New Taxa and Combinations in the Ocotea helicterifolia (Lauraceae) Species Group

Author(s): Henk van der Werff

Reviewed work(s):
Source: Novon, Vol. 9, No. 4 (Winter, 1999), pp. 571-583
Published by: Missouri Botanical Garden Press
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 New Taxa and Combinationsin the Ocotea helicterifolia
(Lauraceae) Species Group
Henk van der Werff
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

butwhichdid nothave the pu-

a treatment of 0. helicterifolia,
ABSTRACT. In thecourseofpreparing
of the Ocotea helicterifoliagroup for Flora bescentleaves and twigs.He notedthattheO. hel-
Mesoamericana, theexistingtreatment was foundto ictertfolia groupis related to the Ocotea sinuata
be outdated.A reviewof the groupis here pre- group,whichdiffersin havingtongue-shapedan-
sentedand includesthedescription ofthefollowing therseach witha conspicuoussteriletip.
new species: O. congregatavan der Werff, O. cor- In thecourseofwriting a treatment ofOcoteafor
rugata van der Werff, O. gordonii van der Werff, Flora Mesoamericana, I found several undescribed
and O. patula van der Werff, as well as thefollow- species in the O. helicterifolia groupand several
ing new combinations: 0. betazensis(Mez) van der otherspecies thatneeded to be transferred to Oco-
Werff, O. bourgeauviana (Mez) van der Werff, O. tea. A fewspecies of this groupoccurredoutside
purpurea(Mez) van der Werff, and O. tonii(Lun- the area coveredby Flora Mesoamericana.In ad-
dell) van der Werff.A key to the species of the ditionto thenovelties,I decided it mightbe useful
groupis presented, newsynonymy is given,and the to publisha key to all species I recognizeas be-
specimensstudiedare listed. longingto theO. helicterifolia group,as wellas new
synonyms. In the mostrecenttreatment of Central
Amongthespecies ofOcoteain CentralAmerica
American Lauraceae (Allen, 1945), most species of
is a groupcharacterized the O. helicterifoliagroup were included in Phoebe,
bythepresenceofan erect
indumenton the leaves (especially on the lower and these species were mostly separated based on
thatis discernible to the with dense- leaf size and leaf shape. I foundthese characters
surface) touch,
less important than Allen did, and rely more on
ly to moderatelypubescenttwigs,tepals thatare
partiallypapillose (sometimesonlyalong the mar-
such characters as inflorescence type(racemoseor
paniculate-cymose), flower characters (flowersgla-
ginor nearthetip),glabrousor somewhat papillose
antherswiththe fourcells arrangedin twosuper- brous or pubescent; inner surface oftepals glabrous
and, at least in some or pubescent; receptacles glabrous or pubescent in-
posed pairs species,relatively
side), and leaf position (alternate or clustered). Use
well-developedstaminodia.The termpapillose is
used here fora dense cover of veryshort,curly of thesecharactersleads to better-defined species,
hairs. In a few species the antherseach have a although some of the species appear quite variable
small,steriletip and theanthercells do notfillthe and may be furtherdivided at some later point.
anthercompletely, as is thecase in mostspecies of Specifically, the Costa Rican specimensof O. hel-
Ocotea. Althoughthis groupis easy to recognize, icterifolia seem slightly different and occurat lower
its taxonomy is confused.The presenceofwell-de- altitudes than specimens collected northof Costa
staminodia has resulted in the inclusionof Rica. Likewise, specimens of 0. purpureafrom
veloped
several species in Phoebe, and later those were Panama seem different (fewerlateralveins,forex-
transferred to Cinnamomum.Otherspecies have ample) than those from Hondurasnorthward. How-
been placed in Nectandra,sharinga commonpap- ever, splitting these species can only be done using
illosityof the tepals and anthers. Nectandra beli- vegetative characters (leaf shape and size), and be-
zensis(Lundell)Allenresemblesin vegetative char- cause I regardthese charactersas weak and not
acters the O. helicterifolia group,but has typical reliable,I am reluctanttofurther dividetheserath-
Nectandrastamens(short,broad,withthe anther er variable species.
cells arrangedin a shallowarc, not in twosuper- In severalspecies the upperrimofthe recepta-
posed pairs as in Ocotea). Nectandrabelizensisis cle carriesa ringof hairs. These hairs are easily
knownfromBelize, Costa Rica, and Panama. Roh- visibleand maysuggestthereceptacleitselfis pu-
wer(1991) gave a briefdiscussionofthegroupand bescent or the tepals are pubescenton the inner
provideda list of taxa includedin it. He also re- surface.It is necessaryto split a receptacleopen
vised the species withthe generalflowerstructure in orderto ascertainwhetherthe receptacleis pu-
NovoN 9: 571-583. 1999.
572 Novon

bescentinside:a viewfromabove is notsufficient. lengthofthetepals;a tuftofhairsvisibleat ornear

I only describetepals as pubescenton the inner the base of the tepals is not enoughto score the
surfaceif the hairs extend at least halfwaythe innersurfaceofthe tepals as pubescent.

KEY TO SPECIES OF THE OCOTEA HELICTERIFOLIA GROUP IN CENTRAL AMERICA

la. Inflorescences
racemoseor rarelywithone or twolateralcymes.
2a. Leaves obovate;tuftsor lines ofwhitehairscommonalong majorveinson lowerleaf surface......
............................................................ 15. O. sp. nov.
2b. Leaves elliptic;axillarytuftsofhairslackingor onlya fewon each leaf.
3a. Leaves bullate-rugose, the majorveinsstrongly impressedon the upperleaf surface........
......................................................... 4. 0. corrugata
3b. Upperleaf surfacesmoothor nearlyso, the majorveinsimmersed, notstrongly impressed.
4a. Outersurfaceof tepalsglabrous;antherssessile ................... .. 5. O. gordonii
4b. Outersurfaceof tepals at least sparselypubescent;stamenswithfilaments 1/3or moreof
the lengthof theanthers.
5a. Hairs on lowerleaf surfaceascending,coveringmostof thelamina;indument gray ..
................................................. 8. 0. mollicella
5b. Hairs on lowerleaf surfaceerect,mostof the lamina visible; indumentbrownor
ferruginous .........................................11. 0. purpurea
lb. Inflorescences
paniculate.
6a. Leaves clustered.
7a. Outersurfaceof tepalsglabrous................................ 2. O. bourgeauviana
7b. Outersurfaceof the tepalspubescent.
8a. Petiolesto 6 mmlong;leaves 15-25 x 5-6 cm; twigswithyellowishbrownindument
12. O. tonii
8b. Petiolesat least 10 mmlong;leaves 9-17 x 4-7 cm; twigswithbrownindument........
.................................................... 3. 0. congregata
6b. Leaves alternate,evenlydistributed alongthetwigs.
9a. Receptaclepubescentinside.
10a. Leaves elliptic,to 15 cm long ................... ............. .1. O. betazensis
10b. Leaves obovate,18-40 cm long.
lla. Indumentcompletely coveringyoungtwigsand inflorescences...... .14. O. valerioides
llb. Surfaceof twigsand inflorescences visiblebetweentheindument........ 7. 0O.lentii
9b. Receptacleglabrousinside.
12a. Outersurfaceoftepalspubescent.
13a. Leaves broadlyelliptic;tuftsof whitehairslackingin axils of lateralveins or along
the majorveins................... .................... 9. 0. patula
13b. Leaves obovate;tuftsof whitehairs presentin axils of lateralveins and along the
majorveins......................................... 15. 0. sp. nov.
12b. Outersurfaceof tepalsglabrous.
14a. Innersurfaceoftepalspubescent;filaments evident,ca. 1/3thelengthoftheanthers.
15a. Leaves to 10 cm long,the tipsobtuse,acute or shortly acuminate.........
.......................................... 10. O. praetermissa
15b. Leaves 13-20 cm long,acuminateor graduallynarrowedintoa slendertip .....
.......................................... 2. 0. bourgeauviana
14b. Innersurfaceof tepalsglabrous;antherssessile.
16a. Surfaceofyoungtwigscompletely coveredbytheindument; cupule cup-shaped
............................................ 13. O. valeriana
16b. Surfaceofyoungtwigspartiallyvisiblebetweentheindument; cupuleonlyshal-
lowlybowl-shapedor platelike ...................... 6. O. helicterifolia

1. Ocotea betazensis (Mez) van der Werff,

comb. Ocotea betazensis (as Phoebebetazensis) has long
nov. Basionym: Phoebe betazensis Mez, Jahrb. been includedin Ocoteahelicterifolia, fromwhichit
Koenigl.Bot. Gart.Berlin5: 192. 1889. SYN- differs in theellipticto broadlyelliptic(notobovate)
TYPES: Mexico. Oaxaca: Liebmann 2(C), leaves,thestamenswithwell-developed the
filaments,
3(C), 22(C), 23(C), Galeotti 2885 not seen, dense tomentellous-tomentose indument on theyoung
Juergensen575 not seen. Figure 1. twigsthatcompletely coversthe surface(nothirsute
withthe surfacepartially visible),and the generally
Oreodaphnemexicana Meisner var. diminutaMeisner,
Prodr.15(1): 118. 1864. Ocoteamexicana(Meisner) longerpetiolesthanseen in 0. helicterifolia. Ocotea
Hemsley var. diminuta (Meisner) Hemsley,Biol. betazensisis currentlyonly known from cloud forests
Centr.Amer.,Bot. 3: 73. 1882. TYPE: Juergensen between1900 and 2600 m altitudein Oaxaca, but
575 notseen. can be expectedin neighboring Chiapas.
Volume9, Number4 van derWerff 573
1999 Ocotea helicterifolia
Species Group

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_1-:11:

Figures1-4 (clockwisefromtop left).-1. Ocotea betazensis:Cedillo 1197. -2. Ocotea bourgeauviana:Breedlove

50583. -3. Ocoteacongregata:MendezTon9594. -4. Ocoteacorrugata:Wendt et al. 6765 (isotype).
574 Novon

2. Ocotea bourgeauviana (Mez) van der Werff, the hairs erect,soon becomingglabrous,the pu-
comb. nov. Basionym:Phoebe bourgeauviana bescence denser and persistingalong the major
Mez, Jahrb.Koenigl.Bot. Gart.Berlin5: 194. veins,lowersurfacemoderately denselypubescent,
1889. Cinnamomumbourgeauvianum(Mez) the hairs erect and discernibleto the touch,the
Kostermans, Reinwardtia6: 20. 1961. TYPE: surfacereadilyvisible betweenthe hairs,the in-
Mexico.Veracruz:Bourgeau2234 (isosyntype, dumentdenserand tomentose alongthemidriband
MO). Figure2. lateralveins;domatialacking;petioles10-25 mm,
round,witha similarindumentas the twigs.Inflo-
Nectandralongicuspis Lundell,Wrightia 5: 34. 1974. Syn.
nov.TYPE: Guatemala.Izabal, Contreras 11186 (iso- rescences5-12 cm,paniculate-cymose, theflowers
type,MO). in compact clusters, densely hirsute-tomentose,
mostlyin the axils of bracts,infrequently in the
Phoebechinantecorum R. E. Schultes,Bot. Mus. Leafl.9:
axils ofleaves. Flowers7-9 mmdiam.,white,per-
170. 1941. Syn. nov. Cinnamomum chinantecorum
(R. E. Schultes) Kostermans,Reinwardtia6: 20. fect, the receptacle densely pubescent outside;
1961. TYPE: Mexico.Oaxaca: Schultes& Reko827 pedicels ca. 3 mmlong.Tepals 3 mmlong,elliptic,
(holotype,
GH). the outsidemoderately to sparselypubescent,the
insidepubescentnearthebase, otherwise glabrous,
Ocotea bourgeauviana can be recognized by the
spreading at anthesis,outer6 stamens1.6 mm,ses-
combinationof paniculate-cymose inflorescences,
sile or nearlyso, witha fewhairs near the base,
glabrousflowers, pubescentinnersurfaceofthete-
otherwiseglabrous,the cells arrangedin 2 pairs,
pals, and the mostlyclusteredleaves. The indu-
at the tip witha narrow,sterileborder,
menton thetwigsis hirsute,withthesurfaceofthe introrse,
inner3 stamens1.7 mm,thefilament 0.5 mm,with
twigspartiallyvisiblebetweenthe indument.Leaf
a few hairs,the cells in 2 pairs, extrorse-lateral,
size rangesfrom13 to 20 cm long.It is knownfrom
at the base of the filaments, stami-
Mexico (Veracruz,Chiapas), Guatemala,and Hon- glandspresent
nodia 3, minute,stipitiform, hidden betweenthe
durasat altitudesrangingfrom100 to 1200 m,but
hairs on the top of the receptacle;pistil 1.5 mm,
is infrequently collected.Provisionally placed here
glabrous,thestyle0.4 mm,receptaclecup-shaped,
are somecollectionsfromVeracruz(Mexico)witha
appressedpubescentor glabrousinside. Fruitel-
glabrousinnersurfaceof the tepals and slightly
lipsoid,2 x 1.3 cm,thecupule deeplycup-shaped
smallerflowers(4-5 mmdiam. vs. 5-6 mmin O.
whenyoung,bowl-shapedat maturity, 1 cm diam.,
bourgeauviana).Because thesespecimensdiffer in
witha singlemargin,thetepalsdeciduous.Flowers
onlyone solid character(theglabrousinnersurface March,
April,November;fruitsAugust,October.
ofthetepals)fromO. bourgeauviana, I am reluctant
to recognizethem as a distinctspecies and list Ocotea congregatais namedafterthe erectsta-
themin thespecimensstudiedas O. aff.bourgeau- mens groupedin a rathertightcluster.The new
viana. taxon is currently only knownfromthe Mexican
stateofChiapas and occursbetween800 and 1370
3. Ocotea congregata van der Werff, m altitude.It can be recognizedbytheslightly bul-
sp. nov.
TYPE: Mexico. Chiapas: Mpio. Oxchuc,Cas- late,loosely clustered leaves, thepubescent flowers
cada de Coralito,ShilomTon8930 (holotype, arrangedin paniculate-cymose inflorescences, the
brownto dark brownindumenton the twigs,and
MO). Figure3.
theratherlong(10 mmormore)petioles.Its closest
Ocoteaetoniisimilis, sed foliisbrevioribus,latioribus,relativeis Ocoteatonii,whichdiffers in itsnarrow-
petiolisplusquam10 mmlongis, indumento castaneo dif- er,
longer leaves, shorter petioles (to 6 mmlong),
fert.
and the yellowishbrownindumenton the young
Small to medium-sized trees,to 15 m tall.Twigs twigs.
terete,solid,denselybrown-tomentose ortomentel- Paratypes. MEXICO. Chiapas: Mpio. Ocosingo,
lous, the hairserectand twisted,coveringthe sur- Breedlove15672 (CAS), 33017 (CAS, MO), 52589 (CAS,
face of the youngtwigscompletely, terminalbuds MO),MartinezS. 17084 (MO), Quintanilla49 (MO); Mpio.
denselybrown-tomentose. Leaves 9-17 x 4-7 cm, San Cristobalde las Casas, MendezTon9594 (CAS, MO);
Mpio. Oxchuc,ShilomTon8503 (MO).
ellipticto broadlyelliptic,firmly chartaceous,clus-
tered,thebase roundedor obtuse,rarelyacute,the 4. Ocotea corrugata van der Werff,sp. nov.
apex obtuseto acute,midriband lateralveins im- TYPE: Mexico.Oaxaca: Mpio. Sta. MariaChi-
pressed,tertiary venationweaklyimpressedon the malapa, Sierrade Tres Picos, alt. 1150-1250
uppersurface,midriband majorveinsprominently m, T Wendt, HernandezG., Tenorio, Torres, Sa-
raised, smallerveins raised on the lowersurface; lazar, Soto & Rocha 6765 (holotype,MEXU;
upper surfacemoderately pubescentwhenyoung, isotype,MO). Figure4.
Volume9, Number4 van derWerff 575
1999 Ocotea helicterifolia
Species Group

Ad gregemOcoteaehelicterifoliae sed foliis locality.However,the remotenessof this locality

pertinens,
diversaest.
corrugatis to bringback morematerial.
made it difficult
Small tree,to 4 m. Twigsterete,denselybrown-
thesurfaceoftheyoungtwigsnotvis-
5. Ocotea gordonii van derWerff, sp. nov.TYPE:
tomentellous,
Panama. Chiriquf:vicinityofFortunaDam, G.
ible, solid; terminalbuds densely tomentellous.
Leaves 7-14 x 2.5-7 cm,ellipticor broadlyellip-
McPherson10421 (holotype, MO). Figure5.
tic,firmlychartaceous,alternate, thebase variable, InterspeciebusOcoteaehelicterifoliae gregisinflores-
fromacute or obtuseto rounded,theapex acute or centiaracemosa, receptaculo intuspubescente etpedicel-
the surface lis longisrecedit.
obtuse, upper sparselypubescent with
erector ascendinghairswhenyoung,soon becom- Small trees,to 10 m tall. Twigs terete,solid,
ing glabrous,thelowersurfacesparselypubescent, denselytomentose, the surfacenotor scarcelyvis-
the hairs erect and discernibleto the touch,the ible betweenthe hairs; terminalbuds denselyto-
indumentdenserand tomentellous alongthemajor mentose.Leaves 12-22 x 5-8 cm, narrowlyto
veins, midrib,lateral veins, and tertiary venation broadlyelliptic,chartaceous,alternateand evenly
impressed on the upper surface,strongly raised on distributed alongthetwigs,themarginflator some-
the lowersurface,theleaves bullateto rugose,lat- timesfoldeddownward, thebase acute or obtuseto
eral veins 5 to 7, domatiaabsent,petioles 7-15 almostrounded,thetip acuminatewithan acumen
mm,flat above, with a similarindumentas the to 2 cm longor acute,theuppersurfacewithsome
twigs.Inflorescences 3-5 cm, racemose,less than erect,curledhairswhenyoung,but thisindument
10-flowered, denselyhirsute,the flowersgrouped rapidlywearingoff,the hairsdenserand becoming
near the tip of the inflorescence.Flowers7-8 mm tomentosealongthe majorveins,thelowersurface
diam., cream-colored, perfect,receptacledensely copiouslypubescent,thehairserectand softto the
pubescenton the outside;pedicels 2-3 mmlong. touch,denserand tomentosealong the midriband
Tepals 3 mm long, elliptic,on both surfacespu- lateralveins,midriband lateralveinssomewhat im-
bescentnear the base, otherwiseglabrous,spread- pressed,tertiary venationslightlyraisedon theup-
ing at anthesis;outer6 stamens1.5 mm,the fila- per surface,midrib,lateralveins,and tertiary ve-
mentsca. 0.4 mm,withsome hairsnear the base, nationraised or prominently raised on the lower
otherwiseglabrous,the cells introrse, arrangedin surface;lateralveins 6 to 8; domatiaabsent;peti-
2 pairs,a steriletip lacking,inner3 stamens1.5 oles 8-14 mm,flattened on the upperside, witha
mmlong,the filament 0.4 mm long,with2 glands similar indument as the twigs. Inflorescences 8-16
near the base, the cells in 2 rows,the upperrow cm, racemose,moderately denselyto sparselypu-
lateral,thelowerone lateralextrorse, staminodia3, bescent,the hairs erect or spreading,to 0.6 mm
to see amongthe long;in axils ofbractsor,less frequently, ofnormal
clavate,witha fewhairs,difficult
hairson the rimof the receptacle,pistilglabrous, leaves. Flowers 8-10 mm diam., white, glabrous,
ca. 1 mmlong,thestyleveryshort,receptaclecup- fragrant, perfect, pedicels ca. 1 cm long. Tepals 6,
inside. 4-4.5 mmlong,elliptic,spreadingat anthesis,the
shaped,appressedpubescent Fruit and cu-
outer3 witha basal triangular papillosepatch,oth-
pule unknown.FlowersApril.
erwiseglabrous,the inner3 uniformly papilloseon
6
AlthoughOcotea corrugatais onlyknownfrom the innersurface;stamens9, 4-celled,the outer
the typecollection,it is heredescribedlargelybe- weakly papillose, the cells arranged in 2 rows,
cause of its verydistinct,bullateto rugoseleaves; openingintrorse-lateral, theantherssessile ornear-
additional charactersare the racemose inflores- ly so, steriletipshort,0.2-0.3 mm,inner3 stamens
cences and the pubescentinsideofthereceptacle. 1.5 mmlong,the anthersessile, thecells arranged
It belongsto the Ocoteahelicterifolia group,where in 2 rows,laterallyextrorse, theanthersweaklypa-
it does notseem to have a close relative.Racemose pillose and with a few hairs near the base, glands
inflorescences are uncommonin thiscomplex,but present at the base of the inner 3 stamens,stami-
I do notthinkthat0. corrugatais closelyrelated nodia not seen, receptaclecup-shaped,pubescent
to the otherspecies withracemoseinflorescences inside.Fruitsand cupulesnotknown.FlowersFeb-
such as 0. purpureaand 0. gordonii.These species ruary-April.
differgreatlyfromeach otherin othercharacters, Ocotea gordoniiis part of the Ocotea helicteri-
and racemoseinflorescences have probablyarisen folia complexbecause of the papillose innersur-
several timesin this complexby reductionof the face of the tepals,the relativelylarge flowers, the
more commonpaniculate-cymoseinflorescence. long spreadingindumentof stemsand leaves, and
This species was describedas abundantat thetype the spreadingtepals. Withinthis groupit stands
576 Novon

'i
?

X -i

?: i ii !
-

!
?

Figures5-8 (clockwisefromtop left).-5. Ocoteagordonii:McPherson10421 (holotype).-6. Ocoteahelicterifolia:

MendezTon 5902. -8. Ocotea helicterifolia
-7. Ocotea helicterifolia:
Linden1641 (syntype). (Costa Rica): Herrera
4866.
Volume9, Number4 van derWerff 577
1999 Ocotea helicterifolia
Species Group

apartdue to its racemoseinflorescences, the long at lowerelevations(50-600 m) and has moreob-

pedicels,and pubescentreceptacle.Onlyone other ovate leaves. These collectionsfromCosta Rica
species in thisgroupwithracemoseinflorescences possiblyrepresent an undescribedtaxon.Provision-
is knownfromPanama, and this species, 0. pur- ally includedin 0. helicterifolia is Ocotea teneja-
purea,has pubescentflowersca. 5 mmin diameter pensis,knownonlyfromthe typecollection.This
and leaves less than11 cm long.Vegetatively,there collectiondiffersfromtypical0. helicterifolia in
is a strongerresemblancebetweenthenewspecies thatthe outer6 stamenshave filaments about 1/3
bothofwhich the lengthof the anthers,the lowerleaf surfaceis
and 0. valerianaand 0. helicterifolia,
occur in Costa Rica. These two species have pa- verysparselypubescent,and small axillarytuftsof
niculate-cymose inflorescencesand glabrousrecep- hairsare oftenpresenton the lowerleaf surface.I
tacles.Twootherspecies fromCosta Rica and Pan- have not foundothercollectionswiththese char-
ama, 0. lentiiand 0. valerioides,have pubescent acters,and therefore include 0. tenejapensisas a
receptacles like 0. gordonii,but in
differ theirpa- somewhat aberrant form in 0. helicterifolia.
niculate-cymose inflorescencesand obovateleaves. has a widerangeofdis-
Because 0. helicterifolia
This species is dedicatedto GordonMcPherson, tributionand shows some variationin vegetative
whoseexcellentcollectionshave contributed much characters, I includethreefigures, includingone of
to our knowledgeofthe Panamanianflora. the type,ofthisspecies.

Paratypes.PANAMA.Veraguas: mountains above 7. Ocotea

Sta.Fe, Hammel& Kress8534 (MO).Chiriqui:vicinity lentii W. C. Burger,FieldianaBot.,n.s.
ofFortunaDam,McPherson 10573(MO). 23: 86. 1990. TYPE: CostaRica. Cartago:Lent
794 (isotype,MO). Figure9.
6. Ocotea helicterifolia(Meisner)Hemsley,Biol. Ocotealentiiis best recognizedby its large(18-
Centr.Amer.,Bot. 3: 73. 1882. Oreodaphne 40 cm long),obovate,alternateleaves and thehir-
Meisner,Prodr.15(1): 123. 1864. sute indumenton twigsand inflorescences.
helicterifolia Partof
(Meisner)Mez,Jahrb.Ko- the surfaceremainsvisiblebetweenthehairs.Oco-
Phoebehelicterifolia
enigl. Bot. Gart.Berlin5: 193. 1889. Cinna- tea lentilis an infrequentlycollectedspeciesknown
momumhelicterifolium (Meisner)Kostermans, onlyfromtheprovincesCartagoand Guanacastein
Reinwardtia6: 21. 1961. TYPE: Mexico. Costa Rica. Most collectionshave been made be-
Chiapas: Linden1641 (syntype, K). Figures6, tween700 and 1400 m altitude.
7,8.
Nectandra corzoana
Lundell,Wrightia4: 102. 1969.Syn. 8. Ocotea mollicelia (Blake) van der Werff, Fiel-
nov.Phoebecorzoana (Lundell)Lundell,Wrightia5: diana Bot., n.s. 23: 88. 1990. Phoebe molli-
342. 1977.Cinnamomum corzoanum Kos-
(Lundell) cella Blake, Contr.GrayHerb. 52: 64. 1917.
termans, 10:422. 1988.TYPE:Mexico.
Reinwardtia Cinnamomummollicellum (Blake) Koster-
Chiapas:ShilomTon3560 (holotype, LL).
mans,Reinwardtia6: 22. 1961. TYPE: Costa
Ocoteatenejapensis
Lundell,Wrightia4: 108. 1969.Syn. Rica. Tonduz11676 (isotype,G). Figure10.
nov.TYPE: ShilomTon779 (holotype, LL).
Ocotea mollicellais characterizedby its race-
Phoebenectandroides
Mez,Jahrb. Koenigl.Bot.Gart.Ber- mose inflorescences, ascending(noterect)hairson
lin5: 194.1889.SYNTYPES:Botteri 1018notseen, the lowerleaf surface,and the graycolor of the
Galeotti7004(BR),Juergensen 937 notseen,Lieb-
21 notseen. indument.Its leaves are narrowly ellipticto ellip-
mann4 notseen,Oersted
tic-lanceolateand generallyless than 8 cm long.
Phoebeobtusata Contr.
Lundell, Univ.Mich.Herb.6: 21. Ocoteamollicellais a rarelycollectedspecies only
1941.Syn.nov.Cinnamomum obtusatum (Lundell) knownfromcloud forestsin Costa Rica and occur-
Kostermans, 6: 22. 1961.TYPE: Mex-
Reinwardtia
ico. Chiapas:Matuda1887 (isotype,CAS). ringbetween1400 and 2300 m altitude.

Ocoteahelicterifolia by its alter- 9. Ocotea patula van der Werff,

is characterized sp. nov.TYPE:
nate leaves, glabrousflowersand inside of the re- Costa Rica. Puntarenas:Cantonde Osa, Fila
ceptacle, as well as the hirsuteindumentof the Costena,Aguilaret al. 2715 (holotype,MO).
twigs,withat least a small partofthe surfacevis- Figure11.
ible betweenthe hairs. In Costa Rica it can be
confusedwith0. valeriana,whichhas a denserin- similis,sed floribus
Ocoteaeuvalerianae pubescentibus
brevioribusque
pedicellistomentellis recedit.
dumenton twigsand inflorescences. Ocotea helic-
terifoliaoccursfromsouthernMexicoto Panama at Small trees, to 8 m. Twigs tereteor slightly
altitudesof 1000-1900 m; in Costa Rica it occurs ridged,solid, denselyyellowishbrowntomentose
578 Novon

I~t~;l"c~??re.rt,,,,1
i
i

i~
i.~?-~
X

; gc2~Url?!"
rii
3f
-"b
~lrl+~

~t.:HI~.:::::?,:.fj??,:.:li.B:?Bl~l~~il . 10~ iBls aas~e~ ~

~":"i~
:f

~ ""a"'
M01(13~1$18rb
81 alrrl89ag
R EtlBs78S16

e ? r,

~ %9r;n

";"aY~r?ic~r?fbd;j~ ~~~;::

pl%'

:-i:ps~r

Figures9-12 (clockwisefromtop left).-9. Ocotea lentil:Herrera8938. -10. Ocotea mollicella:Tonduz11676

(isotype).-11. Ocoteapatula: Aguilar2715 (holotype).-12. Ocoteapraetermissa:
Burgeret al. 12065 (holotype).
Volume9, Number4 van derWerff 579
1999 Ocotea helicterifolia
Species Group

whenyoung,thesurfacecompletely coveredbythe ternate,rathersmall leaves (rarelyexceeding10

indument, theindument becomingwhitishwithage, cm), paniculate inflorescences,glabrous flowers
terminalbuds denselyyellowishbrowntomentose. witha pubescentinnersurfaceofthetepals,and a
Leaves 12-24 x 9-14 cm, broadlyelliptic,char- glabrousreceptacle.It is a species ofmontanefor-
taceous,alternate,the base obtuseto rounded,the ests from2000-3200 m elevation,mostlyfoundin
apex obtuseorshortly acuminate,theuppersurface Costa Rica, butwitha fewcollectionsfromPanama.
withsome erecthairs when young,soon glabres- This species has been misidentified as Ocotea (or
cent,themidriband lateralveinstomentellous, the Phoebe)pittieriand was includedin Burgerand van
lowersurfacesparselyto moderately pubescent,the der Werff (1990) underthatname.
hairserect,the indumentdenserand tomentellous
along the midriband lateralveins,midrib,lateral 11. Ocotea purpurea (Mez) van derWerff, comb.
veins,and tertiary venationimmersedon theupper nov. Basionym:PhoebepurpureaMez, Jahrb.
surface,raised to prominently raised on the lower Koenigl. Bot. Gart. Berlin 5: 196. 1889.
surface,domatialacking,lateralveins7-8; petioles TYPE: Guatemala. Alta Verapaz,von Tuer-
17-26 mm long, witha similarindumentas the kheim371 not seen (B not seen, typephoto-
twigs,shallowlycanaliculateon theuppersurface. graph,MO). Figure13.
Inflorescences 10-16 cm long,paniculate-cymose, 5: 33. 1974.
Nectandra capituliforma Lundell,Wrightia
denselyyellowishbrowntomentellous, in the axils Syn.nov.TYPE: Guatemala. Contreras11235(iso-
of cataphylls,rarelyin the axils ofleaves. Flowers type,MO).
ca. 7 mm diam., white,perfect,sparselyto mod-
Ocoteapurpureais characterized byitsracemose
eratelypubescent;receptacle densely pubescent
inflorescences (one or twocymesare rarelypresent
outside;pedicels ca. 2 mmlong,tomentellous. Te-
at thebase oftheinflorescence), pubescentflowers,
pals ca. 2.5 mmlong,elliptic,spreadingor some-
whatreflexedat anthesis,the innersurfacemod- and rather small (to 11 cm), (narrowly) elliptic,flat
leaves. The leaf and inflorescence charactersare
eratelyto sparselypapillose; stamens9, 4-celled,
visibleon the typephotograph. It is among
the outer6 ca. 1.2 mmlong,weaklypapillose,the clearly
thefewspecies in the0. helicterifolia complex with
cells arrangedin 2 pairs,introrse, a short(0.2 mm)
racemoseinflorescences and pubescentflowers; the
steriletip present,the antherssessile or nearlyso,
ones are an undescribed species from Mexico
inner3 stamensas longas theouter6, also weakly other
withlarge,obovateleaves and conspicuousdomatia
papillose, sessile or nearlyso, the cells in 2 pairs,
alongthe majorveins,0. corrugatawithbullateor
extrorse,glandspresentat thebase, staminodianot
and 0. mollicellawithascending,
seen. Pistil ca. 1.5 mmlong,the styleas long as rugose leaves,
gray pubescence on the lowerleaf surface(brown
the ovary,glabrous,receptacle cup-shaped,gla-
brousinside.Fruitsnotknown.FlowersDecember. and erect in 0. purpurea). Ocoteapurpureaoccurs
in southernMexico, Guatemala,Honduras,and
Ocoteapatula is named afterthe spreadingte- Panama at altitudesrangingfrom1400 to 2600 m.
pals at anthesis.It is knownonlyfromtwocollec- The name Phoebe bourgeauvianahas frequently
tions made on the same day in the same general been appliedtothisspecies,butthetypeofP. bour-
area, but probablyfromdifferent trees.One ofthe geauviana has glabrousflowersand a paniculate-
collectorsindicatedit was growingon a limestone cymoseinflorescence.
substrate.It is part of the 0. helicterifolia group
and resembles 0. valeriana. However,it differs 12. Ocotea tonii (Lundell)van der Werff, comb.
fromthisspecies in its denselytomentellous inflo- nov. Basionym: Nectandra tonii Lundell,
rescences(withthe surfaceentirelyor almosten- Wrightia4: 106. 1969. TYPE: Mexico.Chia-
tirelycoveredby the indument), by its shorter(2- pas: Municipiode Tenejapa, A. Shilom Ton
3 mm vs. 6-8 mm)and tomentellous pedicels, as 2014 (isotype,NY). Figure14.
well as its pubescentflowers. Ocotea toniiis characterizedby its narrowly el-
Paratype.COSTARICA.Puntarenas:Fila Costena, liptic,clustered leaves with short petioles, yellow-
Hammel etal. 19217(MO). ish browntomentum on the youngtwigs,and pa-
niculate-cymoseinflorescenceswith pubescent
10. Ocotea praetermissa van der Werff, Novon flowers.It is onlyknownfromChiapas at altitudes
between800 and 1500 m.
6: 482. 1996. TYPE: Costa Rica. Cartago:
Burgeret al. 12065 (holotype, MO). Figure12.
13. Ocotea valeriana (Standley)W. C. Burger,
Ocoteapraetermissa can be recognizedby its al- Fieldiana Bot.,n.s. 23: 96. 1990. Phoebeval-
580 Novon

u" t~r~al

~tr?~:rr

"a

i'ln?ii.i?:
r?-i*-?i*
*PI?-?,r

D,tllara~lnr?

mra~wr
Itag~ ~faE"9
~~IKPAbt~~WA

II a3311PP9 ? rJ~tsoacf

4,

ii

:oa~lns
~ rad~Pa

ifra.~a
~ri: sr?i-~
?
I
rzre~wr
xs~s~wa~
Qbl~X
1$WIAI~B
:figf

Figures13-16 (clockwisefromtop left).-13. Ocoteapurpurea:Breedlove9467. -14. Ocotea tonii:MendezTon

5264. -15. Ocoteavaleriana:Herreraet al. 475. -16. Ocoteavalerioides:Grayum3519.
Volume9, Number4 van derWerff 581
1999 Ocotea helicterifolia
Species Group

eriana Standley,Publ. Field Mus. Nat. Hist., are racemoseor have sec-
veins. Its inflorescences
Bot. Ser. 18(2): 460. 1937. Cinnamomum val- ondaryaxes terminating in a cyme.Flowersare pu-
erianum(Standley)Kostermans,Reinwardtia bescent.A descriptionofthisspecies is beingpre-
6: 24. 1961. TYPE: CostaRica. El Copey,Ton- pared by F. Lorea-Hernandez.The new species is
duz 11746 (holotype, F notseen). Figure15. onlyknownfromthe stateof Guerrero, Mexico.
Nectandraaustinii
Allen,J.ArnoldArbor.
26: 374. 1945.
Syn.nov.TYPE: CostaRica.AustinSmithP2226 IMPERFECTLYKNOWNSPECIES
A notseen).
(holotype,
Ocotea macrophylla Kunth,Nov. Gen. Sp. 2:
PhoebesmithiiAllen,J. ArnoldArbor.26: 317. 1945.
smithianum Kostermans,
Reinwardtia 131. 1818. Nectandra macrophylla(Kunth)
Cinnamomum
6: 23. 1961.TYPE: CostaRica.Austin SmithP.C. Nees, SystemaLaurinarum292. 1836. TYPE:
367 (holotype,F notseen). Colombia.Quindiu,Bonplands.n. (isotype,P).
OcoteavalerianaresemblesO. helicterifolia in its The typeof 0. macrophyllais a fruiting speci-
paniculate-cymose inflorescences,alternateleaves, men; thus,flowering specimenscan onlybe asso-
glabrousflowers, and glabrousinside oftherecep- ciated withthisname based on vegetativesimilar-
tacles. They are best separatedby the typeof in- ities. Rohwer (1991) recognized several recent
dumenton the youngtwigs.In O. valeriana this collectionsfromColombiaas O. macrophyllaand
indument consistsofshort,mattedhairsand longer, acceptedO. macrophylla as theonlyspecies in the
erect hairs; usually the short,mattedhairs cover groupoccurringsouthof Panama.
0. helicterifolia
the surfaceof the twigscompletely. The indument However,recent collections show that probably
of0. helicterifolia
consistsonlyoflong,erecthairs, morethanone species is presentin northern South
whichmaybe denselydistributed, butdo notcover America; lack of sufficient floweringcollections
the surfaceofthetwigscompletely. The cupules of makesit impossibleto accuratelydescribethetaxa
0. valerianaare also deeperthanthoseof O. hel- involved.All collectionshave alternateleaves and
icterifolia.Ocotea valeriana is not rare in Costa paniculate-cymose inflorescences.
Rica and is knownfroma veryfewcollectionsin I distinguish
Provisionally thefollowing entities:
Panama. It occurs from800 to 2200 m. I accept 1. Ocoteamacrophylla sensu typi.The typecol-
theconceptof0. valerianaused in Burgerand van lectionhas a ratherlax, hirsuteindumenton the
der Werff (1990), whereW. C. Burger,whohad ac- twigs,whichdoes notcompletely coverthesurface.
cess to the holotype,transferred Phoebevaleriana The hairson the lowerleaf surfaceare also rather
to Ocotea. AlthoughI have not seen the typesof long and straight.The inflorescence or infructes-
Nectandraaustiniiand Phoebesmithii,I place both cence is sparselyhirsute.In additionto thetype,I
species in synonymy of 0. valerianabased on the place hereSanchez1371 (MO) fromAntioquia,Co-
paratypesofthesespecies I have seen (forN. aus- lombia,and Ruiz Teran1558 (MO), Breteler4928
tinii:AustinSmithP2114 (GH); forP smithii:Aus- (MO), bothfromM&rida,Venezuela.The Ruiz Teran
tinSmithH523 (MO)). specimenis a fruiting one, whilethe Sanchez and
Bretelercollectionshave flowers. These flowers are
14. Ocotea valerioides W. C. Burger,Fieldiana glabrous,even on the inside ofthe tepals.Anthers
Bot., n.s. 23: 97. 1990. TYPE: Costa Rica. of the outersix stamensare sessile or nearlyso.
Hartshorn 1530 (isotype,MO). Figure16. 2. Sanchez1891 (MO) fromAntioquia,Colombia.
This species is similarto O. lentiiin its large, Indumenton the twigsis denselytomentellous/to-
the surfacecompletely, as is the
obovateleaves, paniculate-cymose inflorescences, mentose,covering
indument on theinflorescences.The flowers are pu-
and pubescentinside of the receptacles.Ocotea
bescent.
valerioidesdiffersin itsdenserindument,complete-
3. A groupof several specimenscharacterized
ly coveringthe surface,on the twigsand inflores-
cences. It is knownfroma fewcollectionsin Costa by a dense, tomentellous indumenton the twigs,
Rica and Panama,whereit occursin lowlandrain- covering the surfacecompletely. Inflorescences are
forestbetween50 and 500 m altitude. sparsely pubescent,and the flowersare glabrous or
nearlyso. This groupoccursin Colombia(Boyaca:
Lawrance217 & 281 (MO); Antioquia:Fernandez
15. Ocotea sp. nov.
7 (MO) and Giraldo287 (MO); Valle: Croat70677
This species can be readilyrecognizedbyitsob- (MO) and Albert2451 (MO)) and Ecuador (Carchi:
ovateleaves withconspicuoustuftsofwhitishhairs Palacios 12794 (MO), and van der Werff 10772
in theaxils ofthelateralveinsand alongthemajor (MO)).
582 Novon

OCOTEA SPECIES, POSSIBLY UNDESCRIBED Beaman5402 (2); Bello 640 (13), 1977 (13), 4316 (13),
4340 (13); Bourgeau 2234 (2); Brant2871 (2), 2886
A few collections fromPuebla (Mexico) resemble (11); Breedlove6307 (12), 6969 (11), 9467 (11), 15672
Ocoteahelicterifolia but differ
vegetatively, in hav- (3), 21696 (6), 23329 (6), 24742 (6), 25130 (6), 27588
(6), 27637 (11), 28182 (11), 31020 (6), 31371 (6), 32107
ing rathersmall flowers(only5-6 mmdiam.) and (6), 32239 (6), 32668 (6), 32699 (6), 33017 (3), 33543
in havingstamenswithdistinctfilaments (filaments (11), 34800 (6), 35276 (6), 38043 (2), 38781 (6), 42794
1/3as longas theanthers).I have notseen fruiting (11), 44395 (6), 44411 (6), 47853 (6), 48210 (6), 49332
materialofthistaxon.These collections(Rzedowski (6), 49348 (6), 49371 (6), 49746 (6), 50583 (2), 51313
31863, Tenorio8686, Ventura783, and Campos (6), 52589 (3), 52683 (6), 52998 (6), 53010 (6), 55689
(6), 55690 (6), 57479 (6), 58528 (6), 60245 (6), 60289
225, all in MO) maywell representan undescribed (6), 66260 (6), 68637 (6), 68935 (11), 70059 (12), 72693
species, but I preferto waituntilmorecollections (11); Burger11747 (6), 12065 (10), 12097 (13).
are at hand beforedescribingit.
Cascante 644 (13); Chavarria1153 (13); Campos853
Acknowledgments. I thankthecuratorsofCAS, (6), 954 (6), 1020 (6), 1328 (6), 1877 (6), 2569 (6), 3012
F, GH, K, and TEX forthe loan of specimens,B. (6), 3175 (6); Cedillo 667 (1), 1197 (1), 1204 (1), 1257
(1), 3060 (aff.2), 3323 (aff.2), 3406 (aff.2); Chacon1557
Stannard(K) forhis efforts in locatingthe typeof
(13), 1634 (14); Choc 89 (6); Churchill4326 (6), 5926
and T. Wendt(TEX) forduplicates
0. helicterifolia, (6); Contreras5209 (11), 11186 (2), 11205 (2), 11233
of his excellentLauraceae collections. (11), 11235 (11), 11318 (11); Croat46913 (13), 47613
(6), 64215 (6).
Cited
Literature
Davidse 28547 (10), 28551 (10), 28627 (10), 28809
Allen,C. K. 1945. Studiesin theLauraceaeVI. A prelim- (10), 28922 (10), 30477 (6), 34826 (11); Dryer1336 (13).
inarysurveyoftheMexicanand CentralAmericanspe-
cies. J. ArnoldArbor.26: 280-434. Estrada515 (14); Evans 1474 (11).
Burger,C. W. & H. van der Werff.1990. Flora Costari-
Galeotti7004 (6); Gentry13575 (6), 44065 (6); Gereau
censis,Lauraceae. Fieldiana Bot.,n.s. 23: 1-129.
Rohwer,J. G. 1991. Borderlinecases betweenOcotea, 2096 (1); Gomez 18814 (7), 23619 (14); Gonzalez 159
(14); Grayum3519 (14), 8335 (6), 8870 (14); Grijalva
Nectandra,and Phoebe(Lauraceae):The marginalspe- 3705 (6).
cies oftheO. helicterifolia
group,includingtheO. hey-
deana group.Bot.Jahrb.Syst.112(3): 365-397. Haber 314 (13), 5064 (13), 5591 (13), 5665 (13), 5701
(13), 5826 (13), 5998 (13), 6643 (13), 7938 (13), 8198
(13), 9155 (13), 11267 (13); von Hagen 2022 (11), 2031
LIST OF SPECIES RECOGNIZED (11); von Hagen 2070 (11); Hammel7013 (10), 8534 (5),
15679 (6), 16504 (6 vel aff.),17836 (14), 18049 (13),
1. Ocotea betazensis (Mez) van der Werff 19217 (9), 20062 (6), 20553 (8); Hartshorn1463 (13),
2. Ocotea bourgeauviana (Mez) van der Werff 1530 (14), 2130 (13); Hawkins452 (11), 588A (11), 647
3. Ocotea congregata van der Werff (11), 712 (11); Hazlett 582 (11), 1341 (6), 2516 (11);
4. Ocotea corrugata van der Werff Heath 594 (6), 2033 (6); Herrera475 (13), 3778 (13),
5. Ocotea gordonii van der Werff 4866 (6), 4946 (13), 4986 (14), 5144 (14), 8938 (7);
6. Ocotea helicterifolia(Meisner) Hemsley House 975 (11), 1187 (11), 1300 (11).
7. Ocotea lentii W. C. Burger
8. Ocotea mollicella (Blake) van der Werff Ishiki 1389 (11), 1396 (11), 1488 (11), 1551 (11).
9. Ocotea patula van der Werff
10. Ocotea praetermissa van der Werff 578 (6); Juzepczuk
Jimenez564 (13), 827 (13); Johnson
11. Ocotea purpurea (Mez) van der Werff 1513 (12).
12. Ocoteatonii(Lundell)van der Werff
13. Ocoteavaleriana(Standley)W. C. Burger Kirkbride486 (14).
14. Ocotea valerioides W. C. Burger
Laughlin528 (6); Lent 794 (7), 2070 (7); Liebmann
770/3 (1), 770/22 (1), 771/2 (1), 771/23 (1); Liesner
SPECIMENS STUDIED 1789 (6); Linden 1641 (6); Lopez L. 181 (11); Lorea
5542 (6); Lorence4711 (1); Lot 222 (6); Lundell 18944
If collected by a team, only the last name of the (11), 19425 (6), 20435 (11), 20999 (11), 21170 (11).
firstcollector is listed. For example, Breedlove &
Smith is included as Breedlove. Type collections MacDougall H288 (6); MartinezS. 17084 (3), 20768
are in bold face. The number in parentheses fol- (6); Matuda 1887 (6), 5020 (11), 5125 (6), 5352 (11),
5375 (11), 5400 (6), 5437 (11), 6095 (6), 16224 (6); Maya
lowing the collector refers to the species listed 1226 (6), 1316 (6), 2094 (6), 2865 (6), 2882 (6), 2989
above, forexample, Aguilar 438 is species #6 from (6), 3081 (6), 3098 (11), 3951 (6), 3967 (6), 4176 (11),
the list (= 0. helicterifolia). 4232 (11), 4276 (6), 4292 (11); McPherson9259 (13),
10421 (5), 10573 (5); Meave 1314 (6); Mejia 12 (11),
Aguilar438 (6), 2715 (9), 2819 (6); Allen 15209 (6); 256 (11), 283 (11), 294 (11), 334 (11), 379 (11); Mendez
Almeda3762 (10); Angulo200 (10); Antonio1602 (10); G. 7966 (6), 8503 (3), 8888 (6), 8930 (3); MendezTon
Aranda1247 (11), 1261 (11), 1265 (11), 1317 (11); Aus- 4214 (2), 4802 (11), 5035 (12), 5068 (11), 5264 (12),
tinSmithH523 (13), H679 (10), P2114 (13). 5902 (6), 5931 (11), 9594 (3); Miller2653 (6), 2943 (6
Volume9, Number4 van derWerff 583
1999 Ocotea helicterifolia
Species Group

s.1.);Molina203 (6), 8138 (6), 8186 (6), 11521 (6), 11769 316 (6); Standley
Soto 226 (6); Sousa 9337 (1); Stafford
(6), 12289 (6), 23320 (11), 24212 (11), 24289 (11), 31102 8044 (6), 20611 (6), 69379 (6), 69525 (6), 71446 (6),
(6); Montenegro 1845 (6); Mora110 (14), 416 (7), 508 (7); 89985 (6), 91546 (6); Stevens 17055 (6), 20397 (6),
Moraga171 (7); Morales39 (13), 248 (13), 377 (6), 436 22153 (6); Steyermark 44207 (6), 44280 (6), 44648 (6).
(10), 665 (6), 1053 (6), 4806 (10), 5812 (8), 5989 (8);
Moreno7477 (6), 8253 (6), 15427 (6), 15647 (6), 15826 Taylor4460 (10); Tenorio3457 (12 vel aff.),5870 (1),
(6), 15862 (6), 19376 (6), 19396B (6), 19424 (6), 21103 11276 (1); Thorne40325 (6); Tonduz 11676 (8); Torres
(6). 476 (1), 494 (1), 1400 (6), 2066 (1), 2928 (1), 4832 (11),
4871 (11), 4953 (1), 4973 (1), 6598 (6), 8866 (6), 9153
Nee 24906 (6), 29832 (aff.2); Nelson2419 (6). (6), 10444 (6), 10830 (6), 11620 (6), 11911 (6); von
Turckheim II 1651(11), II 2164 (6).
Poveda 3920 (14).
Vazquez Torres378 (2).
Quesada 368 (6), 941 (7); Quintanilla49 (3). Wendt2677 (aff.2), 3000 (aff.2), 3000A (aff.2), 3861
(2), 4315 (6), 4653 (11), 4866 (11), 5137 (6), 5622 (2),
Ramirez139 (6), 384 (6), 452 (6); Ramirez-Marcial 656 209 (13); Williams13888 (10),
6765 (4); Wheelwright
(6); Raven 19804 (6), 21630 (6); Rios 179 (7); Rivera260 15760 (6), 16469 (13), 20540 (6), 28623 (13), 28996 (13),
(10), 664 (6). 40390 (6); Wilson40770 (6), 40859 (6).
Sandino2330 (6), 4706 (6); SantizRuiz836 (6); Schul- Yncker5777 (11).
tes 827 (2); Shank55 (6); Shilom Ton 779 (6), 2014
(12), 3645 (12), 3560 (6), 7399 (11); Sinaca 777 (aff.2); Zamora674 (14), 1287 (6); Zuniga282 (6).