragrostis (Eragrostideae,
Eleusininae,Poaeeae)inAustralia.Austral.Syst.Bot.
10:
77-187.
Maiden,.1.H.&E.Betehe.1904.AnewAustraliangrass
(DiplachnePeacockii,MaidenandBetehe).Agrie.Gaz.
NewSouthWales15:925—926,plates1,2.
Nowaek.R.1994.RevisionofleptochloaBeauv.(inch
DiplachneBeauv.)inMalesia.Rheedea4(2):79—92.
-.1995.AnewcombinationinMalesianLeptochloa
Beauv.Rheedea5(1):93.
Simon,B.K.1993.AkeytotheAustralianGrasses.2nd
80
Novon
ed.
Queensland
Department
of
Primary
Industries,
Bris¬
bane.
Snow,N.1997a.PhylogenyandSystematic^ofleptochloa
P.Beauvoissensulato(Poaceae,Chloridoideae).Ph.I).
Dissertation.
Washington
University,
St.
Louis,
Missou¬
ri.
-. 1997b. Application of the phylogenetic species
concept:
botanical
A
monographic
perspective.
Austrobaileya5:1-8.
-. 1998. A new species of Leptochloa from Sri Lan¬
ka(Poaceae,Chloridoideae).Novon8.inpress.
-& G. Davidse. 1993. Leptochloa mucronata is the
correct
name
for
Leptochloa
filiformis
(Poaceae).
Taxon
12:
413-117.
- & -. 1998. Proposal to reject the name Poa
malaharica(Graniineae).Taxon47,inpress.
-& B. K. Simon. 1997. leptochloa southwoodii (Po¬
aceae:Chloridoideae),anewspeciesfromsouth-east
Queensland.
Austrobaileya
5:
137-143.
- & -. In press. Taxonomic status and Aus¬
traliandistributionoltheweedyneotropicalgrasslep¬
tochloafuscasubsp.uninervia,withanupdatedkeyto
Australian
Leptochloa
(Poaceae,
Chloridoideae).
(Aus¬
trobaileya)
Stanley,T.I).&E.M.Ross.1989.Eloraofsouth-eastern
Queensland.VolumeIII.QueenslandDepartmentof
Primary
Industries,
Brisbane.
Steyermark,,1.A.1963.FloraofMissouri.IowaState
Univ.
Press,
Ames.
Veldkamp,J.F.1971.NotesonMalesiangrassesV.New
species
and
combinations
in
Pheidochloa,
Hyparrhenia
,
andleptochloa.Blumea19:61-64.
Croton
martinianus
(Euphorbiaceae),
a
New
Species
from
Mexico
Victor W. Steinmann
Rancho Santa Ana Botanic Garden, 1500 N. College Avenue,
Claremont, California 91711, U.S.A.
ABSTRACT. A new species of Croton (sect. Geiseleria ) from western Mexico is described and illus¬
trated. The relationship between this species and
C. cupulifer McVaugh is discussed, and a key dis¬
tinguishing these two species and C. glandulosus
L.isprovided.
Thetropicaldeciduousforestfloraofsoutheast¬
ern Sonora, Mexico, is diverse and still relatively
little known. This area was first floristically treated
by the late Howard Scott Gentry in his 1942 pub¬
lication Rio Mayo Plants. During the past few
years, the University of Arizona Herbarium has
conductedaprojecttoreviseandupdatethiswork.
Tripstoinventorythearea’splantshaveresultedin
a number of collections of an interesting herba¬
ceousCroton;onespecimenfromnorthernSinaloa
and two specimens from Jalisco have also been lo¬
cated. These collections cannot be referred to any
known species and are here described as new.
Croton martinianus V. W. Steinmann, sp. nov.
TYPE: Mexico. Sonora: Mpio. Alamos, Sierra
de Alamos, N side of the range ea. 2 km SW
of Alamos along the trail from El Chalaton to
Ea Huerta, 27°00'N, 108°58'W, ca. 750 m, 31
Aug. 1996, Steinmann 952 (holotype, AR1Z;
isotypes, BM, DAV, IBUG, MEXU, MICH, MO,
NY,’ RSA, USON). Figure 1.
Herbaperennisvelfortasseannuausquead30cmalta,
erecta
vel
ascendens,
stellato-pubescens;
stipulae
attenuato-subulatae;
foliaprincipaliaalterna,longipetiolata,
laminaovata,dentato-serrata;foliasuperioraminoraet
saepe
angustiora
et
lanceolata,
opposita;
racemus
terminalis,floribuspistillatis(1—)4—8.staminatisusquead40;
bracteae
attenuato-subulatae;
florum
staminatorum
petala
alba,stamina10—11;(lorespistillatiapetali,sepalisviridibus,
inaequalibus,
oblanceolatis,
obovatis,
vel
spathulatis,ovariasubglobosa,stellato-villosa,styli3,bipartiti;
eapsulae
subglobosae;
semina
oblonga,
nitida,
carunculata.
Crotoneeupuliferoaffinis,aquodiffertbracteislongis
ovario
et stellato-villoso.
SpeciesinhonorumPaulS.Martin(1928—).
Monoecious perennial herbs (but flowering in
the first year) to 30 cm high, erect to ascending,
stellate-pubescentthroughout;fromanarrowtap¬
root reaching 6 mm diam.; stems little branched
below, often diffusely branched above; lower in¬
ternodes to 6.5 cm long, decreasing in length
above; stipules linear-subulate to 5.3 mm long,
usually terminated by stellate trichomes; lower
leaves alternate, on petioles (0.6—)1.1-3.8 cm
long, with a pair of slender stipitate cupuliform
petiolar glands 0.1—0.3 mm diam. on the sides of
the petioles just below the point of attachment
with the lamina, the lamina ovate, palminerved
with 3—5 prominent veins, rounded at the base,
serrate-dentate,generallyacuteattheapex,1.6—
4.7 cm long, 1.4—3.5 cm wide, densely hoary eanescent to green and sparsely pubescent, the tri¬
chomes stellate with rays to 0.6 mm long; upper
leaves nearly sessile or on short petioles, serratedentate,(1.0—)2.0—3.4mmlong,mostlyopposite
or below the flowering branches appearing
whorled;inflorescencesexceedingthesubtending
leaves, terminal, racemose, 2.5—7.0 cm long, with
(1— )4—8 pistillate flowers and up to ca. 40 staminate flowers; bracts linear-subulate, 1.1—2.5 mm
long, subtended at the base by clusters of orangebrown glands ca. 0.1 mm long, tips of the bracts
with1—5straightbristle-likehairs;staminateflow¬
ers on pedicels 1.1—2.0 mm long, calyx 1.7—2.2
mm long, the sepals united toward the base, their
free lobes deltoid, 1.2—1.7 mm long, petals white,
narrowly obovate-elliptic, 1.7-2.2 mm long, 0.9—
1.0 mm wide, villous especially along the margins
and toward the base; small ovate glands ca. 0.2
mm long opposite the sepals and alternating with
the petals; receptacle densely villous within; sta¬
mens10—11,filamentsglabrous,1.0—1.4mmlong,
anthers ca. 0.5 mm long, ca. 0.3 mm wide, elliptic,
basifixed and inflexed in the bud; pistillate flowers
apetalous, on pedicels 0.6—2.1 mm long, the se¬
pals 3.2—4.1 mm long, 1.5—2.9 mm wide, oblanceolate to spatulate, green and somewhat foliaceous, unequal, the proximal generally smaller
and narrower, occasionally small linear-filiform
appendages to 0.8 mm long present between the
sepals; ovary nearly globose, stellate-pubescent,
appearing villous, the rays of the trichomes to 1.9
mm long; styles 3, 1.7—2.0 mm long, biparted, paNovon 8: 81-83. 1998.
82
Novon
1cm
Figure1.CrotonmartinianusV.W.Steinmann.—A.Habit.—B.Pistillateflower.—C.Seed.AllfromSteinmann
93-309.
83
Volume 8, Number 1
1998
Steinmann
CrotonmartinianusfromMexico
pillate and often beset with stellate trichomes;
capsule nearly globose 3.6—3.9 mm diam.; colu¬
mella2.6—3.0mmlong;seedsoblong,3.0—3.3mm
long, 2.0-2.2 mm wide, dorsoventrally com¬
pressed. shallowly foveolate, blackish brown to
mottled black-gray, shiny, carunculate; caruncle
narrowlyandtransverselyelliptic.
la.Pistillateflowersinatightclusterandappearing
fasciculate;inflorescencegenerallylessthan2
cmlongandnotexceedingthesubtendingleaves
.
C.
glandulosus
lb.Pistillateflowersmoderatelyspacedandar¬
rangedinaraceme;inflorescencegenerally3—
10cmlongandexceedingthesubtendingleaves.
2a.
Plants
perennial
herbs,
sometimes
appearing
annual;staminatebracts1.1—2.5mmlong;
pubescenceoftheovaryappearingvillous,
atleastsomeoftheraysofthestellatetri¬
chomes greater than 1 mm long.
.
C.
martinianus
2b.Plantsshrubsorlessfrequentlyperennial
herbs;staminatebracts0.6mmorless;pu¬
bescence
of
the
ovary
appearing
puberulent,
theraysofthestellatetrichomeslessthan
0.3
mm
long
.
C.
cupulifer
Phenology. Flowering and fruiting overlap
broadly. Reproductive plants have been collected
in March and June to September.
Distributionandhabitat.SoutheasternSonora,
northernSinaloa,andnorthernJalisco,150—1200
m. On the north side of the Sierra de Alamos, this
species is frequent along the trail from El Chalaton
to La Huerta, where the plants mostly occur in
rocky, red-orange soil on relatively dry hillsides in
tropicaldeciduousforestandloweroakwoodland.
Etymology. The specific epithet honors the
well-known paleontologist Paul S. Martin, an avid
student of the flora of Sonora and Chihuahua and
atreasuredfriend.
Following the sectional delimitations of Croton
as circumscribed by Webster (1993), C. martinianus belongs to section Geiseleria (Klotzsch) BailIon, where it is most similar to C. cupulifer McVaugh,aspeciesofwesternMexicoknownfromthe
statesofColimaandJalisco.Bothofthesetaxapos¬
sess coarsely toothed leaves that are quite diverse
in form; the lower ones are usually alternate, longpetiolate, and relatively large, while those associ¬
atedwiththeinflorescenceareoppositetowhorled,
subsessile,smaller,andnarrower.Thetwospecies
alsohaveattenuate-subulatestipules,nearlyiden¬
tical cupuliform petiolar glands, and unequal pis¬
tillate sepals that are somewhat foliaceous. In the
protologue of C. cupulifer , McVaugh (1961: 163)
mentioned that 1 or more extra calyx lobes often
develop in the pistillate flowers. Although this un¬
usual characteristic has not been observed in C.
martinianus,linear-filiformappendagesarerarely
presentbetweenthesepals,andthesemayrepre¬
senthighlyreducedsepals.Thesalientdistinguish¬
ing features of these two species are given in the
key.
The leaves of Croton martinianus vary from
denselyhoary-canescenttogreenandsparselypu¬
bescent. This character appears environmentally
influenced,withthehoary-canescentleavesoccur¬
ring in drier periods of the year.
Crotonmartinianusalsobearsaresemblanceto
C. glandulosus I,., to which the first collections
were initially referred. The following key will sep¬
arate C. martinianus from C. glandulosus and C.
cupulifer:
Paratypes. MEXICO. Jalisco: 1 km a NE de San
CristobaldelaBarranca,900m,13Eeb.1993,LomelCn
2010(DAV);SanCristobaldelaBarranca,12kmalSde
lapoblacion,1200m,26Aug.1987,OrnelasACervantes
1199(IIIMO).Sinaloa:Mpio.SalvadorAlvarado,eerros
al N de Terrero, 150 m, 12 Aug. 1988, Bojdrquez 68ft
(MEXU).Sonora:1.25mi.NWofAlamosinElRincon
area. 1350 (t., 3 Sep. 1973, Fish 82 (EC); ca. 2 mi. NE
ofAlamosalongroadtoCuehuhuari,27°OI'N,108°54'\V.
410m.19Aug.1994,Fishbein1892(AKIZ);Sierrade
Alamos, 22 July 1989, Martin s.n. (ARIZ); 8 mi. W of
Alamos,roadfromMinasNuevastoAduana,24June
1984.Starr779(ARIZ.DAV);samelocalityasthetype.
27Aug.1993,Steinmann93-309(ARIZ,DAV,MEXU);
ArroyoGochico,ca.10km(byair)EofSanBernardo,Sfacingslopeabovearroyo,alongthetrailfromSanBer¬
nardo to Gochico Nuevo, in the vicinity of 27°24'N,
108°44'30"W, 750 m, 15 Mar. 1995, Steinmann 606
(ARIZ, DAV. E, HIIMO, MEXU, MO. NY, RSA, UCR);ca.
5km(byroad)NofGiiirocobaalongtheroadtoChoquincahui,inthevicinityof26°57'N.108°41'30"W,ca.
500m,13Mar.1995.Steinmann613(ARIZ);Sierrade
Alamos.ElRinconViejo,ArroyoElAguaje,ca.3.4km
(byair)NofAlamos.27°03'55"N,108°56’W,480-520m.
23Sep.1993,VanDevender93-1066(ARIZ).
Acknowledgments. I thank Paul S. Martin,
Thomas R. Van Devender, and Rebecca K. Wilson
forencouragementandsupport.RupertC.Barneby
kindly assisted with the Latin diagnosis and de¬
scription. Marshall R. Crosby, Mark Fishbein, Gor¬
don McPherson, and Grady L. Webster provided
manyusefulcommentsaboutthemanuscript.Anne
Gondorrenderedtheillustration.Fortheprivilege
of access to their collections, I am indebted to the
curators and staffs at ARIZ, DAV, HUMO, MEXU,
andUC.
literature
Cited
Gentry,H.S.1942.RioMayoPlants.CarnegieInstitution
of
Washington
Publication
527.
Washington,
D.C.
McVaugh,R.1961.EuphorbiaceaenovaeNovo-Galicianae.
Brittonia
13:
145—205.
Webster,G.U.1993.Aprovisionalsynopsisofthesec¬
tionsofthegenusCroton(Euphorbiaceae).Taxon42:
793-823.
Acanthosyris
annonagustata
(Santalaceae),
Eastern
a
New
Species
from
Ecuador
Carmen Ulloa Ulloa and Peter Miller j0rgensen
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. A new species, Acanthosyris annona¬
gustata C. Ulloa & P. j0rgensen, from Napo, Ec¬
uador, is described and illustrated. This species is
characterizedbyitsgoldenbrownpuberulentinflo¬
rescence and fruit. The plant is used in local med¬
icine,andthefruitstastelikeAnnonaandareeaten
by monkeys. In Ecuador, Acanthosyris is also rep¬
resented by A. glabrata (Stapf) Stauffer from the
drywesternforests,atreethatwearealsoreporting
forthefirsttimefromPeru.
RESUMEIM. Se describe e ilustra la nueva especie
Acanthosyris annonagustata C. Ulloa & P.
j0rgensen (Santalaceae) de la provincia de Napo,
Ecuador, que se caracteriza por tener las inflorescencias y frutos pardo-dorado puberulentos. La
planta es medicinal, los frutos tienen sabor a An¬
nona y son comidos por monos. En el Ecuador
Acanthosyristambienest<irepresentadoporA.gla¬
brata(Stapf)Stauffer,unarboldelosbosquessecos
occidentales que ademas se cita por primera vez
para
Peru.
The genus Acanthosyris (Martius & Eichler)
Grisebach in Ecuador was known only from the
type collection of Acanthosyris glabrata (Stapf)
Stauffer made by Baron von Eggers at “Hacienda
El Recreo,” province of Manabf in 1893. The “Ha¬
cienda Ed Recreo” is the type locality of several
species, located north of the town of Bahfa de Cardquez, between San Vicente anti Canoas, and in
1977 only an abandoned house was left by the
beach (H. H. litis, pers. comm.). A century after
the publication of this species in the genus Cervantesia Ruiz & Pavon, we saw the isotype at the
U.S.NationalHerbariumandcouldthereforeiden¬
tify several recent collections from Ecuador and
northernPeru,mostofthismaterialbeingmisidentifiedasXimenia(Olacaceae).Acanthosyrisglabra¬
taisalsoknownfromtwoPeruviancollections,Lao
5153 (MO) and Vargas 9 (F, MO) from the depart¬
mentofTumbes.DuringpreparationoftheSanta¬
laceae for the Flora of Ecuador ; we have studied
specimens from Guayas, Manabf, anti Loja prov¬
inces.Acanthosyrisglabrataisalsoknownfromthe
departmentofAtldnticoinnorthernColombia.This
Novon B: 84-86. 1998.
species grows in dry forests from near sea level to
500 m elevation. Nee (1996) published a key of the
woody Santalaceae for South America and de¬
scribed A. asipapote M. Nee from Santa Cruz, Bo¬
livia. Recent collections of an edible tree from the
Amazonian province of Napo, Ecuador, have re¬
vealed a new species of Acanthosyris, making six
thetotalnumberofspeciespresentlyknowninthe
genus. Acanthosyris is a South American genus of
more or less spiny shrubs and trees, with edible,
drupaceous fruits, distributed from Colombia to
northernArgentina,Uruguay,Paraguay,andBrazil.
In Ecuador it is represented by two species, which
canbeseparatedbythefollowingkey:
la.fruitea.2cmlong,glabrous,smooth;flowers
whitishgrayvelutinous,rachiswhitishgrayvil¬
lous;leavesovate;youngleavessparselyvillose
onthemidribandpetiolebelow;oldtwigswith
exfoliatingbark;Pacificdryforest....A.glabrata
lb.Fruit2.9-4cmlong,denselypuberulent.rugulose;flowersandrachisgoldenbrownpuberu¬
lent;leaveselliptic;youngleavesglabrousor
rarelywithafewminutehairsonthemidriband
petiolebelow;oldtwigswithstriatebark;Ama¬
zonian rainforest. .4. annonagustata
Acanthosyris annonagustata C. Ulloa & P.
j0rgensen, sp. nov. TYPE: Ecuador. Napo:
ParqueNacionalYasunf,carreterayoleoducto
de Maxus en construct ion, km 20, 250 m, 28BO July 1993 (fr), M. Aulestia & G. Grefa 232
(holotype, QCNE; isotypes, AAU, GB, MO,
US).Figure1.
Arbores30maltae,spinosae.Foliaelliptica,5—15cm
longa,2.1-6.8cmlata,glabra,petioiis4—9mmlongis.
Inflorescentiaspicata2.1-6.8cmlonga.Floresextustrichomatibusfulvisdensepuberuli.tepalis5(6).2.0-2.5
mmlongis,staminibus5(6),nectario5(6)-lobalo,lobis
1.5—1.8mmlongis.Fructusdrupaceus,2.9—4cmlongus,
trichomatibus
fulvis
dense
puberulus.
Tree to 30 m tall, 30-45 cm diam.; old branches
withlightbrownorwhitishgray,thinlystriatebark,
spiny; twigs drying dark brown, finely striate, and
angulate, spiny; spines one or two per leafy twig,
4-10.1 mm long, axillary to a leaf and subtending
an axillary bud. Leaf blades elliptic, 5-15 X 2.16.8 cm, acute to acuminate at apex, acute to atten-
Volume 8, Number 1
1998
Ulloa Ulloa & Jorgensen
Acanthosyris annonagustata from Ecuador
85
Figure1.AcanthosyrisannonagustataC.Ulloa&P.Jprgensen.—A.Habit,floweringbranch.—B.Apicalportionof
inflorescence.—C.Openbud.—I).Open(lower.—E.Fruits,withlongitudinalsection.(A—I),basedonAulestiaetal.
IA18:E,basedonAulestiaAGrefa232andl)ik522.)
uate and slightly decurrent at base, margin flat to
slightlyrevolute,paperywhenyoung,ehartaceous
when mature, glabrous with a few scattered hairs
onthemidnerveandpetiolewhenyoung,soonglabrescent,abovedryingbrownorolive,lustrous,be¬
low dull, midnerve impressed or flat above, raised
androundedbelow,secondarylateralnerves4—7
per side, raised on both surfaces, tertiary venation
reticulate,visibleonbothsurfaces;petiole4—9mm
long,flattoslightlycanaliculateabove,roundedbe¬
low. Several spikes generally clustered at the base
of young branches, or 1 or 2 below the leaves, 2.1 —
6.8 cm long, rachis densely golden brown puberulentintheupperhalf,glabrescent,darkbrownand
lustroustowardthebase,14—24-flowered,bracteate; bracts 1.5—2.5 mm long, sparsely to densely
pulverulent.Flowerscampanulate,greenishcream,
5-7mmdiam.,denselycoveredwithminutegolden
86
Novon
brown hairs on the outside; the tepal lobes 5(6),
triangular,2.0—2.5nunlong,glabrouswithinexcept
for a central tuft of long hairs with sticky tips that
adheretotheadaxialsideofthestamens;stamens
5(6),oppositethetepals,1.3-1.5nunlong,inserted
near the base of the tepals, filaments flat, tapered
toward the anther; nectary with 5(6) oblong lobes
alternating with the tepals, 1.5—1.8 mm long, mi¬
nutely puberulent; style 1.2-1.8 mm long, stigma
trilobed. Infructescence rachis to 8.5 X 0.4 cm,
glabrescent,striate;fruitdrupaceous,obovoidtoel¬
lipsoid, 2.9-4' X 2.2—2.8 cm, yellow or orange-yel¬
low, crowned by the remains of the tepals, nectary
and style, which form a depressed scar ca. 3 mm
diam., the exocarp woody, 3-4 mm thick, the outer
surfaceminutelygranulose,denselycoveredwith
golden brown hairs ca. 0.25 mm long, mesocarp
white or light orange; seed single, obovoid, to 3.2
X1.6—1.8cm.
nodocumentationofhemiparasitisminthisspecies,
although it is probably a root hemiparasite, as are
other species in the genus (Barroso, 1968) and the
family (see Kuijt, 1969).
Acanthosyrisannonagustataischaracterizedby
itsellipticleavesandgoldenbrownpuberulentin¬
florescence and fruits. It is the only species that
haspubescentfruitswhenmature.Theseareofme¬
dium size in the genus: A. asipapote and A. pauloalvinii G. M. Barroso from Brazil have fruits almost
twice as large, while the other species have fruits
generally less than 2.5 cm long. The inflorescence
and flowers are similar to those of A. glabrata and
A.asipapote,buttheindumentoftherachis,bracts,
and flowers is golden brown in the new T species,
versuswhitishgrayinA.glabrata,andferrugineous
(bracts) and pale (rachis and flowers) on A. asipa¬
pote.Furthermore,someoftheflowersofA.annon¬
agustata have six tepals, stamens, and nectary
lobes, while only five have been reported for the
other species in the genus. The leaves are elliptic
in A. annonagustata, as in A. paulo-alvinii. This is
the only species in the genus that grows in tropical
rainforestandhasfruitsknowntobeeatenbymon¬
keys.
Etymology,commonnames,anduses.According
to Dik 522 the fruit is very sweet with a taste of
“GuanAbana” (Annona muricata L.), hence the
name of this new species. The Huaorani Amerin¬
dians call it “Aguencatue” (Aulestia & Goriti 2053)
or “Oreclamohue” (Aulestia et al. 1318) and pre¬
pareateatoeliminateintestinalworms(Aulestiaet
al.1318).
Distribution, habitat, and dispersal. This spe¬
cies has been collected in eastern Ecuador along a
roadopenedbyanoilcompanyintheAmazonrain¬
forest at the Yasunf National Park and Huaorani
Ethnic Reserve, Napo province, at ca. 250 m ele¬
vation.MaterialinflowerhasbeencollectedinDe¬
cember and in fruit in March, July, September, and
December.Thisnewspeciesiscommoninthearea
wherespider(Ateles)andwoolly(Lagothrix)mon¬
keysfeedonthefruitsandpresumablyaretheprin¬
cipal seed dispersal agent (D. Neill, pers. comm.).
According to Dawson (1944), fruits of the Argen¬
tineanspeciesofAcanthosyrisareeatenbymam¬
mals and birds, but seed dispersal has not been
verified.FruitsofA.glabrataarerelishedbyagouti
(Dasyprocta) (Smith, 1950), and the fruits of A. asipapote are generally eaten by wild animals (Nee,
1996). As this new species has been reported hav¬
ingatastyfruit,presumablyotherlargeAmazonian
frugivoresconsumeanddisperseit.GaloTipaz,an
Ecuadorianbotanist,andhisassistantsgerminated
seedsaspartoftherevegetationoftheMaxuspipe¬
line road, and the plants were established in the
revegetatedareasofthatroad,whereatleastsome
of them survived (I). Neill, pers. comm.). There is
Paratypes.F.CUADOR.Napo:ParqueNacionalYasunf,carreterayoleoductotieMaxuseneonstruccionkm
40,ParcelapermanenteNo.10,235m,10Dec.1904(fr),
M.Aulestia2987(QCNE);km46,244m,17Sep.1993
(fr), A. Dik 522 (COL. F, MO, QCNE, USM); km 10-12.
250m.10July1993(fr).O'.Tipaz2724(QCNE);Keserva
etnicaHuaorani,earreterayoleoductodeMaxusencon¬
struccionkm67—69.250m,1—3Dec.1993(fl).M.Au¬
lestia.A.Audi&E.Nenquerei1318(AAU,COL,OILMO.
QCA,QCNE.US.ISM);km92-96,250m,20Mar.1994
(fr)..1/.AulestiaA().Conti2053(LPB.MO.NV.QCA,
QCNE,
S).
Acknowledgments. We thank Roy Gereau for
correcting the Latin description, Henk van tier
Werll for revising the manuscript, David Neill anti
Hugh litis for sharing their observations with us,
and Job Kuijt and Michael Nee for their review
comments.
Literature
Cited
Barroso,G.VI.1968.Acanthosyrispaulo-alvinii—uma
novaespecietieSantalaceae.AnaisSoc.Bot.Brasil
(XIXCongressoNacionalticBotanica1968):107—109.
Dawson,0.1944.LasSantalaceasargentinas.Revista
Mus.LaPlata,Secc.Bot.4(23):1-80.
Kuijt.J.1969.TheBiologyofParasiticFloweringPlants.
Univ.
California
Press,
Berkeley.
Nee,M.1996.AnewspeciesofAcanthosyris(Santala¬
ceae)fromBoliviaantiakeytothewoodySouthAmer¬
ican
Santalaceae.
Brittonia
48:
574—579.
Smith,A.C.1950.AColombianspeciesofCervantesia
R.&P.Trop.Woods51:12-14.
A
New
Species
of
Ceratozamia
(Zamiaceae)
from
Veracruz,
Mexico
MarioVazquez-Torres
Institute tie Investigaciones Biologicas, Universidad Veracruzana,
Apdo. Postal 294 Xalapa, Veracruz, 9100 Mexico
Andrew P. Vovides
Institute <le Ecolegfa, A.G., Apdo. Postal 63, Xalapa, Veracruz, 91000 Mexico
ABSTRACT. Ceratozamia morettii is described
fromacloud-forestenvironmentinVeracruz,Mex¬
ico. The main morphological characters are illus¬
trated,andcommentsonrelatedspeciesaremade.
The new species differs from others in the genus
by the near prostrate habit, circinate vernation of
the leaves, and wide leaflets with translucent ve¬
nation. This taxon is apparently related to a group
of species that are relatively small trunked,
branched,producefew7leaves,andhaverelatively
small strobili. The non-sympatrie species of the
group also inhabit moist to very moist habitats, as
in the case of C. miqueliana, C. microstrobila, and
C.mexicanavar.robusta.
Ceratozamia morettii Vazquez-Torres & Vovides,
sp. nov. TYPE: Mexico. Veracruz: 7 Jan. 1992,
M.Vdzquez-Torres&H.Barney4097(holotype,
GIB; isotypes, GIB, MEXU, XAL). Figure 1.
Triincussemihypogaeus,humilisad30cmalius;folia
vernatacircinata,pauca,usque10,glabra.Foliolasubopposilavelalterna,12—25juga,remota,linearisvelfaleata,
translucida,
tenuia,
basicuneata.
Plant palm-like, trunk erect to procumbent,
short,globosetocylindrical,semihypogeousupto
30 cm long, 8 cm diam., typically with 1^4 branch¬
escoveredwithpersistentcataphyllandleafbases;
cataphylls triangular, 2 cm wide at base, 2.6 cm
long; vernation circinate; leaves light green,
pubescent when juvenile, decurrent to prostrate
forming an open crown with a maximum of 10(4—
7) leaves per crown, 1-1.4 m long, 40—65 cm wide;
leaflets12—25pairs,ovoidwhenimmature,becom¬
inglineartofalcateuponmaturity,chartaeeous,ve¬
nationparallel,translucid,paleyellow,dichotomous
principally in the lower third of leaflet, 25—35 cm
long at median position of leaf, distal portion un¬
evenlysinuous,apexacute,baseattenuate,2.7^4.8
cm wide, petiole terete or subterete, 45-60 cm
long,armedwithshortstoutprickles;microstrobilus
typicallyconiform,elongate,yellowishgreen,10—
15 cm long, 2.5^4 cm diam.; peduncle terete, 5-7
cm long, l cm diam., reddish brown tomentum; microsporophyllseuneate,10—12mmlong,8—9mm
wide,distalendwithtwoerecttocurvedprominent
coniform
protuberances;
microsporangia
numerous,
generally in sori of three covering Vi to % of abaxial
surface, dehiscence longitudinal; megastrobilus
coniform,greenwhenjuvenileturningbrownatma¬
turity, 12—16 cm long, 4.5—5 cm diam.; peduncle
terete, dark brown tomentose, 5—7 cm long, 1 cm
diam.;megasporophyllspeltate,reddishbrown,dis¬
tal end almost hexagonal with two erect or curved
corniform protuberances; ovules ovoid, two per
megasporophyll;seedsirregularlyovoidwithoutany
definedfaces,sarcotestayellowishwhitewhenim¬
mature turning to gray brown, delicately papyra¬
ceous and transparent when mature, sclerotesta
hard, light gray, 1.5—1.8 cm long, 1.2 cm diam.;
chromosomenumber2n=16.
ChromosomalStudies
ThechromosomenumberandkaryotypeofCer¬
atozamiamorettiiweredeterminedfromthreees¬
tablished specimens held at the Jardin Botanieo
Feo. J. Clavijero (Botanic Garden of the Instituto
deEcologfa)undertheaccessionnumbers81-397,
81-857, and 81-852; vouchers are deposited at
XAL. The root tip mitosis technique used was that
describedbyVovides(1983),andthechromosome
classificationbasedoncentromerepositionwasthat
of Levan et al. (1964) modified by Schlarbaum and
Tsuehiya (1984). The diploid idiogram (Fig. 2) was
constructed by taking the average arm lengths of
the best three metaphase cells examined (Fig. 3).
PhotomicrographywasdoneonaZeissphotomicroscope (Fomi III) equipped with phase contrast op¬
ticsandplanapochromaticobjectives.Printswere
madeusingKodakbromidepaper.Armlengths,to¬
talchromosomelength,chromosomeindex(short
arm divided by long arm), and symmetry index
(lengthoflongestpairdividedbylengthofshortest
pair)werecomputedusingtheaveragearmlengths
from the three metaphase cells (Table 1). The
Novon 8: 87-90. 1998.
88
Novon
FigureI.a-i,CeratozamiamorettiiVazquez-Torres&Vovides.—a.Hal>itofplant.—b.Circinatevernationofleaf,
petiole,andtrunk.—e.Leafletarticulationsandrachis.—<1.Detailofleafletveins.—e.Non-expandedmalestrobilus.
—f.Abaxialviewofmicrosporophyllshowingdehiscedsporangia.—g.Femalestrobilus.—h.Megasporophyllwith
immature
ovules.
—i.
Seed.
Volume 8, Number 1
1998
Vazquez-Torres & Vovides
CeratozamiamorettiifromMexico
89
1'igure2.DiploididiogramolCeratozamiamorettii(2n
= 16), bar = 2 /ant.
karyotype shows 12 median region (m) chromo¬
somes,1submedian(sm),1sub-terminalregion(st)
chromosome, and 2 terminal point (T) chromo¬
somes. A maximum of 5 satellites were recorded
butwerenotconsideredinthecalculations.
Habitat
This cycad occurs in cloud forest on humus-rich
grayish yellow clay soil of volcanic origin. The
plants are found on steep 45-60° slopes or on ver¬
tical rocky walls of loose, weathered basalt, at an
elevation of 1200 to 1400 m. The closest climato¬
logical station to this habitat has recorded an av¬
erage temperature of 17.3°C and over 1900 mm
annual
precipitation.
Other vascular plantsassociatedwith thecycad
in this vegetation type are typical of cloud-forest
species distributed on the windward slopes of the
Sierra Madre Oriental facing the Gulf of Mexico.
These are: Alnus jorullensis Kunth, Clethra mexicana DC., Dendropanax arboreus (L.) Decaisne &
Planchon,DichsoniagiganteaMaxon,Ilexdiscolor
Hemsley,LiquidambarmacrophyllaOersted,Mag-
I'igure3.MitoticmetaphasecellofCeratozamiamoret¬
tii , bar = 4 gm.
nolia schiedeana Schlechtendal, Marattia laxa
Kunze,Oreopanaxcapitatus(Jacquin)Decaisne&
Planchon, Ostrya virginiana (Miller) K. Koch, PodocarpusguatemalensisStandley,Quercusgermana
Chamisso & Schlechtendal, Q. laurina Humboldt
&Bonpland, Q. xalapensisHumboldt &Bonpland,
Table1.Karyotypicdataatmetaphaseinroot-tipmitosisofCeratozamiamorettii(meanofthreemetaphasecells).
Pair
index=0.70.
90
Turpiniainsignis(Kunth)Tulasne,andUlmusmexicana(Liebmann)Planchon.
Discussion
Thespecificepithetwaschoseninrecognitionof
the scientific contributions of Aldo Moretti in the
field of cycad biology. Prof. Moretti is a researcher
of the Orto Botanico of the University of Naples,
Italy.
Ceratozamia morettii differs from the rest of its
congeners by the circinate vernation of its leaves.
Like C. microstrobila, C. morettii has few nearly
prostrateleavespercrownandprofusebranching
of the trunk. Ceratozamia morettii belongs to the
group of species having wide leaflets: i.e., C. eu ryphyllidiaVdzquez-Torres,Sabato&Stevenson,C.
microstrobila Vovides & Rees, C. hildae Landry &
Wilson, C. miqueliana H. Wendland, C. latifolia
Miquel, C. mexicana var. robusta (Miquel) Dyer,
and C. whitelockiana Chemnick & Gregory. With
the exception of C. microstrobila, C. hildae, and
perhaps C. mexicana var. robusta, the other taxa
are distributed south of the neovolcanic belt of
Mexicoinmoisthabitats.
ThefollowingkeyseparatesCeratozamiamorettii
fromotherCeratozamiaspecieswithwideleaflets.
DiagnosticKit
la. Emerging
leaves
presenting
circinate
vernation,
leaves
spreading,
leaflets
ovoid
when
immature,
linear,falcatetosubfalcate,notgreaterthan5
cm
wide.
C.
morettii
lb. Emerging
leaves
not
presenting
circinate
verna¬
tion,leavesascendingorspreading,leafletslan¬
ceolate,elliptic,obovate,orbroadlyoblanceolate.
2a. Persistentleafbasesdarkbrown,notop¬
pressed
to
trunk.
3a.
Leaflets
coriaceous.
4a.Leafletslanceolate,lessthan4cm
wide. C. mexicana var. robusta
4b.
Leaflets
not
lanceolate.
5a.Leafletsobovatetobroadlyoblaneeolate,greaterthan4cm
wide.
C.
miqueliana
5b.Leafletselliptictooblanceolate.lessthan4.5cmwide..
.
C.
latijolia
3b.
Leaflets
papyraceous.
6a.Leafletslinearlanceolate,lessthan
4 cm wide. C. whitelockiana
6b.Leafletsbroadlyoblanceolate,
greater than 8 cm wide .
C.
euryph
vllidia
Novon
2b.Persistentlealbaseslightbrown,tightlyappressed
to
trunk.
7a.Leafletspinnate,elliptictolanceolate
.
C.
microstrobila
7b.Leafletsfasciculate,lanceolate..C.hildae
Thechromosomecountandkaryotypearecon¬
sistent with that reported for the genus (2n = 16)
byMarchant(1968),Vovides(1983,1985),Vovides
et al. (1993), and Moretti (1990). The karyotype of
C. morettii (12m + Ism + 1st + 2T) is nearly
typical for the genus Ceratozamia (12m + 2sm +
2T) and appears to be stable within the genus (Vo¬
vides et al., 1993; Moretti, 1990). Satellite number
andpositionappeartovarywithcellsobservedand
much care is needed in recording them. A maxi¬
mumoffivewererecordedandmanycellsshowed
three.
Thepreciselocalityhasbeenintentionallyomit¬
tedtodiscourageindiscriminatecommercialcol¬
lectingofthisendangeredspecies,whichcouldre¬
sult in its extinction. The common names of this
speciesare“tepetmaizte,”or“tepemaizte”(forest
maize), because of a similarity between the corn¬
cobandthecycadcones.
Acknowledgments.WethankManuelEscamilla
for the botanical illustration of this species. This
research was partially funded by CONACyT pro¬
jects No. 0063-N9106 and 1837P-N9507, I.E. Sistematica Vegetal project No. 904-14 and U.V.
F641-M940.
Literature
Cited
Levan,A.,K.Fredga&A.A.Sandberg.1964.Nomen¬
clature
for
centromerio
position
on
chromosomes.
Ilereditas52:201-220.
Marchant,C.J.1968.Chromosomepatternsandnuclear
phenomenainthecycadfamiliesStangeriaceaeandZamiaceae.
Chromosoma
(Herb)
24:
100—134.
Moretti,A.1990.Karyotypicdataonnorthandcentral
American
Zamiaeeae
(Cycadales)
and
(heir
phylogenet¬
icimplications.Amer.J.Bot.77:1016—1029.
Schlarbaum,5.E.AT.Tsuchiya.1984.Thechromosomes
ofCunninghamiakonishi,C.lanceolalaandTaiwania
cryptomeroides
(Taxodiaceae).
PL
Syst.
Evol.
145:
169—
181.
Vovides,A.P.1983.SystematicstudiesontheMexican
Zamiaeeae
I.
Chromosome
numbers
and
karyotypes.
Amer.J.Bot.70:1002-1006.
-. 1985. Systematic studies on the Mexican Zamiaceae
11.
Additional
notes
on
Ceratozamia
kuesteriana
fromTamaulipas,Mexico.Brittonia37:226-231.
-. M. V azquez-Torres, B. Schutzman A C. G. Iglesias.1993.AnewspeciesofCeratozamia(Zamiaeeae)
fromQueretaroandHidalgo,Mexico.Novon3:502—
506.
Trifolium
jokerstii
(Leguminosae,
from
Butte
Papilionoideae),
County,
a
New
Species
California
Michael A. Vincent
W. S. Turrell Herbarium, Department of Botany, Miami University, Oxford, Ohio 45056, U.S.A.
Randall Morgan
3500 Main St., Sequel, California 95073, U.S.A.
Abstract. Trifolium jokerstii (Leguminosae, Pa¬
pilionoideae), a new species from Butte County,
California, is described and illustrated. It is mor¬
phologically similar to T. barbigerum and T. grayi,
from which it differs in stipule shape, flower color,
seed size, and lack of pubescence.
ThegenusTrifolium(Leguminosae,Papilionoi¬
deae)consistsofapproximately240speciesfound
inmainlytemperateandmontaneregions,withar¬
easof“Mediterranean”climate(theMediterranean
basin,California,andChile)beingconsideredcen¬
ters of diversity in the genus (Zohary & Heller,
1984). In North America, 93 species are known
(Kartesz, 1994), of which 64 are native and 29 are
introductions. Isely (1993) recognized 45 species
in California, 32 of which are native to the state;
ofthese,14belongtosectionInvolucrarium,aNew
Worldendemicsectioncharacterizedbythepres¬
enceofaninvolucreoffusedbractssubtendingthe
inflorescence.
During the course of a revision of Trifolium bar¬
bigerum Torrey and related species of section In¬
volucrarium,severalspecimensfromthevicinityof
North Table Mountain, Oroville, Butte County, in
north-centralCalifornia,weredeterminedtorep¬
resent an undescribed species. A single collection
was examined by Isely and mentioned in his treat¬
ment of the genus for the Jepson Manual as a yel¬
low-floweredvariantofTrifoliumbarbigerumvar.
andrewsii A. Gray (Isely, 1993). The new species
wasalsomentionedbyOswaldandAhart(1994)as
possiblythesamevariety,withacommentthatthe
plants might deserve taxonomic recognition. No
collections of this taxon were mentioned by Mc¬
Dermott (1910) or Zohary and Heller (1984) in
their monographs of the genus. The species is sig¬
nificantlydifferentfrombothT.barbigerumandT.
grayi Lojacono (71 barbigerum var. andrewsii A.
Gray) and is described here as new.
Trifolium jokerstii Vincent & R. Morgan, sp. nov.
TYPE: U.S.A. California: Butte County, North
Table Mountain, N of Oroville, 29 Mar. 1996,
M. A. Vincent 7227, Rhode & Snowden (holotype, MU 177695; isotypes, F, ISC, MO, NY,
RSA, UC, US). Figure 1.
Trifoliumannuum,cauliserectisvelascendentietsimplicovelranioso,glabro;stipulisovatis,serratis,persistentibus;
foliolis
elliptieis
vel
obovatis,
serrulatis;
capitulis
semiglobosis;
pedunculis
foliis
longioribus;
involuoris
lobatis;
lobis
ovatis,
dentatis;
calycibus
campanulatis,
pubescentibus,tenuis;lobissimplioibussubulatis,plumosis;
corollisluteisadsulphureas.Affinis7.barbigeroTorrey
el/.'grayiLojacono;abutroquecaulibusetloliisglabris,
stipulisovatis,serratis,corollisluteisadsulphureas,et
seminibus
3.1—3.4
mm
longis
differt.
Erect-ascendingannualwithsimpletobranched
glabrous stems to 20 cm, from a fibrous taproot.
Stipules thin, pale to green, 8-13(—20) X 7—10
mm, rounded to acute and serrate at apex, persis¬
tent, adnate to tlie petiole, distinct or sometimes
basally fused into a cylinder for % their length.
Leaves trifoliolate, petioles to 90 mm, glabrous.
Leafletssessile,glabrous,serrulatetorarelynearly
lobed,elliptictoobovate,(5—)8—17(—32)X(4—)6—
8(—15) mm, with or without a prominent white to
dark purple chevron. Inflorescence involucrate,
subglobose,10—30-flowered,12—30mmwide,pe¬
dunclelongerthantheleaves;involucrewide-campanulatetonearlyflat,(13—)15—17(—22)mmwide,
glabrous,lobed,thelobesrounded,toothed.Calyx
7—9 mm long, expanding in fruit, tube membra¬
nous,sparinglypubescent,campanulate,5-nerved,
oblique;teethplumose,nearlyaslongastoslightly
longer than tube, subulate-setaceous, the upper
shorterthanthelower,simple,lateralteethsimple
to bifid, lower tooth bifid to trifid. Corolla 10-15
mmlong,golden-yellowtosulphur-yellow;standard
broadly ovate, inflated in fruit with a constricted
throat above the mouth of the calyx; wings auriculate, longer than keel; keel with or without a pur¬
ple spot on each side. Ovary glabrous, 4—5 mm
Novon 8: 91-93. 1998.
92
Novon
■.
figure1.TrifoliumjokerstiiVincent&K.Morgan.—A.Stipules(fromAliurl7569<£•Cunningham).—15.Leaflet
showingteethantichevron(fromtheholotype).—C.foldedinvolucre,showingrounded,toothedlobes(fromthe
holotype).—I).Inflatedfruitingcalyxandcorolla(fromMorgan2566c).—K.field-collectedseedsofTrifoliumjokerstii
(right),and/.’grayi(left),showingsi/edifferences.
Volume 8, Number 1
1998
Vincent&Morgan
TrifoliumjokerstiifromCalifornia
long, ovules 2. Fruit 3.3—3.5 X 2.0—2.2 nun, stipitate, ovoid, (1—)2-seeded. Seeds 3.1—3.4 X 2.3—
2.7 mm, dark brown, somewhat rough. Flowering
Mareh-May.
I’aratypes.U.S.A.California:UniteCounty,North
'lahleMountain,NofOroville,2Apr.1995,/,.Abort7569
&Cunningham(Cl15(7MU).30May1995,I..Aliarl7583
(CHSC), 29 Mar. 1996, M. A. Vincent 7205. Rhode &
Snowden(GH.ISC,MO.MU,USA),7215(CM.ISC,MU,
USA.UC),7219(MU),7240(MU):alongCottonwood
Boat I, N of Oroville, 7 Apr. 1989, I.. Abort 0202 (CAS.
CHSC.MO.U.Morganpersonalherbarium.UC),29Mar.
1985,./.I).Jokerst2180(ISC),29Mar.1996.M.A.Vin¬
cent7245.Rhode&Snowden(F,ISC.MO.MU,USA.UC,
US):cultivatedplant,12May1995.R.Morgan2566c
(MU.
U.
Morgan
personal
herbarium).
TrifoliumjokerstiihasaffinitieswithT.barbigerum and T. grayi. It differs from the former in the
larger size of all parts. It can be distinguished from
bothspeciesbyflowercolor,lackofpubescenceon
thestemandfoliage,stipuleshape,andseedtraits.
Flowers of T. barbigerum and T. grayi are lavender
to purple with white to cream tips, or rarely all
white. Stipules of T. grayi are acute to attenuate at
the apex, with very large, jagged teeth, while stip¬
ules of T. jokerstii are rounded to slightly acute,
with smaller teeth. Seeds of T. grayi measure 1.6—
2.0 X 1.4—1.5 mm and are pale brown and mottled,
whilethoseofT.jokerstiiaremuchlarger(thereare
no intermediates) and are dark brown. Petioles of
the cotyledons of T. jokerstii are from 13 to 25 mm
long, while those of the cotyledons of T. grayi are
3—10mmlong.
ThetwoknownpopulationsofTrifoliumjokerstii
are in Butte County, north of Oroville, and are at
least 100 miles from the closest known population
ofT.grayi.
Allozyme banding patterns (Vincent, in prep.)
are vastly different in T. jokerstii from those of both
T. barbigerum and T. grayi, and bear out the dis¬
tinctnessofthisspecies.
The new’ species is named in honor of the late
James D. Jokerst [1956-1995 (Beedy & Preston,
1996)],whocollectedtheearliestknownspecimen
of the clover and published a flora of North Table
Mountain (Jokerst, 1983). The common name
“Butte County Golden Clover” was coined for the
taxon by Oswald and Ahart (1994).
93
Acknowledgments.Thefollowingpeopleassist¬
ed with this project: Lowell Ahart, R. James Hick¬
ey, Julie Horenstein, Vernon Oswald, Delores and
EdwardRhode,andJamesSnowden.Thefollowing
herbaria graciously lent specimens: CHSC, ISC,
MO, l)C. The W. S. Turrell Herbarium Fund (Ml!)
supportedfieldworkassociatedwiththisproject.
Literature
Cited
Beedy,K.C.&U.K.Preston.1996.JamesDentJokerst.
Amer.
Soe.
PI.
Taxonomists
Newslelt.
10(1)
unpaginated
(electronic
publication).
Iselv.I).1993.Trifolium(Clover).Pp.646—655inJ.C.
Hickman(editor).TheJepsonManual:HigherPlantsof
California.
Univ.
California
Press.
Berkeley.
Jokerst.J.I).1983.ThevascularfloraoffableMountain.
ButteCounty,California.Madrono30:1—18.
kartesz.J.T.1994.ASynonymizedChecklistoftheVas¬
cularFloraoftheUnitedStates.Canada,andGreen¬
land.2nded.TimberPress,Portland.
McDermott.L.F.1910.AnIllustratedkevtotheNorth
AmericanSpeciesofTrifolium.Cunningham,Curtis&
Welch.
San
Francisco.
Oswald,\.II.&I..Ahart.1994.ManualoftheVascular
PlantsofButteCounty,California.CaliforniaNative
Plant
Society,
Sacramento.
Zoliary.M.vS,I).Heller.1984.TheGenusTrifolium.Israel
Academy
of
Sciences,
Jerusalem.
New
Taxa,
New
Combinations,
(Poaceae:
and
Observations
in
Kengyilia
Triticeae)
Chi Yen and Jun-Liang Yang
Triticeae Research Institute, Sichuan Agricultural University,
Dujiangyan City 611830, Sichuan, China
Bernard R. Baum
Eastern Cereals and Oilseeds Research Centre, Agriculture and Agri-Food Canada, Central
Experimental Farm, K.W. Neatby Building, Ottawa, Ontario, Canada, K1A 0C6
ABSTRACT. A new species is described and five
new combinations are made in Kengyilia. The new
taxa are Kengyilia eremopyroides and K. batalinii
var.villosissima.KengyilialongiglumisandK.nana
are reduced to K. alatavica var. longiglumis and K.
nana to K. batalinii var. nana, respectively. Roegneria carinata, Elytrigia kryloviana, and Elytrigia
pulcherrima are transferred to Kengyilia as K. car¬
inata, K. kryloviana, and K. pulcherrima, respec¬
tively.
We first described Kengyilia Yen & J. L. Yang
in 1990 (Yen & Yang, 1990) and soon afterward
added a new species to it (Baum et al., 1991). Sub¬
sequently, we provided a synopsis and key to the
16 species of the genus known to us at the time
(Yang et al., 1992). Cai and Cui (1995) have since
addedtwomorenewspecies.Thisgenus,nowwith
approximately 20 species, is distributed primarily
inwestChina,withsomespeciesfoundincountries
west of China. We recently investigated the taxo¬
nomicrelationshipswithanddifferencesbetween
its congeners Roegneria, Elymus, and Agropyron
(Baum et al., 1995) and provided a key to identify
thesefourgenera.Wearecurrentlypreparingatax¬
onomicmonographofKengyilia.Towardthisgoal
wearesearchingfortypematerialindifferenther¬
baria.
ArecentvisittotheKomarovBotanicalInstitute,
St.Petersburg,Russia,duringthesummerof1995,
by Yen and Yang was carried out to study type ma¬
terialofKengyilia.Morphologicalexaminationsof
specimens,especiallytypematerial,ledtothecon¬
clusion that a number of species in Kengyilia were
hithertomisnamedandthatotherspeciesneeded
to be incorporated within Kengyilia. The purpose
ofthispaperistodocumentthenewcombinations
and revisions in Kengyilia resulting from the ex¬
aminationscarriedoutduringtherecentvisittoLE.
Novon 8: 94-100. 1998.
Materials and Mkthods
Morphologicalexaminationswereperformedon
herbarium materials, including type collections,
from the following herbaria: JSBI, K, LE, NUBl),
PE, SAUTI, TK. Characters on these specimens
were evaluated lor quantitative and qualitative
measurementsandcomparedtovariouspertinent
protologues
for
taxonomic
assessment.
Results—Taxonomic Treatments
1. Kengyilia alatavica (Drobov) J. L. Yang, Yen
& Baum var. longiglumis (Keng & S. L. Chen)
Yen, J. L. Yang & Baum, comb. nov. Basionym:
Roegneria longiglumis Keng & S. L. Chen,
Acta Nanking Univ. (Biol.) 1: 83. 1963. Ken¬
gyilia longiglumis (Keng & S. L. Chen) Yang,
J. L„ Yen & Baum, Hereditas 116: 27. 1992.
TYPE: China. Gansu, on slopes, alt. 2500 m,
July 6 1937, T. P. Wang No. 7080 (holotype,
PE). Figure 1A.
Distribution. China: Gansu, Xiahe; Xinjiang,
Yecheng, Taxkorgan and between Wuqia and Turugar, on dry slopes and scree, alt. 2500—3340 m.
ThistaxonwasknownhithertoasK.longiglumis,
fromtheChinesematerialthatwasexamined.Our
comparative studies proved that K. longiglumis is
a taxonomic synonym of K. alatavica. The Chinese
materialishererecognizedasaseparatevariety;it
differs from K. alatavica var. alatavica by its hairy
leaf sheaths and culms, and by the short rounded
lodicules (Fig. 1A). The pointed lodicules of the
typical variety are shown in Figure IB.
2. Kengyilia batalinii (Krassnov) J. E. Yang, Yen
& Baum var. villosissiina Roshevitz ex Yen,
J. L. Yang & Baum, var. nov. TYPE: Turkestan.
Pamir, near Karakuli Lake, July 5 1901 [col¬
lector&collectionno.unretained](holotype,
LE). Figure 2.
Volume 8, Number 1
1998
95
Yen et al.
New Taxa in Kengyilia
6
E
Figure1.LodieulesofK.alatavica(Drol>ov)J.I,.Yang,Yen&Baum.—A.Typicallodiculesofvarietylongiglumis
(Keng&S.L.Chen)Yen,J.L.Yang&Baum.—B.Lodiculesfoundinvarietyalatavica.
Kengyliaebatalinii(Krassnov)J.L.Yang,Yen&Baum
affinis,sedlaminissupernisdensevillosis,laminisinfernisglabrisvelsparsimciliolatis,spieisdensealbus-villosis,
glumis
dense
pubescentibus,
lemmatibus
dense
vil¬
losis
differt.
This variety differs from the typical variety by its
leaf blades villous on the upper surface and gla¬
brous or sparsely ciliate on the lower surface, the
spikes densely white villous, and the glumes and
the lemmas densely villous. See key below to iden¬
tify this variety and the other two in K. batalinii.
Distribution.Turkestan.
3. Kengyilia batalinii (Krassnov) J. L. Yang, Yen
& Baum var. nana (J. L. Yang, Yen & Baum)
Yen, J. L. Yang & Baum, comb. nov. Basionym:
Kengyilia nana J. L. Yang, Yen & Baum, Canad. T Bot. 71: 339-345. 1993. TYPE: China.
Taxkorgan, Pamir Plateau, in alpine steppe,
alt. 4200 m, Sep. 6 1987, C. Yen et al. 870502
(holotype,SAUTI).
This variety differs from the typical variety in
having leaf sheaths with densely pubescent or cil¬
iate margins, leal blades that are densely hairy on
the adaxial surface and shortly pubescent on the
abaxial surface, and slightly smaller spikes. It
growsathigherelevationsthanvarietybatalinii.
Distribution. China.
This taxon was known hitherto as K. nana, now 1
an established taxonomic synonym of K. batalinii.
The three varieties in K. batalinii may be distin¬
guishedbythefollowingkey.
la. Leaf blades glabrous.var .batalinii
lb.Leafbladesbesetwithhairsatleastonupper
surface.
2a.Leafbladesvillousonuppersurface,gla¬
brousorsparselyciliateonlowersurface
.var.
villosissima
2b.Leafbladeswithshortpubescenceonupper
surface,andwithdensehairsonlowersur¬
face
.var.
nana
4. Kengyilia carinata (Ovczinnikov & Sidoren¬
ko) Yen, .1. L. Yang & Baum, comb. nov. Bas¬
ionym: Roegneria carinata Ovczinnikov & Si¬
dorenko, FI. Tajik SSR 1: 505 (No. 310). 1957.
TYPE: Russia. In the lower part of the north¬
ern slope of Jiptyk valley—a tributary of the
Isfar, 28 June 1938, Mikeshin, G. No. 79 (ho¬
lotype, LE). Figure 3.
This species differs from K. alaica in having
sturdy,denselytuftedculmswith4—5nodes,broad
leaves,densespikes,antiglumessparselycovered
withlonghairs.
Tzvelev (1976) treated R. carinata Ovcinnikov &
Sidorenko as a synonym of Elytrigia batalinii
subsp. alaica (Drobov) Tzvelev, based on the Mi¬
keshin specimen of 26 July 1938, No. 79, probably
bymistakeconfusingitwithNo.81.Theherbarium
of the Komarov Botanical Institute (LE) has two
96
Novon
alaicumandRoegneriacarinataaredifferentspe¬
cies.
5. Kengyilia eremopyroides Nevski ex Yen, J.
L. Yang & Baum, sp. nov. TYPE: China. “Lacus
Orin-Nor sole argilloso-sabuloso porce,” alt.
3962 m (13000 ft.), 10/30 July 1884, N. M.
Przewalski No. 339 (holotype, LE). Figure 4.
£
o
Figure2.SpikeletofKengyiliabatalinii(krassnov)J.L.
Yang,Yen&Baumvar.villosissimaRoshevitzexYen,J.
I,.Yang&Baum,withthehair)'glumesandlemmas.Left,
upper
glume.
sheets collected by J. B. Mikeshin. One is the type.
The other is No. 81, collected on 10 Aug. 1938.
Nevski annotated the type as Roegneria abolinii
(Drobov) Nevski f. breviaristata Nevski. In 1956, P.
N.OvczinnikovannotateditasRoegneriacarinata.
ThesubsequentyearhepublishedR.carinatawith
Sidorenko. Thus, No. 79 is the holotype. In 1958
MelderisannotatedthisspecimenasAgropyroncarinatum. Specimen No. 81 is a loosely caespitose,
delicategrasswith1-2nodes,andculmandnarrow
leaves situated at the base of the plant, identified
as Kengyilia alaica (Drobov) J. L. Yang, Yen &
Baum. In March 1959 Tzvelev affixed the following
annotation on this specimen “Roegneria carinata
Ovczinnikov et Sidorenko Topotype = Agropyron
alaicum Drob." We agree that it is “A. alaicum Drobov,” but it is not “ R. carinata,' 1 ' i.e., Agropyron
Kengyiliaemelantherae(Keng)J.L.Yang,Yen&Baum
affinis,sedculmissubinflorescentiisdensepubescentibus,laminisdensepilosis,spicisbrevioribus(4^1.5cm)
etangustioribus,lemmatibusdensehirsutisnonvillosis
differ!.
This species resembles Kengyilia melanthera
(Keng) J. L. Yang, Yen & Baum but differs in hav¬
ing the culm densely pubescent below the spikes,
leafbladesthataredenselypiloseonbothsurfaces,
narrowerandshorterspikes,anddenselylonghir¬
sute(butnotvillous)lemmas.
Perennials,withshortrhizomes;culmsdensely
caespitose, erect, 31-37 cm tall, about 1.5-2 mm
diam.,glabrousexceptfortheupperintemodepu¬
bescentbelowthespike,with2nodes.Leafsheaths
glabrous;ligulesscariousmembranous,truncate,
about0.5mmlong;leafbladesflat,(1.5—)2—5.5cm
long, 2.5—3 mm wide, their adaxial surface pilose
and their abaxial surface densely pilose. Spikes
erect,oblong,(3.5—)4-^1.5cmlong,8-10mmwide;
raehisinternodesdenselypilose,theuppermostin¬
ternodes 1.5—2 mm long, the lowermost 5-7 mm
long; spikelets ovoid, slightly secund, straw or
purple colored, with 4—6 florets, 10-11 mm long
(excludingawns);rachillaintemodes0.8—1.2mm
long,denselypuberulous;glumesunequal,ovoid,
3—5-nerved, glabrous, acute, mucronate, first
glumes 4—4.5 mm long, second glumes 4.5-5 mm
long;lemmasovate-oblong,7-8mmlong,densely
hirsute,apexacuminateendingwitha3—4mmsca¬
brouslongawn;paleashorterthanlemma,0.5mm
long,withanemarginateapex,withkeelssparsely
ciliolate in their upper part; anthers black, 2 mm
long.
Distribution. China, known only from the type
locality.
The type specimen was initially annotated as a
newspeciesofWheatgrass,namelyAgropyroneremopyroides, by Nevski in 1931, but was never
published.
Volume 8, Number 1
1998
Yen et al.
New Taxa in Kengyilia
97
cm
01
Figure3.Kengyiliacarinata(Ovczinnikov&Sidorenko)Yen,J.[,,Yang&Baum.—A.Matureplant.—B.Spikelet.
—C.Lowerglume.—D.Upperglume.—E.Floretbesetwithsparselonghairs.—F.Ventralviewofafloret.—G.
Ventralviewofapalea.
98
Novon
Figure 4. Kengyilia eremopyroides Nevski ex Yen, J. L Yang & Baum. —A. Mature plant. —B. Flag leaf with
puberulentblade.—C.Spikeletsubtendedbyhirsuteupperpartofinternode.—I).Upperglume.—E.Lowerglume.
—F.Ventralviewofpalea.—G.Ventralviewofafloret.—H.Dorsalviewofafloret.—I.Lodieulepair.—J.Ovary
with
stigma.
—K..
Anther.
Volume 8, Number 1
1998
Yen et al.
New Taxa in Kengyilia
99
Figure5.Kengyiliapulcherrima(Grossheim)Yen,J.LYang&Baum.—A.Matureplant.—B.Ventralviewofpalea,
showingthetwohyalinemembranous,triangularlargeappendagesonbothsidesoftheuppermargins.—C.Dorsal
viewofafloret.—I).Ventralview'ofafloret.—E.Spikelet.—F.Upperpartofleafsheathwithciliatemargins.
100
Novon
Table1.Different1!
6. kengyilia kryloviana (Schischkin) Yen, J. L.
Yang&Baum,comb.nov.Basionym:Agropy¬
ron krylovianum Schischkin, FI. Zapod. Sibir.
2: 353. 1928. (Animadvers. syst. ex Herb.
Univ. Tomsk No. 2). Elytrigia kryloviana
(Schischkin) Nevski, Tr. Bot. Inst. Akad. Nauk
SSSB, ser. 1, 2: 84. 1936. TYPE: Russia. West
Siberia, Altai, Valley of river Chuya (holotype,
TK).
Distribution. Stony and grassy slopes, rocks
and screes of the middle mountain belt in Russia
(West Siberia: Irtysh, Altai; East Siberia: AnganaSatan)andKazakhstan(NorthernBalkhash).
Earlier (Baum et ah, 1991), we suspected that
this species, formerly belonging to Agropyron
(Tzvelev, 1976: 145), might belong to Kengyilia.
EssentialdifferencesbetweenthisspeciesandKen¬
gyilia habahenensis Baum, Yen & J. L. Yang are
given in Table 1.
7. kengyilia pulcherrima (Grossheim) Yen, J. I..
Yang&Baum,comb.nov.Basionym:Agropy¬
ronpulcherrimumGrossheim,TiflisBot.Sada
[Moniteur du Jardin Botanique de Tiflis] 1.3—
14: 42. 1919 (also Plate 4, fig. 1—5). Elytrigia
pulcherrima(Grossheim)Nevski,Tr.Sredneaz
Univ. Ser. 17: 51. 1934. E. intermedia (Host)
Nevskisubsp.pulcherrima(Grossheim)Tzvel¬
ev, Novost. Sist. Vyssh. Rast. 10: 31. 1973.
TYPE: Turkey. “Prov. Kars, ilistr. Ardahan,
propeGuljabert,inlocisstepposis,25/7/1914,
A.Grossheim”(holotypeanilisotype,LE).Fig¬
ure5.
Agropyron
intermedium
var.
ambigens
Hausskneeht
in
HalacsyConsp.PI.Grace.3:437.1904.A.ambigens
(Hausskneeht)Boshevitz.FI.liirkin.I:191.1932.
TYPE:Greece."IterGraeeum,PindusTymphaeus
prope Malakassi. in sehistosis, 18 VII 1880, C.
Haussknecht"
(isotype,
LE).
Identity
and
Typification
of
Dracontium
dubium
Kunth
(Araeeae)
Guanghua Zhu
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Julius (). Boos
1368 Scottsdale Road Last, West Palm Beach, Florida 33417, U.S.A.
Thomas B. Croat
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. A neotype and epitype are selected for
DracontiumdubiumKunth.Dracontiumchanguango G. S. Bunting is treated as a synonym of I),
dubium.
DracontiumdubiumKunthwasfirstcollectedby
RichardSchomburgkduringanexpeditiontoGuy¬
ana (British Guiana) in 1840-1844 at the base of
Mt. Curassawaka of the Kanuku Mountains, south
of Nappi (ea. 3°22'N, 59°34'W), and several living
tubers were sent to the Botanical Garden of Berlin
in 1843 (Roth, 1922: 387-388, 1923: 103). One
plantbloomedintheGardenandwasdescribedby
Kunth in the following year (Kunth, 1844). Schott
(1860:481)notedthattherewerefertilecollections
of this species at the Berlin Herbarium (B). How¬
ever, no such specimen of D. dubium was seen by
Engler in 1911, when he only cited “Bluehte ini
Bot. Garten zu Berlin Sept. 1844” under the spe¬
cies. Schott (1860) might very likely have seen only
live specimens of the species, since at Kunth’s time
gardenplantswereoftendescribedandillustrated
without preparing herbarium specimens (Paul
Hiepko,pers.comm.).Kunthhadapparentlymade
adrawingoftheinflorescencesandsomefloralde¬
tailsofthespecies,whichwasdepositedintheBer¬
lin herbarium (Engler, 1911: 38, fig. 14A-E). More
than a decade after the discovery of D. dubium,
Schott(1857),basedonunspecifiedovaryandstig¬
ma details of the species, described the new genus
Echidnium, giving the single species a new name,
E. schomburgkii. In the following year, Schott
(1858b: pi. 88) published a plate under the name
E.schomburgkii,whichcopiedKunthsdrawingbut
addedanopenedinflorescenceandotherfloralde¬
tails. Later, when he published the first compre¬
hensiveclassificationoftheAraeeae,Schott(I860)
acceptedthenameE.schomburgkii,citingDracon¬
tium dubium as a synonym. Under contemporary
rules,E.schomburgkiiisasuperfluousname,since
Schott should have used the name E. dubium for
tin 1 transfer. Engler (1911) rectified this situation
bypublishingthecombinationEchidniumdubium
(Kunth)
Engler.
EchidniumisostensiblydistinguishedfromDra¬
contium by having a unilocular ovary w r ith two
ovules (Schott, 1857), as opposed to a hi- or pluriloeular ovary in Dracontium. However, these
characters have been shown to be either spurious
(Zhu, 1995) or not to be good generic characters in
this group (Bogner, 1985; Hay, 1988; Zhu. 1996).
Thetwoplatesoriginatedfromthetypicalmaterial
of D. dubium (Genera aroidearum, pi. 88, Schott,
1858b; Das Pflanzenreich, 4 (23C): p. 38, fig. 14AF, Engler, 1911) clearly demonstrated a plant of at
leasttwolocules.Schott’sstatementofaunilocular
ovary was evidently erroneous, based on his own
illustration. Zhu (1995, 1996) noted that unilocular
ovariesdonotoccuriuDracontium,andthisgenus
neverhasmorethanoneovuleineachloeule;these
stand as generic traits of the genus. Therefore,
Echidnium is accepted as a synonym of Dracon¬
tium, and D. dubium is the accepted name for the
treated
species.
The selection of a neotype is indicated for the
nameDracontiumdubiumKunth,becausenoorig¬
inal material exists (Greuter et al., 1994, Art. 9.7).
The fertile collections of this species studied by
Schott (1860) were either living specimens and
never preserved as herbarium specimens or were
lost before Engler’s time. The drawing made by
Kunth (Engler, 1911) at the Berlin herbarium (B)
waspresumablydestroyedduringwartime,oroth¬
erwise lost (Paul Hiepko, pers. comm.). If Schom¬
burgk made any herbarium collection in the field,
this collection is also untraceable. Plate 88 (Fig. 1)
in the Genera aroidearum (Schott, 1858b) was
clearlybasedonaspecimenofSchomburgk’scol¬
lection of this taxon, and thus affords the most re¬
liablereflectionoftheoriginaldescription(Schott,
Novon 8: 101-103. 1998.
102
Novon
Figure1.NeotypeofDracontiumdubiumKunth:GeneraAraidearum,Plate88(Schott,1858b).
1858a). Therefore this plate (Fig. 1) is here desig¬
natedastheneotypeofDracontiumdubiumKunth.
Becausetheneotypeisanillustrationratherthan
aspecimen,somecharactersimportanttothesystematics of Dracontium, such as the texture of the
inner spathe surface, are not discernable. The To¬
kyo Code (Greuter et al., 1994, Art. 9.7) permits
thedesignationofaninterpretativeepitypeforsuch
a situation. The plant depicted in the neotype il¬
lustrationofDracontiumdubiumhasanapicalap¬
pendage on the spadix (Fig. 1), a character known
to occur only in a few species of Dracontium.
Amongspecimensoriginatingfromareasadjacent
to the typical locality of D. dubium, several from
Venezuela possess this character. One of these.
Hunting 3677B (cultivated at Maracav; originally
collected from Sfquita near San Fernando de Atabapo on Rio Orinoco, Atures Department ol Ama-
Volume 8, Number 1
1998
Zhuetal.
DracontiumdubiumKunth
zonas State), also has an identical spathe with the
neotype of D. dubium. Therefore, Bunting 3677B
(NY) is here designated as the epitype of the name
D.
dubium.
DracontiumchanguangoG.S.Bunting(Bunting,
1986) is characterized by having the inner surface
ofthespathecoveredwithdense,translucentscales
(1—2 mm long) and a spadix that often has apical
appendages.Basedonthepreviousepitypification,
nofeaturesseparateD.dubiumandD.changuan¬
go.Therefore,D.changuangoishereconsidereda
synonymofD.dubium.
ThenomenclatureandsynonymyofDracontium
dubiumKunthareasfollows:
We also thank Paul Hiepko, William D’Arcy, Petra
Malesevich,andHongSongforotherassistance.
Dracontium duhium Kunth, Inil. Sem. Hurt. Berol.
1844:283.1844.EchidniumschornburgkiiSchott,
Oesterr.Hot.Wochenbl.8:62.1857,noni.superfl.
Echidniumdubium(Kunth)Kngler,Pflanzenr.IV.
23C(Heft48):38.1911.TYPE:BritishGuiana.Ml.
CurassawakaoftheCanukuRange,SofNappi,
1843.RichardSchomburgks.n.(holotype,B?lost).
PI.88inSchott,Gen.ArouL,1858(neotype,here
designated).
Venezuela.
Cultivated
at
Maracay,
orig¬
inallycollectedfromSiquitanearSanFernandode
AtabapoonRioOrinoco,AturesDepartmentof
Amazonas
State,
Amazonas,
Atures,
Bunting
3677B
(epitype,
here
designated,
NY).
DracontiumchanguangoG.S.Bunting,Phytologia60:
302,figs.13—14.1986.TYPE:Venezuela.Carabobo:
CanoPasoAncho,ca.6kmSofValencia,nearEl
Paito,Bunting2R56(holotype,MY;isotypes,MO.
NY).
Acknowledgments. We thank Mike Grayum for
discussionsandcriticalsuggestionsonthissubject.
103
Literature
Cited
Bogner,J.1985.Onenewnameandfivenewcombina¬
tionsinAraceae.Aroideana8:73—79.
Bunting,G.S.1986.NewtaxaofVenezuelanAraceae.
Phytologia
60:
293-344.
Engler,A.1911.Araceae—Lasioideae.Pp.1—130inA.
Engler,DasPflanzenreich4(23C).
Greuter,W..F.R.Barrie,H.M.Burdet,W.G.Chaloner,
V.Demoulin,I).E.Hawksworth.P.M.Jorgensen,D.II.
Nicolson,P.C.Silva,P.Trehane&J.McNeill(editors).
1994.InternationalCodeofBotanicalNomenclature
(TheTokyoCode),AdoptedbytheXVthInternational
Botanical
Congress,
Yokohama,
August—September,
1993.RegnumVeg.131.
Hay,A.1988.Cyrtosperma(Araceae)anditsOldWorld
allies.Blumea33:427-469.
Kunth,A.C.1844.Plantaenovae:Hortiregiibotanici
Berolinensis.Ind.Sem.Hort.Berol.1844:283.
Roth.W.E.1922.RichardSchomburgk’stravelsinBritish
Guiana(1840—1844),Vol.I.DailyChronicle,Main
Street,
Georgetown.
-. 1923. Richard Schomburgk’s travels in British
Guiana(1840—1844),Vol.II.DailyChronicleOffice,
Main
Street,
Georgetown.
Schott,H.W.1857.Aroideae.Oesterr.Bot.Wochenbl.8:
61-62.
-. 1858a. Aroideen-Skizzen. Oesterr. Bot. Z. 8:
349-351.
-.1858b.Generaaroidearum.Vindobonae.
-. I860. Prodromus Systematis Aroidearum. Vin¬
dobonae.
Zhu,G.H.1995.SystematicsofDracontiumE.(Araceae).
Ph.D.Dissertation,UniversityofMissouri-St.Eouis,St.
Louis,
U.S.A.
-. 1996. The generic affinity of Echidnium spruceanumSchottanditsplacementinDracontium(.Ara¬
ceae).Novon6:308-309.
Volume 8, Number 1, pp. 1-104 of NOVON was published on 23 April 1998.
Volume
8
Number
NOVON
2
1998
Novelties
in
Neotropical
Sapindaceae
Serjania,
and
II.
Notes
on
Averrhoidium,
Porocystis
PedroAcevedo-Rodriguez
Smithsonian Institution, Dept, of Botany, National Museum of Natural History, NHB-166,
Washington, D.C. 20560, U.S.A.
ABSTRACT. The study of recently collected spec¬
imensofneotropicalSapindaceaerevealstheneed
forvarioustaxonomicchanges.Mataybaspondioides Standley from Mexico is transferred to Aver¬
rhoidium; Paullinia lachnocarpa Bentham ex
Radlkofer(Chimborazoalachnocarpa(Benthamex
Radlkofer) H. Beck) from Ecuador is transferred to
Serjania(therefore,Chimborazoaisreducedtothe
synonymy of Serjania ); and Toulicia acuminata
Radlkofer from Amazonas, Brazil, is transferred to
thecloselyrelatedPorocystis.
RESUMEN. El estudio de especfmenes de Sapindaceasneotropieales,recientementecoleccionados
revelalanecesidaddevarioscambiostaxonomicos.
MataybaspondioidesStandleydeMexicoestransferido al genero Averrhoidium-, Paullinia lachno¬
carpaBenthamexRadlkofer(Chimborazoalachn¬
ocarpa (Bentham ex Radlkofer) H. Beck) del
EcuadorestransferidoalgeneroSerjania(comoresultado el genero Chimborazoa es reducido a sinonimiaconSerjania)-,TouliciaacuminataRadlk¬
ofer proveniente de Amazonas, Brasil, es
transferidoalgenerocercanoPorocystis.
When Paul Standley described Matayba spon¬
dioidesin1927,hecastsomedoubtonwhetherthe
species belonged in this genus. He placed the new
species in Matayba because “It appears to agree
better with that [Matayba] than with any other
American group of the family.” Examination of the
type material, as well as of a recent collection, re¬
veals that this species belongs in Averrhoidium
Baillon, a South American genus with two species.
A.gardnerianumBaillonfromnortheasternBrazil
andA.paraguaienseRadlkoferfromParaguay.AverrhoidiumisdistinguishedfromMataybabyits2ovulatecarpels(vs. 1-ovulate);by itsunilocular (by
abortion), tardily dehiscent capsules, with 1(—2)
seedsperlocule,andchartaceoustocrustoseperi¬
carp(vs.2—3-locular,earlydehiscentcapsuleswith
1 seed per locule, and coriaceous to woody peri¬
carp);byitsseedswithslightlyfleshytesta,without
arillode (vs. seeds with a woody testa with a basal
arillode);andbyitsapetalous(ornearlyso)flowers
(vs.flowerswith5well-developedpetals).Because
thesecharacters(exceptfortheflowers,whichare
unknown) are observable in M. spondioides, this
speciesishereintransferredtoAverrhoidium.
Averrhoidium spondioides (Standley) P. Acevedo-Rodriguez & M. S. Ferrucci, comb. nov.
Basionym:MataybaspondioidesStandley,in
R. S. Ferris, Contr. Dudley Herb. 1: 77. 1927.
TYPE: Mexico. Nayarit: Maria Madre Island
(TresMariasIslands),woodedslopesnearthe
ocean below Balleto Point, 25 Oct. 1925 (fr),
R. S. Ferris 5721 (holotype, CAS; isotype, US).
Additionalspecimenexamined.MEXICO.Jalisco:La
Huerta,CumbresdeCuixmala,km45onroadfromRan¬
choCuixmalatoCumbresI,19°25'N,104°58'W,50m,
25 Aug. 1988 (fr), R. Acevedo R. & J. L. Martinez 956
(US).
Chimborazoa H. Beck was described in 1992 to
accommodatetheodd-lookingPaullinialachnocar¬
paBenthamexRadlkofer,whoseschizocarpicfruits
clearly differ from the septifragal capsules that
characterize Paullinia L. Examination ol lruiting
Novon 8 : 105 - 106 . 1998 .
106
Novon
material available at that time showed P. lachnocarpatobedifferentfromanyoftheremaininggen¬
era of Paullinieae. Thus, the new genus Chimbor¬
azoa was proposed by Beck (1992). However,
examinationofadditionalmaterialofC.lachnocarpa showed that its immature fruits contain a prox¬
imal wing, similar to fruits of Serjania Miller. The
only character used to differentiate this species
from other species of Serjania is precisely the ab¬
senceofmericarpiewingsinmaturefruits.Thedis¬
coveryofvestigialwingsinyoungfruitsofP.lachnocarpadefinitivelylinksthisspecieswithSerjania.
It seems at first that the loss of the mericarpie
winginP.lachnocarpawouldbesufficientgrounds
fordescribingitasadistinctgenus,perhapsonthe
assumptionthatadifferentfruitmorphologyshould
result in a different dispersal mode. Therefore, it
would be a character with biological significance.
Inevaluatingwhetherthisphenomenonmeritsge¬
neric recognition or not, I noted that a few other
species of Serjania (S. cissoides Radlkofer, .S', herterii Ferrueci, and S. macrococca Radlkofer) can
containmericarpswithvestigialwingsorlackthem
altogether.
TherecognitionofChimborazoa(whichisbased
onthelossofthemericarpiewing)requiresthatthe
aforementionedspeciesofSerjaniabetransferred
to Chimborazoa. However, these species do not
seem to be closely related, as they differ greatly in
manymorphologicalfeatures,suggestingthatthe
lossofmericarpiewingshasoccurredalongdiffer¬
entlineagesinSerjania.Therefore,Chimborazoaas
currentlycircumscribedwouldresultinapolyphyletic
taxon.
SincetherecognitionofChimborazoawouldre¬
sult in an artificial taxon, it should be regarded as
asynonymofSerjania,necessitatingthetransferof
P.lachnocarpatoSerjania.
deaealongwithsixothergenera,includingToulicia
Aublet, its closest relative. Porocystis and Toulicia
havesimilarhabitandfloralmorphologiesandcan
only be distinguished by their fruits and embryos.
Whereasbothgenerahaveschizocaqiic,membra¬
noustochartaceousfruits,andembryoswithfleshy
cotyledons, they differ in other respects. Toulicia
hasmericarpsthataresamaroid,containingadis¬
tal, slightly flattened cocci and a proximal wing. In
addition, the embryos of Toulicia have a curved,
external cotyledon and a plicate internal one. Po¬
rocystis, on the other hand, has mericarps with a
large, central, inflated or slightly flattened coccus
thatarewingless.Bothcotyledonsoftheembryos
inPorocystisarestraight.
Porocystishastwospecies,ofwhichthefruitsare
well known. In contrast, Toulicia has 14 species,
but fruits are known for only 7 of them. The place¬
ment in Toulicia of species for which fruits are not
known awaits confirmation, because it is possible
that someofthem might belong in Porocystis. Ex¬
aminationoffruitingmaterialofTouliciaacuminata
Radlkoferrevealsthatthisspeciesisbetterplaced
inPorocystis.Therefore,anewcombinationanda
descriptionofthefruitsarehereinprovided.
Serjania lachnocarpa (Bentham ex Radlkofer) P.
Aeevedo-Rodrfguez, comb. nov. Basionym:
PaullinialachnocarpaBenthamexRadlkofer,
Monogr.Paullinia124.1895—1896.Chimbor¬
azoa lachnocarpa (Bentham ex Radlkofer) H.
Beck, Brittonia 44: 308. 1992. TYPE: Ecua¬
dor. Andes, 1857-1859 (fl, fr). Spruce 6011
(lectotype, designated by Beck (1992: 308), K;
isolectotypes, C, F, G, GH, M, NY—2, P, S—
).2
Additionalspecimenexamined.ECUADOR.Bolivar:
Sicoto,alongroadfromGuarandatoSanPablodeAtenas,
cloudforest,1°50'S,79°05'W,2200-2450m,28Aug.
1987(fr),V.Zah&./.Jaramillo2545(US).
PorocystisRadlkoferbelongstothetribeSapin-
Porocystis acuminata (Radlkofer) P. AeevedoRodrfguez,comb.nov.Basionym:Touliciaac¬
uminata Radlkofer, in Mart., Fl. Bras. 13(3):
505. 1900. SYNTYPES: Brazil. Amazonas:
Manaus, in forest, Apr. 1882 (fl), Schwacke
4001 (GOET); s.d. (fl), Glaziou 13631 (B de¬
stroyed?).
Fruits schizocarpic, 2- or 3-carpellate, broadly
obovateinoutline,chartaceous,reticulate-veined,
minutely tomentose; mericarps slightly inflated,
3.2—3.4 X 2.6—2.8 cm, with dorsal suture com¬
pressed into a narrow wing; endocarp glabrous.
Seed(immature)bean-shapedandtomentose.
Additionalspecimensexamined.BRAZIL.Amazo¬
nas:RioCuieiras,5kmupstream,igapoforest,5Apr.
1974(fl),Campbell,I).G.elal.P21847(US);RioCuieir¬
as,28Apr.1975(fr),CarreiraL.57(INPA).
Acknowledgments. I thank John Pruski (US)
and Mark T. Strong (US) for reviewing the manu¬
script, and R. Laurie Robbins lor valuable com¬
ments.
Literature
Cited
Beck,H.T.1992.Chimborazoa(Sapindaceae),anewge¬
nusfromEcuador.Brittonia44:306-311.
Standley,P.1927.In:R.StinchfieldFerris,Preliminary
reportonthefloraoftheTresMarfasIslands.Contr.
DudleyHerb.I:63-88.
Two
New
Species
of
America,
with
Brachymenium
Brachymenium
a
(Bryaceae)
Key
to
the
in
Central
Species
from
Central
of
America
BruceAllen
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. Brachymenium, dehilinerve differs
from B. speciosum by its dark green color, weak
costa,well-spaced,flat,caducousleaveshavingnu¬
merousquadratealarcells,andaweaklydeveloped
limbidium.Brachymeniumexoticosporumdiffers
from all members of the genus in having massive,
mostlylinear,endosporicprotonematawithmostly
verticalend-walls.Theendosporicprotonemataof
B. exoticosporum appear similar to those found in
Ephemeropsistrentepohlioides.Akeytothe16spe¬
cies of Brachymenium in Central America is given.
Brachymeniumisamostlytropicalorsubtropical
genus of about 70 species (Ochi, 1992). Nearly all
Brachymenium species have hexagonal to rhomboidal upper leaf cells, excurrent costae, and apiculate to hair-pointed leaves. Many of the common
species also have quadrate basal leal cells. The
Brachymeniumgametophyteismostlyindistinct,
and sterile material can be difficult to distinguish
fromBryumorAcidodontium.TheBrachymenium
sporophytehaserecttosuberectcapsuleswithconic-apiculatetoshort-beakedopercula.Itsperistome
isdiplolepidouswithnarrow,denselypapilloseexostometeethandareducedendostome.TheBra¬
chymenium endostome has a high or low basal
membrane with the segments and cilia generally
rudimentarytoabsent.Whenthesegmentsandcil¬
iaarerudimentary,theyaremorphologicallyiden¬
tical and as such the two structures can be iden¬
tifiedonlybytheirpositionrelativetotheexostome
teeth. This endostome condition is often termed
erose. A good example of the loss of segment/cilia
differentiation in Brachymenium occurs in the bi¬
zarre endostome of B. columbicum (De Notaris)
Brotherus (see Shaw, 1984). This endostome has a
highbasalmembranefromwhicharise16morpho¬
logically similar, lanceolate structures. Each lan¬
ceolatestructureispositionedoppositeanexostome
toothandrepresentsacilia/segmentcomplexcon¬
sisting of 4 fused parts: 2 cilia (in the center of the
linearstructure)withViofasegmentoneitherside.
Brachymenium has been divided into five sec¬
tions(seeOchi,1980,1992)basedonfeaturessuch
asplantsize,operculumshape,capsuleshapeand
size,sporesize,limbateconditionoftheleaves,and
peristome structure. The sections, however, have
species with reticulating character combinations
that serve as intermediates between the sections,
making it sometimes difficult to separate them
(Ochi, 1980). Some of the present sectional char¬
acteristics may be of minor phylogenetic impor¬
tance (i.e., plant and capsule size, presence or ab¬
sence of leaf border), and an analysis of the genus
focusing on more reliable features (i.e., exostome
andendostomemorphology,sporesize,basalleal
cell shape) within the context ol an in-depth sistergroup character analysis is needed before a more
naturalsubgenericclassificationcanbeproposed.
There are 16 species of Brachymenium in Cen¬
tral America, 2 ol which are described below as
new to science, in preparation for the next volume
of the Moss Flora of Central America (Allen, 1994).
Brachymenium dehilinerve B. H. Allen, sp. nov.
TYPE:Panama.Chiriquf:FortunaDamregion,
along trail to Cerro Homito (Pate de Macho)
on southern ridge of watershed, 8°45'N,
82°15'W, 1800-1950 m, McPherson 13595D
(holotype,MO;isotype,PMA).Figure1A—H.
AB.specioso(Hookerf.&Wilson)Steerefoliisatrovirentibus,planis,caducis,limbidiodebili,cellulisalaribus
quadratis,
costaque
debili
differt.
Plants large, dark green, in loose tufts, moder¬
ately tomentose below. Stems to 30 mm long, epi¬
dermal cells rectangular, rhizoids reddish brown,
papillose.Leavesequallyfoliate,distantlyspaced,
3-5mmlong,erect-flexuous,attimesweaklytwist¬
ed, erect-spreading when wet, plane, ovate to ob¬
long-lanceolate,long-acuminate,oftenasymmetri¬
cally curved above, margins not or faintly and
indistinctlyborderedby1-2rowsofsomewhatlon¬
ger,narrowercells,entirebelow,serratetodentic¬
ulateabove,theteethsingleoroccasionallydouble,
plane; costa thick at base and tapering above, Vi to
% the leaf length; cells firm-walled throughout,
sometimes porose, upper cells long-rhomboidal.
Novon 8 : 107 - 112 . 1998 .
108
Novon
Figure1.BrachymeniumdebilinerveB.H.Allen.—A.Habit.—B.Capsule.—C.Leafapexandupperleafcells.
D&H.Leaves.E.Basalcellsinalarregion.—ELeafmarginandmedianleafcells.—G.Leafmarginshowing
occasional double toothing. Scales in mm: left = 0.5 (B); middle = 0.5 (D. H); middle = 0.1 (C E FV middle = 0 05
(G);right-2.0(A).
Volume 8, Number 2
1998
Allen
Brachymenium from Central America
60-120 X 16—20 jum, basal cells quadrate. Dioicous. Setae 20-25 mm long, red. Capsules 5-6 mm
long, cylindrical, constricted at neck, erect; annu¬
lus not seen, opercula not seen; peristome badly
eroded,onlyremnantsofexostomeandendostome
present. Calyptrae not seen. Spores 10—16 yam,
spherical,smoothtolightlyroughened.
to lanceolate, 5 mm long. Dioicous. Setae to 65 mm
long, yellowish red to orange. Capsules to 7 mm
long, cupulate, neck to 3 mm long, abruptly con¬
stricted,erect;operculumnotseen;exostometeeth
linear-lanceolate,redbelow,whitishabove,densely
papillose, endostome whitish, papillose, basal
membraneshort,segmentsandciliarudimentaryto
absent. Calyptrae not seen. Spores 40-56 gm, thinwalled,oblong,lightlyroughened.Endosporicprotonemata filling the capsule, linear, multicellular,
mostlyuniseriate,240—320yarnX50—60yam.
Habitat. On tree trunks and branches; 1800—
1950m.
BrachymeniumdebilinerveisalargeBrachymen¬
iumspeciesverysimilartoB.speciosum(Hookerf.
& Wilson) Steere. Both differ from all other Central
American species of Brachymenium by the com¬
bination of their large size, extremely long leaf
cells, and occasionally, doubly toothed leaf mar¬
gins. Brachymenium debilinerve is marked by its
darkgreencolor,weakcosta,well-spaced,flat,ca¬
ducousleaveshavingnumerousquadratealarcells,
and a very weakly developed limbidium. In con¬
trast, B. speciosum is yellowish green, has percurrenttoshortlyexcurrentcostae,andcloselyspaced,
persistentleaveswithrectangularalarcellsthatare
distinctively bordered by 3-4 rows of linear, thickwalled cells. Brachymenium debilinerve is known
from Costa Rica (Guanacaste: Dauphin 1833 (CR,
MO); Puntarenas: Dauphin 1454 (CR, MO)) and
Panama.
Brachymenium exoticosporum B. H. Allen, sp.
nov. TYPE: Panama. Chiriquf and Bocas del
Toro: ridge top N of Cerro Pate Macho, ca. 5
km NE of Boquete above Palo Alto area,
8°48'N, 82°24'W, 1950-2200 m, Hammel,
Grayum,McPherson&Smith14404(holotype,
MO; isotype, PMA). Figure 2A-I.
Species
protonematibus
endosporis
grandibus
linearibus,
setisque
longissimis
congeneribus
a
differt.
Plants medium to large, yellowish green, shiny,
in loose, open tufts, tomentose below. Stems to 30
mmlong,epidermalcellsrectangular,rhizoidsred¬
dishbrown,papillose.Leavesdistantlyspacedand
equallyfoliate,3—5mmlong,flexuous-spreading
whendry,erect-spreadingwhenwet,concave,ob¬
long-elliptictoovate,acuminate,endingindentic¬
ulatepoints,marginsdistinctlyborderedbyseveral
rowsoflinear,thick-walledcells,entirebelow,ser¬
ratetodenticulateabove,teethsingleoroccasion¬
ally double, reflexed at base, plane above; costa
taperingtotheapex,shortlyexcurrentintothehairpoint;cellsfirm-walledthroughout,porose,upper
cells long-rhomboidal, 40-70 X 20—24 gm, basal
cellsrectangular,30—50X16—20gm,Perichaetial
leavesstronglydifferentiated,triangular-lanceolate
109
Habitat. Epiphyte on tree branches in forest on
ridge; 1950-2200 m.
Brachymeniumexoticosporum,knownonlyfrom
the type, is a robust species with distantly spaced,
flexuous leaves that are distinctly bordered and
sharplyserrate;themarginalteethattimesaredou¬
ble. In its distantly spaced leaves it resembles B.
debilinerve but that species has flat leaves, a short
costa(%—%theleaflength)andveryweaktoabsent
leaf limbidia. It is also gametophytically close to B.
speciosum,butthatspeciesdiffersinitslongerleaf
cells (80-120 yam long). The outstanding features
ofB.exoticosporumarefoundintheunusuallylarge
size of its sporophytes and its massive endosporic
protonemata.AlthoughthesporesofB.exoticospo¬
rumarealsolarge,thereareseveralBrachymenium
speciesthathavesporesnearlyasbig(B.consimile
(Mitten) Jaeger, 20—30 yam; B. radiculosum
(Sehwaegrichen)Hampe,22—30yam;B.spirifolium
(C. Muller) Jaeger, 30-40 yam), and one species (B.
standleyi Bartram, 20—80 yam) has larger spores.
There are relatively few spores in the capsules of
B.exoticosporum,andmanyoftheseappearinviable because they have collapsed spore walls. Most
ofthecapsulecontentsconsistoflarge,linear,multicelluarstructuresthatrepresentenodosporically
germinatedprotonemata.Thisisthefirstreportof
massive,endosporicprotonematainBrachymen¬
ium.Theendosporicprotonematahavemostlyver¬
ticalend-walls.Occasionallythesestructureshave
horizontalorobliquewalls,andsometimestheyare
irregularly rounded with cell walls in all planes.
Endosporically germinated protonemata are a
rarefeaturerandomlydistributedamongmosses.
Allen (1987) postulated that these structures may
be ecologically adaptive since they have been
foundtoshortenthetimebetweendiasporerelease
andleafygametophyteformation.Mostendosporic
protonemataaremoreorlessgloboseand/ormultiseriateinstructure.Theendosporicprotonemata
ofB.exoticosporumresemblemoreorlessuniseri¬
ate gemmae and appear similar to the type found
110
Novon
Figure2.BrachymeniumexoticosporumB.H.Allen.—A.Habit.—B&E.Vegetativeleaves.—C&F.Endosporic
protonemata.I).Eeafapexandupperleafeells.—G.Basalcellsinalarregion.—H.Leafmarginandmedianleaf
cells. —I. Perichaetial leaf. Scale in mm: = 0.1 (C. I). F. G, H); = 0.5 (I); = 1.0 (B. E). Habit (A) drawn full-sized.
111
Volume 8, Number 2
1998
Allen
Brachymenium from Central America
inEphemeropsistrentepohlioides(Renner)Sainsbury(Sainsbury,1955).
(occasionallywithtaperedendwalls);
upper
leaf
margins
crenulate
toserrulate;leafmarginsoftennar¬
rowly
reflexed
.
klotzschii
(Schwaegrichen)
Paris
10.Leavesimbricate,hair-pointed,the
hairpointhyalinethroughoutorhy¬
alineabove,reddishbelow,0.5—1
mmlong;leafborderoflinearcells
withlongtaperedend-walls;upper
leafmarginstoothedorsharplyser¬
ratetodenticulate;leafmargins
plane
above.
11
11(10).Leaveswithhair-pointhyalinethroughout,
apical leaf cells linear, hyaline .
.
B.
niveum
Beseherelle
1I.Leaveswithhair-pointhyalineinupperhalf,
reddishbrownbelow,apicalleafcellshex¬
agonal
rhomboidal-hexagonal,
to
concolorous
.
B.
morascium
Beseherelle
12(9).Setaeto65mmlong;capsulesto7
mmlong;capsuleswithunicellular
spores(40-56gm)andlinear,multicelluar,
endosporic
protonemata
(240-320
X
56
gm).
. B. exoticosporum B. II. Allen
12. Setae to 30 mm long; capsules to 5
mm
long;
spores
unicellular
(10—80
gm),
multicellular,
endosporic
pro¬
tonemata
absent.
13
13(12).Plantslarge,to30-40mmlong;leavesflexuousandappressedtostemwhendry,oblong
to
oblong-lanceolate;
upper
leaf
cells
80—120
gun
long
.
....B.speciosum(Hookerf.&Wilson)Steere
13. Plants medium, to 15 mm long; leaves spi¬
rally
contorted
or
crisped
and
contorted
when
dry,
elongate-oblong,
obovate-oblong
to
spathulate
(rarely
oblong-lanceolate);
upper
leaf
cells
30-60
gun
long.
14
14(13).Leavescrispedandcontortedwhen
dry,
often
orbicular,
obtuse
to
shortly
and broadly acuminate .
. B. wrightii (Sullivan) Brotherus
14. Leaves spirally twisted when dry,
variously
lanceolate,
oblong,
obo¬
vate-oblong,spathulate,acuteto
acuminate.
15
15(14).Leafmarginsnearlyentiretofinelyserrulate
attheapex;operculumhigh-conicand
obliquelybeaked;exostometeethlinear.
. B. spirifolium (C. Muller) Jaeger
15.Leafmarginssharplyserrulateordenticulate
inupperVyoperculumconictoconic-apiculate;exostometeethlinear-lanceolateto
lanceolate
.
16
16(15).
Exostome
teeth
broadly
lanceolate,
united
at
base;
endostome
segments
andciliarudimentaryorwithendostomalmaterialadheringinpatches
totheupperpartsoftheexostome
teeth....B.consimile(Mitten)Jaeger
16. Exostome teeth linear-lanceolate,
freeatbase;endostomesegments
absent,rudimentary,orwelldevel¬
17
oped
.
17(16).
Endostome
segments
rudimentary
or
absent;
KeytheSpeciesofBrachymeniuminCentralAmerica
1. Plants minute; leaves less than 1.0 mm long
.
2
1. Plants small to large; leaves greater than 1.2
mm
long
.
3
2(1). Leaf cells lax and thin-walled
throughout,uppercellselongate,
alarcellsshortrectangular;leaves
borderedbylong,narrowcells...
.
B.
acuminatum
Harvey
2. Leaf cells firm and thick-walled
throughout,uppercellsrhombic,
alar
cells
quadrate;
leaves
bordered
by short-rectangular cells.
..B.exile(Dozy&Molkenlxter)lioch&
liicoste
3(1).Leavescaducous;costafromVi—Vztheleaf
length. B. debilinerve B. H. Allen
3.Leavesfirmlyattachedtostem;costapercurrent
to
excurrent.
4
4(3).Leavesnotorindistinctlybordered
...
5
4. Leaves distinctly bordered. 9
5(4).Leaveshair-pointed,thehairpointentirely
orinparthyaline,oftenVi—%theleaflength
.
6
5. Leaves cuspidate, the cuspid reddish, less
than ‘4 the leaf length . 8
6(5). Leaves with hair-point hyaline
throughout,apicalleafcellslinear,
hyaline. B. niveum Beseherelle
6. Leaves with hair-point hyaline in
upperhalf,reddishbrownbelow,
apicalleafcellshexagonaltorhomboidal-hexagonal,concolorous....7
7(6).Capsulesnarrowlycylindrical,3—5gmlong;
upper
leaf
margins
toothed
to
sharply
serrate
or denticulate. B. morascium Beseherelle
7. Capsules obovate-oblong, 2.5^t mm long,
upperleafmarginsentiretoweaklyserrulate
. B. systylium (C. Muller) Jaeger
8(5).Leavesovatetooblong-lanceolate,
1.4-2.3mmlong;basalleafcells
quadratetosubquadrate;plants
without
axillary
bulbils;
capsules
cy¬
lindrical,
endostome
cilia/segments
rudimentary
or
absent.
. B. mexicanum Montagne
8. Leaves ovate, lanceolate, or elon¬
gate-triangular,
2—3
mm
long;
basal
leafcellsshort-rectangulartosub¬
quadrate;plantsoftenwithaxillary
bulbils;
capsules
globose,
endosto¬
me
cilia/segments
well
developed.
opposite the exostome teeth .
B.
columbicum
(De
Notaris)
Brotherus
9(4).
Basal
leaf
cells
quadrate.
10
9.
Basal
leaf
cells
rectangular.
12
10(9).Leavestwistedaroundthestem,
shortlyandstoutlyawned,awnred¬
dishthroughout,to0.2mmlong;leaf
borderoftenofnarrowrectangular
cellswith±right-angledend-walls
112
Novon
sporesirregularinshape,spherical,ovoid,or
oblong, 20-80 /u,m. B. standleyi Bartram
17.Endostomesegmentswelldeveloped,linear
with
narrow
perforation
gaps;
spores
spheri¬
cal,
22—30
/Ain
.
. B. radiculosum (Schwaegrichen) Hampe
-. 1994. Moss Flora of Central America. Part 1,
Sphagnaceae-Calymperaceae.
Monogr.
Syst.
Bot.
Mis¬
souriBot.Card.49:1-242.
Ochi,H.1980.ArevisionoftheneotropicalBryoideae,
Musci(FirstPart).J.Fac.Fduc.TottoriUniv.,Nat.Sci.
29:
49-154.
-. 1992. A revised infrageneric classification of the
genusBryumandrelatedgenera(Bryaeeae,Musci).
Bryobrothera
231-244.
1:
Sainsbury,G.0.k.1955.AHandbookoftheNewZea¬
landMosses.Bull.Boy.Soc.NewZealand5:1-490.
Shaw,J.1984.Areinterpretationofperistomestructure
in
Pseudoditrichum
mirabile
Steere
&
Iwats.
(Pseudoditrichaceae).
Bryologist
87:
314—318.
Acknowledgment. This work was supported by
National Science Foundation Grant DEB-9400996
(totheMissouriBotanicalGarden).
Literature
Cited
Allen,B.H.1987.Observationsontheprotonemataof
Drummondia
prorepens
(Musci:
Orthotrichaceae).
Evansia4:33-37.
Five
New
a
Key
Species
of
to
Species
the
Macromitrium
of
(Musci:
Macromitrium
Orthotrichaceae),
in
Central
with
America
BruceAllen
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT.Macromitriumcrosbyorumsp.nov.,M.
echinatum sp. nov., M. frustratum sp. nov., M. picobonitum sp. nov., and M. sejunctum sp. nov. are
described based on Central American material. A
key to the species of Macromitrium in Central
America is given. The 30 Central American species
of Macromitrium are arranged into 7 informal
groups.
Macromitriumisasizablegenus,about350-400
species, of medium- to large-sized pleurocarpous
mosses. The last treatment of the genus that in¬
cluded Central America was Grout (1946), which
attributed 24 species to the region. The genus is
often found on upper tree branches in the forest
canopy,butitalsocommonlyoccursontreetrunks
and can be found on rocks and soil in open, drier
habitats.MostMacromitriumspecieshaveelimbate
leaves with short upper leaf cells and elongate to
linear, tuberculate basal leaf cells, mitrate calyptrae, and short, truncate exostome teeth that are
fusedformostoftheirlengths.However,thegenus
ismorphologicallycomplex,andthereareMacrom¬
itriumspecieswithvariouscombinationsoflimbate
leaves, long upper leaf cells, short basal leaf cells,
non-tuberculate basal cells, cucullate calyptrae,
andlong,narrowlytriangular,non-fusedexostome
teeth. As a result, the genus is difficult to charac¬
terize or cleanly separate from a number of segre¬
gate genera (e.g., Groutiella , Macrocoma, Cardotiella) recognized for groups of species with short
basalleafcellsincombinationwithsomeotherdis¬
tinctive
feature.
Vitt (1994) recognized three distinctive groups
for the 10 Macromitrium species in Mexico. The 30
CentralAmericanMacromitriumspeciesaremore
diversifiedandvariablethanthoseofMexico.They
include 5 species new to science, which are de¬
scribedbelowinpreparationforthenextvolumeof
the Moss Flora of Central America (Allen, 1994).
Macromitrium crosbyorum B. H. Allen & Vitt,
sp. nov. TYPE: Costa Rica. San Jose: along
Inter American Highway, ca. 10 km NW of
summit at La Ascension, 9°37'N, 83°48'W,
Crosby&Crosby6089(holotype,MO;isotypes,
ALTA, CR, NY, US). Ligure 1.
SpecieshaecaM.subcirrosointeraliafoliorumoellulis
isodiametris,
crasse
unipapillosis
costisque
percurrentibus
differt.
Plantslarge,greenishredtoyellowishred.Stems
creeping to 7 cm, branches 2—3 cm long, reddish
tomentosebelow.Leaveskeeled,erectbelow,flexuoustospirallycontortedandundulateabovedry,
erect-patentwet,(3—)4—6mmlong,1mmwide,lan¬
ceolate, acuminate; margins undulate, serrate
above, frequently serrulate to near the base, re¬
curved below, erect to plane above, swollen basal
teethatleafinsertionabsent;costaepercurrent;up¬
per interior cells 8—20 gm, rounded and collenchymatous,isodiametictorhombic,stoutlyunipapillosetomammillose,uppermarginalcellsnarrow
and elongate forming a ± distinct border, basal
cells long rectangular, incrassate and porose,
denselytuberculate,26—44gunlong.Dioicous.Se¬
tae7—10mmlong,smooth.Capsules1.5—2.0mm
long, ovoid to cylindrical, plicate. Annulus non-revoluble,withfragmentsadheringtocapsulemouth.
Exostometeethtruncate,320-424gmhigh,yellow,
densely papillose-striate, united and forming a
membrane, ± reflexed at tips, splitting into eight
pairs of teeth with age; endostome hyaline, lightly
papillose, basal membrane 80-90 gun high, seg¬
ments 60—80 gm high. Opercula rostrate, 1-1.5
mmlong.Sporesanisosporous,14—20gun,smooth
to lightly papillose and 30^8(—54) gun, densely
papillose. Calyptrae mitrate, deeply laciniate, na¬
ked,5mmlong.
Habitat. On tree trunks, logs, and rocks; 3130—
3333m.
Macromitrium crosbyorum is a large moss with
undulatingleavesandisodiametric,collenchymatous,mammillosetostoutlyunipapilloseupperleaf
cells. It has long, narrow marginal leaf cells that
formavariablydistinctborder.Macromitriumsubcirrosum C. Midler differs from it in having elon¬
gated upper leaf cells, an excurrent costa, and
elimbateleaves.MacromitriumscopariumMittenis
similar to M. crosbyorum in having limbate leaves,
isodiametricupperleafcells,andtuberculatebasal
leaf cells but differs in having smaller, narrower.
Novon 8: 113-123. 1998.
Peterson, P M and Planchuelo, A M. 1998. "Bromus catharticus in South
America (Poaceae: Bromeae)." Novon a journal of botanical nomenclature from
the Missouri Botanical Garden 8, 1–60. https://doi.org/10.2307/3391893.
View This Item Online: https://www.biodiversitylibrary.org/item/14668
DOI: https://doi.org/10.2307/3391893
Permalink: https://www.biodiversitylibrary.org/partpdf/24852
Holding Institution
Missouri Botanical Garden, Peter H. Raven Library
Sponsored by
Missouri Botanical Garden
Copyright & Reuse
Copyright Status: In copyright. Digitized with the permission of the rights holder.
License: http://creativecommons.org/licenses/by-nc-sa/3.0/
Rights: https://biodiversitylibrary.org/permissions
This document was created from content at the Biodiversity Heritage Library, the world's
largest open access digital library for biodiversity literature and archives. Visit BHL at
https://www.biodiversitylibrary.org.
This file was generated 16 April 2022 at 20:13 UTC
Related papers
Alexandru I. Philippide, Introducere în istoria limbei și literaturei române [1888]
Iași, Editura Librăriei Frații Șaraga, 1888
Teaching the City: Exploring Pedagogies of Urban Becoming
Teaching and Learning Anthropology, 2024
RAICES CORPORALES DEL DIBUJO
Raices Corporales del Dibujo, 2017
Terapia asistida con animales. Efectos positivos en la salud humana
Journal of Agriculture and Animal Sciences, 2013
Application des commandes non linéaires pour la régulation en vitesse ou en position de la machine synchrone autopilotée
Revue de Physique Appliquée, 1990
Earthing the <i>Anthropos</i>? From ‘socializing the Anthropocene’ to geologizing the social
European Journal of Social Theory, 2016
Refleksje Józefa Andrzeja Gierowskiego o polskiej kulturze i nauce XVIII wieku
Historyka. Studia Metodologiczne, 2023
The Impossibility of Cinema: Intertwinings of “Pre” & “Post” Cinema in Contemporary Art
&&& Journal, 2020
A gramática como lugar de materialização de dizeres sobre
a língua: o caso do substantivo na 1.ª e na 37.ª edição da
Moderna Gramática Portuguesa de Evanildo Bechara
Revista Confluência, 2017
The Sensitive Visualization and Generalized Fractional Solitons’ Construction for Regularized Long-Wave Governing Model
Fractal and Fractional
The Effect of Greenhouse and Biopore on Community Development of Economy and Knowledge of Citeureup Village During the Pandemic
Journal of Innovation and Community Engagement
Photoluminescence Characteristics of Cu+ and Mn2+ in Activated Halosulphate Phosphor
Journal of emerging technologies and innovative research, 2021
A rare etiology of diffuse pulmonary hemorrhage
Advances in respiratory medicine, 2021
The Support Bot Architecture
Theoretical & Applied Science, 2020
Noninvasive ventilation in acute severe asthma
Current Opinion in Pulmonary Medicine, 2014
Call Center and its Significance for Sexual and Reproductive Health
Epidemiology, 2017
Are Descriptions Really Descriptive? An Experimental Study on Misdescription and Reference
Review of Philosophy and Psychology, 2018