An evaluation of the Mesoamerican species of Meriania (Melastomataceae: Merianieae)

12 Pages·1993·5.7 MB·English

by  Frank Almeda

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CALIFORNIA ACADEMY OF SCIENCES
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Vol. 48, No. 7, pp. 141-152, 2 figs December 21, 1993
DEC
2 7 1993
AN EVALUATION OF THE MESOAMERICAM SPECIES OF
MERIANIA (MELASTOMATACEAE: MERIANIEAE)
By
Frank Almeda
DepartmentofBotany, CaliforniaAcademyofSciences, GoldenGatePark,
SanFrancisco, California94118
Abstract: Meriania, one of 17 genera in the neotropical tribe Merianieae, ranges widely from southern
Mexico,CentralAmerica,andtheGreaterAntillessouthtothetropicalAndes,theGuayanaHighlandregion,
and southeastern Brazil. In this study, Meriania isreported from Mexicoforthe firsttime; a newspecies,
M.odorataisdescribedfromCostaRicaand Panama;andtwospeciesofCentronia, C.grandiflora,and C.
phlomoides, are transferred to Meriania. This summary ofthe five Mesoamerican species includes a key,
descriptions, geographical and phenological notes, diagnostic illustrations, and citation of representative
specimens.
Received24May 1993.Accepted23June1993.
Introduction wereknownfromCentralAmericabasedonfew
collections (Gleason 1958; Standley and Wil-
The genus Meriania is a unifying basal group liams 1963).
amongthe 17generacomprisingtheMerianieae, Inthisstudy,therangeofMerianiaisextended
aneotropical tribe characterizedby prevailingly toMexico,anewspeciesisdescribedfromCosta
terminal inflorescences, capsular fruits, and ob- RicaandPanama,andthetwoCentralAmerican
long-pyramidate, angulate or winged seeds with species previously included in Centronia are
a smooth or poorly defined surface patterning. transferredtoMeriania.Tofacilitaterecognition
With over 50 described species, Meriania is the andcomparison,aregionalrevisionispresented
largest genus in the tribe and one oftwo wide- withakey, descriptions,diagnosticillustrations,
spread genera ranging from southern Mexico, andcitationofrepresentativespecimens.Allspe-
CentralAmerica,andtheGreaterAntilles,south ciestreatedhererepresentoutliersofagenuswith
throughthetropicalAndesofSouthAmericaand majordiversityin SouthAmerica. The five Me-
east to the Guayana highlands and southeastern soamericanspeciescanbesortedintothreegroups
Brazil. Preparation ofa treatment ofthe Melas- based on androecial morphology and details of
tomataceae forFloraMesoamericanahasneces- the indument. Each group appears to have its
sitated an evaluation ofMeriania and the five closest affinities with a different assemblage of
other genera ofMerianieae reported for the re- SouthAmericantaxa. Abetterunderstandingof
gion (Almeda 1981, 1989, 1990; Eves 1936; species relationships, therefore, must await a
Gleason 1958;StandleyandWilliams 1963).Pri- comprehensiverevision ofMerianiaandits sat-
or to this study, only two species ofMeriania ellite genera.
[141]
142 PROCEEDINGSOFTHECALIFORNIAACADEMYOFSCIENCES,Vol.48, No. 7
Systematic Treatment fixinggenericlimitsbecausedelimitationofMer-
iania and its satellite genera relies heavily on
Meriania Swartz, Fl. Ind. Occ. 2:823. 1798. A
these characters. reconsideration of generic
(Norn, cons.)
limits in the tribe is beyond the scope of this
Type.—M. leucantha(Swartz)Swartz,typecons.[=Rhexia study. Hopefully, new sources of comparative
leucanthaSwartz]. data such as pollen morphology and chromo-
Treesorshrubswithglabroustovariouslypu- some numbers will provide information needed
bescent distal branchlets. Leaves opposite, de- to reevaluate relationships among these genera.
cussate, and often long-petiolate, those ofa pair Intheabsenceofanoptimalgenericclassification
equal to somewhat unequal, glabrous to vari- ofthetribe,Adelobotrys,Axinaea, Graffenrieda,
ously pubescent. Rowers typically borne in ter- Meriania, and the more distantly related Tess-
minal multiflowered panicles but occasionally mannianthus will be recognized in the melas-
solitary, ternate, or verticillate. Hypanthium tome treatment for Flora Mesoamericana.
mostlyfree from the ovary, campanulate, hemi- The two Mesoamerican species of Centronia
spheric, or urceolate, terete or costate in fruit. have androecial morphology and dorsally in-
Calyxtube usuallywell-developed, truncateand clined anther pores like Meriania and are here
flangelike,calyxlobesandexternalteethobsolete transferred to that genus. They are clearly dis-
or the calyx deeply lobed with well-developed cordant in Centronia, because they do not have
exterior teeth, or calyx sometimes fused in bud arcuateessentiallyisomorphicantherswithven-
andrupturingirregularlyatanthesis. Petals 5 (in trally inclined pores and a connective modified
ourspecies),freeandtypicallyglabrous,oblique- into a solitary acute dorso-basal spur.
ly to broadly rounded or truncate at the apex. Among Mesoamerican genera of Merianieae
Stamens 10 (in our species); nearly isomorphic theonlyspeciesthatcouldbeconfusedwithMer-
todimorphic,glabrous,stronglygeniculateatthe iania is Adelobotrys panamensis Almeda. This
filament insertion; filaments usually declined in species is similarto some speciesofMerianiain
onedirectiontransverselyacrosstheflower;the- being shrubby or arborescent, lacking malpigh-
caesubulatewithadorsallyorventrallyinclined iaceoushairs,andinhavinganantherconnective
modifieddorso-basallyintoanerectspurandan
apical pore; connective usually thickened, pro-
longed,orvariouslymodifieddorsallyintoador- acuteappendagedirectedmoreorlessparallelto
so-basal spur and an ascending blunt or pro- the thecae (Almeda 1981). Adelobotrys pana-
longed dorsal appendage. Ovary superior(3-)5- mensis is readily distinguished from all species
6-celled, glabrous, the apex prolonged and ofMeriania,however,byitscompressed,2-edged
uppermost branchlets and umbelliform pseu-
toothed, truncate, or umbilicate. Style straight,
somewhat sigmoid or declinate with a truncate dolateral inflorescences borne on short stubby
or punctiform stigma. Fruit a many-seeded lo- branchlets.
culicidal capsule. Seeds narrowly oblong-pyr-
amidate to cuneiform, angulate, truncate to Key to the Mesoamerican
somewhat uncinate at the wider distal end or
Species ofMeriania
with tails at both ends.
Among the 17 genera of Merianieae, Adelo- 1. Leafbladesvariouslypubescentontheab-
botrys,Axinaea, Centronia,andGraffenriedaare axial (lower) surface.
close relatives ofMeriania. There is a particu- 2. Abaxial foliar surface covered with
larlycloserelationshipwiththe Centronia/Graf- shaggy-plumulose hairs and a sparser
fenrieda alliance on one hand and a cluster of stellate-hirsute indument between the
Adelobotrysspeciesontheother.Ithasevenbeen elevated veins; pedicels and hypanthia
suggested that the circumscription ofMeriania copiously pubescent; calyx fused into a
couldlogicallybeexpandedtoincludethesefour budcapthatrupturesirregularlyandfalls
closely related genera (Wurdack 1973). away after anthesis; petals white or
As is true of other tribes in the Melastoma- greenish-white; stamensnearly isomor-
taceae, the Merianieae exhibit an impressive phic.
rangeofform incalyxstructureandanthermor- 3. Inflorescencea few-floweredpanicle
phology. Itisthespectaculardiversityandmod- with ultimate units consisting of
ificationsin thesefeaturesthat poseproblemsin simple cymes or paired or solitary
ALMEDA: MESOAMERICANSPECIESOFMERIAN1A 143
flowers; floral buds 15-25 x 10-13 5.5-14 cm, ellipticto elliptic-ovate, (5—)7—9-pli-
mm; petals 2-2.7 x 1.5-3 cm; an- nerved with a well-defined elevated network of
mm
therthecae 10-11 long; fruiting secondary and tertiary veins, base obtuse to
mm
hypanthia 10-14 x 15-20 rounded or varying to obscurely cordate, apex
M. grandijlora acuminatetoattenuate,themarginentiretoden-
3. Inflorescence a multiflowered verti- ticulate, the adaxial surface glabrous and retic-
cillate panicle with ultimate units ulate-bullate, the abaxial surface covered with
consisting of fascicled or umbelli- basallyplumulose hairson the primaryandsec-
formclusters; floralbuds 9-13 x 4- ondary elevated veins and a stellate-hirsute in-
7 mm; petals 1.1-1m.6mx 0.8-1.3 cm; dument between the veins. Inflorescence a ter-
anther thecae 6.5 long; fruiting minal few-flowered, openly branched,
mm
hypanthia 6-7 x 5-7 pedunculatepanicle 10-21 cmlong,theultimate
M. phlomoides units consisting ofsimple cymes or reduced to
2. Abaxial foliar surface uniformly cov- paired or solitary flowers; bracteoles 15-30 x
ered with a pulverulent or floccose in- 2.5 mm, linear-lanceolate, early deciduous and
dument; pedicels and hypanthia gla- typicallyseenonlyonyounginflorescences.Ped-
brous;calyxtruncateandflangelikewith icels 9-15 mm long, lengthening to as much as
persistent lobes that are ill-defined de- 27 mm in fruit. Hypanthium (atanthesis)hemi-
pressed-ovate undulations; petals ma- spheric, 8-12 mmlongtothetorus. Flowerbuds
genta; stamens dimorphic rounded-ellipsoid, 15-25 x 10-13 mm, shortly
M. macrophylla (2-3 mm) blunt-rostrate. Calyx fusedinbudbut
1. Leafbladesglabrousontheabaxialsurface. rupturingintoirregularrounded-deltoidlobesof
4. Leaves entire; calyx on fruiting hypan- varying size that fall away after anthesis. Petals
thia truncate and flangelike, the lobes 2-2.7cmlong, 1.5-3cmwide,whiteorgreenish-
obsolete orconsistingofill-defined un- white, translucent when fresh, sometimes with
dulations; petals 1.7-2.5 x 1-1.7 cm; darkenedconspicuousvenationwhendry,erect,
mm
fruiting hypanthia 6-7 x 7 obovate, ± concave and connivent, rounded to
M. odorata shallowlyemarginateapically,themarginentire.
4. Leavescoarselydentatealongtheapical Stamens nearly isomorphic, very similar in size
halfortwo-thirdsoftheblade; calyxon butdifferingslightlyinconnectivemodification.
fruiting hypanthia consisting ofbluntly Antepetalousstamens: filaments 10-11 mmlong,
roundedortruncatelobes2-4mmlong; anther thecae 10-11 mm long, white, subulate,
petals 2.6-3.9 x 1.6-3.7mcmm; fruiting horizontalandessentiallystraightwithadorsally
hypanthia 9-11 x 12-13 inclined pore; connective thickened, ± flat ven-
M. panamensis trally but somewhat ridged dorsally along the 2
mm
segmentprolongedbelowthethecae,dilated
Meriania grandiflora (Standi.) Almeda, comb. dorso-basally into a divergent longitudinally
nov. ridgedandadaxiallyfurrowedappendage(4x2
(Figs. 1C, D, E) mm) anda smalleradjacent spurelevated 1-1.5
mm
CentroniagrandifloraStandi.,FieldMus.Nat.Hist.,Bot.Ser. mm. Antesepalousstamens:mfimlaments 10-11
22:95. 1940.Type.-Panama.Chiriqui:BajoChorro,2,100 long, anther thecae 10-11 long, otherwise
m, 2 Mar 1938 (fl), Davidson 360 (Holotype: F!; isotypes: like antepetalous stamens in color, posture, and
A!, MO!,US!). pore inclination; connective flattened ventrally
Tree 3-15 m tall. Older cauline internodes ± but vaguely ridged dorsally along the 0.5 mm
terete andglabrate; thebluntly quadrate, upper- segmentprolongedbelowthethecae,dilateddor-
most branchlets, inflorescence axes, floral buds, so-basally into an erect compressed, longitudi-
and hypanthia moderately to densely covered nallyridgedappendage(4x3mm)andasmaller
with brown, basally enlarged hairs (0.5-2 mm adjacent tooth elevated 1 mm. Ovary (at anthe-
long)thatareshaggy-plumulosebasallybutspar- sis) ovoid but truncate apically, glabrous, 5-6(-
ingly barbellate to glabrous toward the apex. 7)-celled, becoming rounded-lobulate to round-
Leaves ofa pair equal to somewhat unequal in ed-truncate apically at maturity, superiororad-
size; petioles 2-7.5 cm long and 2-3 mm wide; nateto thehypanthiumbasallyforabout 3 mm.
mm
blades membranaceous when dry, 11.2-21.3 x Style 21-25 long, glabrous, erect or slightly
PROCEEDINGSOFTHECALIFORNIAACADEMYOFSCIENCES,Vol.48,No. 7
144
stFaimgeunre(lef1t.)Raenpdreasnetnetsaetpiavleousstasmteansm,ense(erdisgh,t)a,ndx hcaai.rs3;ofB,Messeeodasm,erxic1a5n. MMe.rigarnainad.ijlMoerrai:anCi,aa^nmtaecpre^otpahlyolu.lsa:stA^a,meannt^e(lpeeftt)alaonuds
antesepalous stamen (ngJt), x 1 1/2; D, seeds, x ca. 13; E, shaggy-plumulosehair (left)-of Wtfdol§j*£«
hsuyrpfaancet,hmx,s1h5.agMg.yhpaainra(mmeindsdilse:)Fo,faabnatxe.paeltaplroiumsasrtyaamnedns(leecfto)ndaanrdyafnotlieasrepvaeilnosu,sasntdamsetenll(artieg-hth)i,rsuxtecah.ai2r; G, seeds, ca. v. m.
ALMEDA:MESOAMERICANSPECIESOFMERIANIA 145
declinate and gently curved apically below the hair covering between the veins on abaxial leaf
truncate stigma. Fruiting hypanthium campan- surfacesisstellate-hirsute.IfollowGleason(1940:
ulate, terete or irregularly rugose orbullate, 10- 339) in using this term for straight or curved
14 x 15-20 mm. Seeds brown to beige, cunei- essentiallysmooth, simplehairswithabasethat
form or oblong-pyramidate and angulate, 0.5-1 is more or less stellate with numerous radiating
mm
long. arms. (Figs. IE, U). For scanning electron mi-
Distribution and Phenology.—Endemic to crographs of comparable hairs see Wurdack
Panama where it is local in cloud forests at (1986:53).
(880)1,350-2,500 m. Floweringcollectionshave The stamens ofM. grandiflora and M. phlo-
been made from January through August; fruit- moidesare also noteworthy in havinga pseudo-
ingspecimens havebeen collectedin all months inversion ofthe connective that was first noted
except March, April, June, and December. for other species ofMeriania by Triana (1871:
RepresentativeSpecimensExamined.—PANAMA. Bocas 165). In each stamen the two anther cells are
delToro: Robalotrail,N slopesofCerroHorqueta, 5-7 Aug completely separated by the thickened connec-
1947 (fl, fr), Allen 4992 (MO, NY); E slope ofLa Zorra to tive. The anthercells are not only separatedbut
divide on Chiriqui trail, 20 Apr 1968 (fl), Kirkbride& Duke stronglypositionedupwardtowardthedorsalap-
833(NY).Chiriqui:vicinityofCerroPunta,24May 1946(fl), pendage-bearing side (Figs. 1C, 1H). This in-
Ak1l5ml3e7nN(3WD4U9oK2fE(C,eBrMMr,Oo)B;PRut,nratiGal,,toN2YC8e)rM;rao1y-P2a1tk9e7mM2aNc(fhl)oo,f,ALl8a°ms4e9dN'uaN,b&e82s°Wa2i4nl'bdWu,5r vmearkteesdtphoestduorrsealofsitdheeapapnethaerrascetlhlsousguhpeirtfiiscivaelnl-y
13Mar 1988(fl),Almedaetal.6140(CAS,MO,PMA,TEX); tral.
Monte Azul, 1.4 mi N ofEntre Rios on E slopes ofCerro Gleason (1958) erroneously applied the name
Punta,22NovN19W79(fr),Antonio2702(CAS);vicinityofUs Centronia tomentosa Cogn. to this species and
NCurboaets,222.3797mi(C, F, MoOf,RiNoY)C;hirBiaqjuoiCVhioerjroo,,2B7oqFueebte19di7s3tri(fclt),, relegated C. grandiflora(= M. grandiflora)to its
25Jan 1938(fr),Davidson212(A,F,MO);GuadalupeArriba, synonymy. Wurdack (1976) subsequently trans-
aboveCerroPunta,8°52'N,82°33'W,23Jul1985(fl),deNevers ferred C. tomentosa to Meriania and correctly
& Charnley6055(CAS); E ofGuadalupe alongRio Chiriqui noted that these two species are not conspecific
Viejo,2miNEofCerroPunta, 13Jan 1971 (fl), Wilburetal. (Wurdack 1980). Among South American spe-
13093(CAS, DUKE);slopesaboveRioCalderabeyondBajo
Mono, 17Jan 1970(fl), Wilburetal. 11082(DUKE, F, GH, cies of Meriania, M. tomentosa resembles M.
MO);vicinityofBajoChorro,20-22Jul1940(fl,fr), Woodson grandiflorainhavinganirregularlyrupturingde-
& Schery 6574 (NY). Code: Cerro Pilon, spring 1968 (fr), hiscent calyx, and anther thecae that show the
Lallathin9(NY). pseudo-inversion described above. It consis-
Merianiagrandifloraanditscloserelative,M. tently differs from M. grandiflora in having or-
phlomoides, differ from other Mesoamerican ange-redpetals,yellowantherthecae, largerseeds
congeners in androecial morphology and com- (1.5 mm) with an elongate lateral raphe, and an
plexity ofthe indument. The upper branchlets, appressed-setose hypanthial indument of fine
inflorescences,floralbuds,andhypanthiaofthese hairs with an expanded roughened base and an
species are covered with hairs that are here de- attenuate cauducous apex.
scribed as shaggy-plumulose. These hairs have
an enlarged roughened or shaggy base (Figs. 1E, Meriania macrophylla (Benth.) Triana, Trans.
U) and a smooth tapering shaft. The elevated Linn. Soc. London 28:66. 1871.
primary and secondary veins on abaxial foliar
(Figs. 1A,B)
surfaces ofthese species are also covered with
similarhairsbutthebaseofeachisnotasprom- DaGvyUaATmaEcMrAopLhAyl.laQuBeeznatlht.e,naPnI.goH:aRrtawn.ch7o5.Pal18o4H1.uecToY,PHEa.r--
inently expanded and the roughening often ex- twegs.n. (Holotype: K!).
tendsforagreaterdistanceupthehairshaft. The ConostegiaexcelsaPittier,Jour.Wash.Acad.Sci. 14:450. 1924.
phlomoides:H,antepetalousstamen(left)andantesepalousstamen(right), x ca.11;I,seeds, xca.11;J,shaggybasallyenlarged
hair(left)ofbranchlets,inflorescences,andhypanthia,shaggyhair(middle)ofabaxialprimaryandsecondaryfoliarveins,and
stellate-hirsutehair(right)ofabaxialfoliarsurface, x ca. 2 1/4.(AfromBreedlove47700;BfromBreedlove&Almeda47727;
CfromAlmeda& Wilbur1537; DfromAntonio2702;EfromAlmedaetal. 6140;FfromAntonio1040;GfromLao399; H
fromAlmedaetal. 2728; IfromAlmeda&Daniel7076;JfromAlmeda&Nakai4674.)
146 PROCEEDINGSOFTHECALIFORNIAACADEMYOFSCIENCES,Vol. 48,No. 7
TYPE.—PANAMA.Chiriqui:southernslopeofCerroHor- spheric, terete to bluntly costate when dry, 6-8
queta,Pittier3196(Holotype: US!;isotype:F!). mm long to the torus and 6-7 mm wide. Seeds
brown, vernicose, straight to somewhat curved,
Tree 6-21 m tall with terete to bluntly quad- cuneiform and angulate, 0.5-1.5 mm long.
rate cauline internodes. Distal branchlets, pe- Distribution and Phenology.—Local and
dunclesand pedicels ofthe inflorescence, young uncommon from Chiapas, Mexico, and western
hypanthia,andabaxialfoliarsurfacescaducously Guatemala, disjunct to central Costa Rica and
pulverulentorfloccose.Leavesofapairtypically western Panama, in montane forests at 1,400-
somewhatunequalinsize; petioles(2.3-)4-9 cm 2,600 m. Flowering from September through
mm
longand3.5-6 wide;bladescoriaceouswhen JanuaryandJuly; fruitingfromJanuarythrough
dry, 9.8-34 x (5-)10-28 cm, elliptic to elliptic- April.
ovate, sometimes varying to subrotund, 5(-7)-
nerved with secondaries also prominent and RepresentativeSamplesExamined.—MEXICO.Chiapas:
elevatedabaxially, base broadlyroundedtocor- SWslopeofVolcanTacanaaboveTalquian, 13Dec 1974(fl),
date and typically subpeltate, apex obtuse to Breedlove42550(CAS,DS);aboveTalquianonslopesofVol-
rounded and commonly mucronate, the margin coafnVoTlaccaannaT,ac1a8naN,o2v31N9o8v8(1f9l)8,0Br(fel)e,dBlroeveed7lo1v5e3&5(AClAmSe)d;aS4E7s7i0d0e
entire, glabrous adaxially at maturity. Inflores- (CAS, TEX); SEsideofVolcan Tacana, municipioofUnion
cence a terminal, erect, multiflowered panicle Juarez, 23 Nov 1980 (fr), Breedlove& Almeda 47727(CAS,
mostly 8-30 cm long, the ultimate units con- TEX).GUATEMALA.SanMarcos:VolcanTajumulco,8Apr
sistingofumbelliformorsubverticillateclusters; 1965 (sterile), Andrle 521 (US); nearAldea Fraternidad, be-
bracts and bracteoles 0.5-2.5 x 0.25-0.5 mm, t1w9e6e3n(Sfla),nWRialfaleilamPsieeldeall.a2C6ue2s0t0a(aFn,dGP,alUoS)G.orCdOo,ST1A0-1R8ICDAe.c
subulate, sessile, caducous and typically absent Cartago: Volcan Turrialba, between Hacienda Volcan and
at(atotraunastn.htehCseaislsiy.sx)Petcduaibmceepla(sn0u.4l5-—a1)t21e—,m2m4m-6lmonmgl.omnHg,ylpoanasgncettnhodiitunhmge C(RPhauNifeYrab,ielqlUo,uSiS):1a.7nMtHSoaeenrpCterdeu1iz9aR,6:e71yN4(flaOW)cb,toLsvle1eon9pt6eB7so1q2(ofu9fle)1,VtMeo(,alCcgR2au,1niFrJB,eualM&rOv1M)a9a.7ng1euPai(ArfrlN)Re,iA6oCM1r4SAo1aa.n5t
or spreading and flangelike; calyx lobes evident 15713(F,MO);Boquete,20Mar1977(fr),Folsom2207(CAS);
as ill-defined undulations or depressed-ovate to N side ofRio Calderabetween Horqueta and Bajo Mono, 3
semicircular, 1-1.5 mm long and 4-5 mm wide Jan 1975 (fl), Luteyn& Wilbur4583(CAS, DUKE, F, MO);
basallybetween sinuses. Petals 1.5-1.8 cm long, valleyofRioChiriquiViejo, 14Apr1938(fr), White79(MO,
NY); slopes above Rio Caldera beyond Bajo Mono, 17 Jan
0.9-1.2 cm wide, magenta, glabrous, oblong- 1970(fl), Wilburetal. 11086(DUKE,GH, LL, MO, NY).
ovate, obliquely rounded apically, the margin
entire. Stamensdimorphicandgeniculate at the All Mesoamerican populations ofthis species
point offilament insertion; filaments (6—)7.5—10 are assignable to the nominate subspecies that
mm
long, magenta, glabrous and complanate; does not occur in South America. The Colom-
antherthecaepurple,glabrous,stronglysubulate, bian population of M. macrophylla, which is
mm mm
7 longand 1.5-2.5 widewithadorsally known only from fruiting material, appears to
inclined apical pore. Connective ofthe antepe- representanundescribedsubspecieswhereasthe
talousthecaeprominentlythickeneddorsallyand populationsknownfromthenorthernandsouth-
modified at the filament insertion into an erect, westernmountainsofVenezuelaareattributable
mm
acutely bifid appendage 3 long with a for- to subspecies costanensis and subspecies meri-
mm
wardly projecting arm 4 long and an out- densis respectively (Wurdack, 1978).
mm
wardly projecting arm mostly 2 long. Con- Despite its broad but disjunct distribution,
nectiveoftheantesepalousthecaealsothickened subspecies macrophylla is rare throughout its
dorsallyandmodifiedintoan incurvedbifidap- known range. Ongoing destruction ofits cloud
mm
pendage 4 long with each divergent arm foresthabitatpresentsaseriousthreatto itssur-
mm
typically 1.5-2 long. Ovary (at anthesis) vival. Unfortunately, it is not yet known from
subglobose,glabrous, 5-celled, becominground- any protected park or reserve in the Mesoamer-
ed-lobulate and umbilicate apically at maturity, ican region.
superior or adnate to the hypanthium only ba- Thelarge,handsomefoliage,straightbolewith
mm
sally. Style 16 long,glabrous, somewhatde- sparingly flaky bark, and showy floriferous pan-
cimate and slightly incurved apicallyjust below icle makeM. macrophylla one ofthe most spec-
thetruncatestigma. Fruitinghypanthiumessen- tacular arborescent melastomes in the neotrop-
tially glabrous, shallowly campanulate to hemi- ics.
ALMEDA: MESOAMERICANSPECIESOFMERIANIA 147
Thestaminalmorphologyofthisspeciesisex- sepalous) stamens inserted opposite the calyx
mm
traordinary. The dorsally thickened connective lobes.Antepetalousstamens:filaments9-10
mm
on the antepetalous stamens (Fig. 1A) is es- long,antherthecae9-12 long,yellow,linear-
pecially thick and fleshy but nothing is known subulateandarcuatewithadorsallyinclinedapi-
mm
about its functional significance, ifany, in pol- cal pore, connective prolonged 1 belowthe
lination. thecae and dilated dorso-basally into an acute
mm
red spur 0.5 long and an oblong red ap-
pendage 3 mm long directed ± parallel to the
Meriania odorata Almeda, sp. nov. thecae. Antesepalous stamens: filaments 13-14
(Fig. 2) mm long, anther thecae 5 mm long, yellow, su-
Type.—Panama. BocasdelToro:vicinityofFortunaDam, bulate, and somewhat rostrate with a ventrally
alongroadtoChiriquiGrande,08°50'N,82°15'W,ca.470m, inclined apical pore; connective dilated dorso-
6 Sep 1987(fl),McPherson 11651(Holotype:CAS!;isotypes: basallyintoan acuteredspur0.25 mm longand
MOA!r,boPrMA8!-1,5UmS!;).ramulirotundato-quadrangulatidemumter- amfmorwardly projecting oblong appendage 1.5-2
etessicutfoliainflorescentiahypanthiaqueubiqueglabri. Pe- long.Ovary(atanthesis)subcylindrictonar-
tioli 2-5 cm longi; lamina 6.5-25.6 x 4.2-10.8 cm elliptica rowly ellipsoid, glabrous, 5-celled, becoming
vel elliptico-ovata apice acuminata basi acuta vel obtusa, rounded-lobulateandumbilicateapicallyatma-
5-nervata, coriacea et integra. Panicula 6-20 cmmlomnga mul- turity, superior. Style 12-14 mm long, glabrous,
tHiyflpoaran,tfhlioruems(5a-dmetroir,umpe)d4ic-e5llmism(aldonagnutmh;esciaml)yx6-19.5mmlloonnggiuss. declinateandhookedapicallybelowthetruncate
truncatus. dentibusexterioribus non evolutis. Petala 1.7-2.5 stigma. Fruiting hypanthium glabrous, subur-
x 1-1.7cmasymmetriceobovata.Staminadimorphaglabra: ceolate, terete, 6-7 x 7 mm. Seeds dark brown,
amlm1na.otm5inhogemrarlamtoi;n,sugmfaopi.bdlooearSnnmotgteaonedmt-obiarsasnu9sabaal-uli1lmit0aie0tnrm.io5isrmnacmp:lloimonfrnaiogtalaoca0;;um.teac7onno5,ttn-aah1npee1prcm3eat-nrmi1duv4imucdmmetiham0aem.sc5.caleeomnSn9dgtm-eaan;1mtp2iarinnoxa--3 cvtoehuprenpnoewisaciitodteseeerw,eientsdnht,draa2bin-guh3titntmfoalrmpateseroldeomdnwegiaw.nnhgdlait±keccuaarpuvpdeeadtn,edlaaigtneetahra-et
therarumthecae5 x 1.5mm,poroventraliterinclinato;dente Distribution and Phenology.—Local and
basali0.25mmlongoacuto,appendiceascendentmim1.5-2mm uncommoninmontanerainforestfromsouthern
longa.Ovarium 5-loculareglabrum;semina2-3 longa. Costa Rica (Limon) to western and central Pan-
Tree8-15 mtall,essentiallyglabrousthrough- ama at elevations of470-1,450 m. Floweringin
out with rounded-quadrate internodes. Leaves May, July, and September; fruitingin February,
ofapairequaltoslightlyunequalinsize;petioles May, and December.
2-5 cm long and 2 mm wide; blades coriaceous
when dry, 6.5-25.6 x 4.2-10.8 cm, elliptic to Paratypes.—CostaRica. Limon: ParqueInternacionalLa
elliptic-obovate, 5-nerved with a prominulous Amistad,FilaTsiurabeta,entreRiosUrenyLari,9°27'30"N,
networkofsecondaryandhigherorderveinsev- 83°00'00"W, 25 Jul 1989 (fl), Chacdn 288 (CAS. CR, MO).
PANAMA.Chiriqui:vicinityofFortunaDam,8°45'N,82°5'W,
identabaxially,baseacutetoobtuse,apexblunt- 15Dec1987(fr),McPherson11794(CAS).Code:CerroGaital,
ly acuminate, the margin entire. Inflorescence a E slope and ridges leadingto the summit, 8°40'N, 80°07'W,
terminalmultifloweredpanicle6-20cmlong,the 24 Feb 1988 (fr),Almeda etal. 5900(CAS, CR, MO, PMA,
ultimate units consistingofumbelliform orver- US); hillsN ofEl Valle, E slope and ridges leadingto Cerro
ticillate clusters; bracts and bracteoles evidently VGaeirtaaglu.a8s°:40a'bNo,ve80S°a0n7t'aW,Fe30anMdaaybo19v8e2A(lftl)o,KdneaPpiped5ra3,28on(CrAiSd)g.e
earlymcamducousandnotseenatanthesis. Pedicels trail to top ofCerro Tute-Arizona, 8°30'N, 81°10'W. 6 Feb
6-9 long. Hypanthium (at anthesis) cam- 1988(fr).McPherson 12080(MO).
mm
panulate, 4-5 long to the torus. Calyx tube
1.5mmlong,spreadingandflangelike;calyxlobes Allbutoneofthesixknowncollectionsofthis
obsoleteorconsistingofill-defineddepressedun- species were made in the last six years. It seems
dulations on flowering and fruiting hypanthia; surprising that an arborescent melastome with
calyx teeth obsolete. Petals 1.7-2.5 cm long, 1- showy, fragrant flowers should have gone un-
1.7 cm wide, pale pink, glabrous, obovate, noticed for so long. Rainy season flowering and
rounded to obliquely emarginate or irregularly theapparentrestrictionofM. odoratatothewet-
lobulate apically, the margin otherwise entire. ter,lessaccessibleslopesoftheCaribbeandrain-
Stamens dimorphic and unequal in length with ageinsouthernCentralAmericaaccount,inpart,
thelarger(antepetalous)stamensinsertedonthe for the paucity ofcollections.
torus opposite the petals and the smaller (ante- ThedistinctivefeaturesofM. odoratainclude
148 PROCEEDINGSOFTHECALIFORNIAACADEMYOFSCIENCES,Vol. 48,No. 7
Figure2 Merianiaodorata.A,habit, x ca. 1/3;B,representativeleaf(abaxialsurface), x 1/4withenlargement(left)ofarea
betweentwotransverse secondaryveins; C, fruitinghypanthium, x 2; D, seeds, x 7; E, petals, x 1; F, antesepalousstamen
(left)andantepetalousstamen(right), x ca. 2.(A,B,E,Ffromtheholotype;CfromKnapp5328- DfromAlmedaetal. 5900.)
ALMEDA: MESOAMERICANSPECIESOFMERIANIA 149
itsoverallglabrosity,entireleaves,palepinkpet- minal, openly branched multiflowered panicle
als, flangelike calyx with obsolete or ill-defined mostly 9-20 cm long, the ultimate units con-
lobes(Fig. 2C), absenceofcalyxteeth, andcom- sisting of simple cymes; bracts and bracteoles
parativelylarge,appendaged,linear-cuneateseeds 0.3-2.5 cm longand 0.5-5 mm wide, linear-ob-
(Fig. 2D). long with a cuneate base and rounded apex, the
Among its congeners, M. odorata appears to petioles 1-1.5cmlong,earlycaducousandrarely
mm
be most closely allied to M. panamensis, which seen at anthesis. Pedicels 2-5 long. Hypan-
mm
is known only from higher elevations (1,850- thium (at anthesis) campanulate, 4-7 long
3,000 m) on the northern slopes ofVolcan Baru tothetorus. Calyxtruncateandflangelikeinbud
(= Volcan Chiriqui) in western Panama. Both butrupturing(onfruitinghypanthia)intobluntly
mm
species are glabrous throughout and share a di- rounded or truncate lobes 2-A long and 5-
mm
morphic androecium with each set of stamens 7 wide basally between sinuses, each calyx
differingin apical pore orientation (Figs. IFand lobe inconspicuously beset with a blunt subter-
2F). Inadditiontothediagnosticcharacterspre- minal abaxial tooth or callosity. Petals 2.6-3.9
sented in the key, M. panamensisalso differs in cm long, 1.6-3.7 cm wide, magenta or violet,
having an ill-defined dorso-basal appendage on glabrous, obovate, rounded to emarginate api-
theantesepalousstamens(Fig. 1F)andmarkedly cally, themarginentire. Stamensdimorphicand
different seeds (compare Figs. 1G and 2D). unequal in length with the larger (antepetalous)
Label information on Chacon 288, Knapp stamensinsertedonthetorusoppositethepetals
5328, and McPherson 11651 note that the an- andthe smaller(antesepalous)onesinserted op-
thers ofthis species are yellow. The latter two positethecalyxlobes.Antepetalousstamens: fil-
mm mm
collectorsdescribe the anthersasyellowand the aments 11-12 long,antherthecae 10-12
anther tails as red. I interpret this to mean that long, magenta dorsally and white ventrally, lin-
theantherthecae areyellowand theappendages ear-subulate and somewhat arcuate with a dor-
are red. Ifcorroborated by additional field ob- sally inclined apical pore; connective prolonged
mm
servations, coloration of the androecium will below the thecae for 1 and dilated dorso-
provide another difference between M. odorata basallyintoashortacute ± erectyellowspurand
mm
and M. panamensis; the latter has magenta an- abluntlinear-oblongyellowappendage3-3.5
ther thecae and yellow appendages. long directed ± parallel to the thecae. Antese-
mm
The epithet for this species is derived from palous stamens: filaments 15-16 long, an-
mm
odorus, Latin for fragrant or having a smell, to ther thecae 6.5-7 long, brownish dorsally
emphasizetherose-scentedfragranceoftheflow- and white ventrally, subulate with a ventrally
ers. inclined apical pore; connective dilated dorso-
basally into a yellow erect spur and a forwardly
mm
projecting rounded appendage 0.5 long.
Meriania panamensis Gleason in Woodson and
Ovary (at anthesis) ellipsoid, glabrous, (4-)5-
Schery, Ann. Missouri Bot. Gard. 28:436. 1941.
celled, becoming rounded-lobulate and umbili-
(Figs. IF,G) cate apically at maturity, superior or adnate to
Type.—Panama.Chiriqui:CasitaAhatoCerroCopete,Vol- the hypanthium basally for about 1 mm. Style
mm
candeChiriqui.2,300-3,300m, 10Jul 1940(fl,fr), Woodson 15-17 long, glabrous, somewhat declinate
&Schery361 (Holotype:NY!;isotype: MO!). and strongly hooked apically. Fruiting hypan-
thium glabrous, campanulate, essentially terete
Tree4-10mtall,essentiallyglabrousthrough- orbluntlycostatewhendry, 9-11 x 12-13 mm.
outwithtereteinternodes. Leavesofapairequal Seedsbrown, vernicose, straight or curved, nar-
oronly slightly unequal in size; petioles 1.8-4.5 rowlycuneiformandangulate, taperedtoabeak
cm long and 1.5-2 mm wide; blades coriaceous at the wider end when dry, 1-2 mm long.
when dry, (3.2)7-18.5 x 3.2-8 cm, broadly el- Distribution and Phenology.—Known only
liptic varying to elliptic-ovate, 5-nerved with a from western Panama where it is local and un-
prominulous network of secondary and higher common in cloud forests at 1,860-3,000 m.
order veins especially evident abaxially, base FloweringcollectionshavebeenmadefromJan-
acute, apex cuspidate, the margin coarsely and uarythroughMarchandinJuneandSeptember;
sometimesremotelydentatealongtheapicalhalf fruiting specimens have been gathered in Janu-
or two-thirds ofthe blade. Inflorescence a ter- ary, June, and September.
150 PROCEEDINGSOFTHECALIFORNIAACADEMYOFSCIENCES,Vol.48,No. 7
Representative Specimens Examined.—PANAMA. Chi- obscurely cordate, apex acute to short-acumi-
riqui:8kmWofCerroPuntainvicinityofLasNubes, 11 Feb nate,themargindenticulatetosubentire,thead-
1978 (fl), Almeda & Nakai 3526 (CAS); Bajo Chorro, E of axialsurfaceglabrousbutnotbullate,theabaxial
C1e0r4r0o(PCuAnSt,a,M8O°)5;3'cN,am8i2n°o32d'eW,Lo5sJPuonzo1s97a9C(oflt,itfro),yAnNtuoenviao surface covered with shaggy-plumulose hairs on
ZelandiaN,W13 Mar 1990(fl),AWrandaetal. 1074(CAS, PMA); the primary and secondaryelevated veinsand a
2.7 mi ofRioChiriqui, ofCerroPunta, 27 Feb 1973 stellate-hirsute indument between the veins. In-
(fl), Croat22351(CAS,MO,NY); CerroPunta, 14Sep 1971 florescence a terminal multiflowered, long-pe-
M(Bfcor)qP,uhLeeatroeso3nne9a9r9(3BM3aO4j,o(UGCSrA)aS,n,daelM,oOn8)g°;5r0oa'aNbd,ofv8re2o°mC3e5Cr'erWro,roP5PuJunutnnat,a1t98o8°w65a0r('fdNl)s,, d2u1nccumlaltoeng(,9-t1h7e culmt)imvaertteicuinliltasteappapneiacrlien(g9—f)as1c3i—-
82°35'W, 9 Feb 1986 (fl), McPherson& Morello8380(CAS, cledorumbelliform;bracteoles20-30 x 3-4mm,
MO); PaseodeRespingaonwaytoBoqueteattopofdivide, linear-lanceolate, early deciduous and usually
14 Jan 1971 (fl), Wilburet al. 13196 (DUKE, F, GH, MO, seen only on young inflorescences. Pedicels 6-
NY). mm mm
12 long, lengthening to as much as 23
Meriania panamensis is readily recognized in fruit. Hypanthium (at anthesis) campanulate,
evenin sterilecondition by itsglabrous leathery 5-6 mmlongtothetorus. Flowerbudsrounded-
leaves that are coarsely dentate along the apical ellipsoid, 9-13 x 4-7 mm, rounded to bluntly
halfortwo-thirdsoftheblade. Inadditiontothe acute orbluntly rostrate apically. Calyx fused in
line drawings of stamens and seeds presented bud but rupturing into two irregular rounded-
here (Figs. IF, 1G) see Gleason (1958:208) for deltoid lobesthat fall away afteranthesis. Petals
otherillustrationsofsomediagnosticcharacters. 1.1-1.6cmlong,0.8-1.3cmwide,greenish-white,
SeeM. odorataforadiscussionoftheprobable somewhat translucent when fresh, the venation
relationships ofM. panamensis. darkened and conspicuous when dry, erect, ob-
ovate, ± concave and connivent, rounded api-
Meriania phlomoides (Triana) Almeda, comb, cally, the margin entire. Stamensnearlyisomor-
nov. phic, very similar in size, differing slightly in
(Figs. 1H,I,J) detailsofconnectivemodification.Antepetalous
mm
CentroniaphlomoidesTriana, Trans. Linn. Soc. London 28: stammenms: filaments 8-9 long, anther thecae
72. 1871. 6.5 long, white, subulate, horizontalandes-
sentially straight with a dorsally inclined apical
Tytpyep.e:—CBo,sdteastRriocyae,d;wipthhootuotgreaxpahcst:loCcAaSli!t,y,FO,erUsS!t)e.d5A5no(tShyenr- pore; connective thickened, ± flat ventrally but
syntype,Hoffmanns.n.,wascitedintheprotologuebut,to romunmdedandsomewhatridgeddorsallyalongthe
date,Ihavebeenunabletolocateoriginalorduplicatema- 1 segment prolonged below the thecae, di-
terial ofeithercollection. Oersted'scollectionsfrom Costa lated dorso-basally into a spreading ridged and
RicaarepreservedatCopenhagen(C)butnotypematerial adaxiallyfurrowedappendage(2 x 1.5 mm)and
o2f5thAiusgspe1c9i8e8s).apApecacrosrdtionbgetaomoSntagntdlheeym((1B.93H7a)n,seCanr,linHoliftft.-, a blunt adjacent spur elevated aboutm0.m5 mm.
mann'scollectionswereatBerlin(B),allofwhichwereev- Antesepalous stamens: filaments 7-8 long,
mm
identlydestroyedduringWorldWarII. Irefrainfromneo- antherthecae 6-6.5 long, otherwiselikean-
typifying this species pending a more extensive search of tepetalous stamens in color, posture, and pore
Europeanherbaria.
inclination; connective flattened ventrally but
m mm
Shrub or tree 2-12 tall. Older cauline in- somewhatelevatedandroundedalongthe 1
ternodes ± terete to rounded-quadrate and gla- segmentprolongedbelowthethecae,dilateddor-
brate, the rounded-quadrate uppermost bran- so-basally into a spreading, flattened, longitu-
chlets, inflorescence axes, pedicels, floral buds, dinally ridged appendage (1.5-1.75 mm) and a
andhypanthiacoveredwithbrownishbasallyen- smaller adjacent tooth elevated 0.5 mm. Ovary
mm
larged hairs (0.5-1.5 long) that are shaggy- (at anthesis) ovoid, glabrous, 5-celled, rounded-
mm
plumulose basally but sparingly barbellate to lobulateapically,superior.Style 16-17 long,
glabrous toward the apex. Leaves ofa pair es- glabrous, erect or only slightly declinate and
sentiallyequaltoslightlyunequalinsize;petioles barelycurvedapicallybelowthetruncatestigma.
1.5-1.75 cm long and 1.5-4 mm wide; blades Fruiting hypanthium cupulate, costulate, 6-7 x
membranaceous when dry, 10.5-26 x 6-15.5 5-7 mm. Seeds beige, cuneiform or oblong-pyr-
mm
cm, broadly elliptic to elliptic-lanceolate, 5-7- amidate and angulate, 0.5-1 long.
—
plinervedwithawell-definednetworkofsecond- Distribution and Phenology. Local and
aryandtertiaryveins,baseobtusetoroundedor uncommon fromnorthwesternCostaRica(Cor-