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Two New Species in the Tillandsia utriculata Complex(Bromeliaceae) from MexicoAuthor(s) Ivoacuten M Ramiacuterez M and Germaacuten Carnevali F CSource Novon A Journal for Botanical Nomenclature 17(1)72-78 2007Published By Missouri Botanical GardenDOI httpdxdoiorg1034171055-3177(2007)17[72TNSITT]20CO2URL httpwwwbiooneorgdoifull1034171055-317728200729175B723ATNSITT5D20CO3B2

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Two New Species in the Tillandsia utriculata Complex (Bromeliaceae)from Mexico

Ivon M Ramırez M and German Carnevali F C

Herbarium CICY Centro de Investigacion Cientıfica de Yucatan AC Calle 43 130 ColoniaChuburna de Hidalgo Merida 97200 Yucatan Mexico ramirezcicymx carnevalcicymx

ABSTRACT Two new species in the Tillandsiautriculata (L) L complex (Bromeliaceae) from Mexicoare proposed Both are herein described and illus-trated with their affinities discussed The first newentity T aesii I Ramırez amp Carnevali from the statesof Jalisco Guerrero and Oaxaca is similar to Tmakoyana Baker but the new taxon has an openrosette with leaves that are green adaxially and whitelepidote abaxially (vs a funnelform rosette with grayleaves on both surfaces in T makoyana) actinomor-phic flowers with a light apple-green corolla (vszygomorphic with a purple to light purple corolla) Thesecond taxon proposed here T pinicola I Ramırez ampCarnevali from the state of Oaxaca is also similar toT makoyana but is a smaller plant with a morecompact growth habit and leaves with proportionallyshorter leaf blades that abruptly attenuate from a broadsheath (vs gradually attenuate into the proportionallylonger blade) into a sub-acicular apex It is alsocharacterized by typically growing on pines hence theepithet

RESUMEN Dos nuevas especies en el complejoTillandsia utriculata (L) L (Bromeliaceae) de Mexicose proponen como nuevas Ambas se describen sediscuten sus afinidades y se documentan conilustraciones y fotografıas La primera especie T aesiiI Ramırez amp Carnevali de los estados de JaliscoGuerrerro y Oaxaca es similar a T makoyana Bakerpero la nueva especie tiene rosetas abiertas con hojasverdes adaxialmente y blanco lepidotas abaxialmente(vs rosetas en forma de embudo con hojas grises enambas superficies en T makoyana) flores actinomor-ficas con corola verde manzana claro (vs floreszigomorfas y con petalos purpura o purpura claro) Lasegunda especie T pinicola I Ramırez amp Carnevalidel estado de Oaxaca es tambien similar a Tmakoyana pero es una planta mas pequena con unhabito de crecimiento mas compacto y hojas propor-cionalmente con laminas foliares mas cortas las cualesse atenuan abruptamente en una vaina ancha (vsvainas atenuandose gradualmente en laminas foliaresproporcionalmente mas largas) con un apice acicularEsta nueva especie tambien se caracteriza por creceren pinos de allı su epıteto

Key words Bromeliaceae Mexico Tillandsia

The genus Tillandsia L was classified in sevensubgenera by Mez (1896) and later supported bySmith and Downs (1977) in their Flora Neotropica

treatment of the entire Bromeliaceae The high levelsof homoplasy in several morphological characters

(such as petal appendages) considered important intraditional classifications (including that of Smith and

Downs) have blurred our understanding of the limits ofthe supraspecific groups within the genus Some of thesubgenera recognized by Mez (1896) and Smith and

Downs (1977) have been recently treated as distinctgenera ie Racinaea M A Spencer amp L B Smith

formerly Tillandsia subg Pseudocatopsis Baker (Spen-cer amp Smith 1993) but without the necessary

phylogenetic work to back up the proposals despitetheir wide acceptance by several taxonomists (egLuther 1995 Morales 2003)

Recent novel research and the analysis of newcharacters (ranging from the re-evaluation of fresh

floral material to newly generated DNA nucleotidesequence data) have allowed a better understanding ofthe relationships within the genus (Gardner 1986

Terry et al 1997 Barfuss et al 2005) but morestudies are required before more taxonomic and

nomenclatural changes are warranted within theTillandsioideae Furthermore in the most recentphylogenetic study based on DNA sequence data

from seven plastid regions none of the subgenera ofTillandsia were found to be monophyletic (Barfuss et

al 2005)

Studies at the population and at the species-complex levels using field data (eg habitat and

distribution analysis ecology) and the careful analysisof fresh material either in the field or under

cultivation can provide a wealth of new data that areusually unavailable when only working with herbar-

ium material (see Gardner 1986 for a good example)Within Tillandsia some species complexes includingthe relatives of T utriculata (L) L can only be

understood with this kind of approach because most ofthe relevant features required to diagnose the taxa in

the group become lost upon vouchering including

NOVON 17 72ndash78 PUBLISHED ON 23 APRIL 2007

plant architecture In many cases the taxa thatcompose these species complexes are allopatric orparapatric but because herbarium data are scarcethey fail to reveal the often subtle ecological andgeographical patterns which can only be disclosedwith extensive fieldwork and an abundant herbariumrecord

A brief description of the Tillandsia utriculatacomplex was provided by Ramırez et al (2004)Described species in the T utriculata complexinclude T makoyana Baker T limbata Schlechten-dal T dasyliriifolia Baker T pringlei S Watson(treated by Gardner (1984) as Tillandsia utriculatasubsp pringlei (S Watson) C S Gardner) Tutriculata T calcicola L B Smith amp Proctor Tcucaensis Wittman T pulvinata Baker T geniculataBaker and a few other related species some of whichare probably still undescribed A few additional taxasuch as T flexuosa Swartz T karwinskyana Schultesf and T albida Mez amp Purpus have inflorescencesand flowers that recall those of this complex but theplants are strikingly different Phylogenetic analysesare required to better ascertain the position of thesetaxa Tillandsia utriculata is the type species of thegenus

As a result of extensive fieldwork in Mexico as wellas cultivation of plants of the group for several yearsthe delimitation of taxa in the Tillandsia utriculatacomplex in Mexican populations is now betterunderstood Species whose identities and circum-scriptions have come into focus with the study of thiscomplex include T dasyliriifolia (Ramırez et al2004) T limbata Schlechtendal (Espejo et al 2005)and T makoyana The type of the latter species isa watercolor painting by Edouard Morren housed inthe Kew library without collection data the name hasbeen applied in herbaria and in the literature (egSmith amp Downs 1977) to several species of the Tutriculata complex that display purple corollas Ourinterpretation of Morrenrsquos plate is that it is conspecificwith a species that commonly features a rosette ofstraight erect (not twisted nor spreading) gray leavesand purple or light purple corollas that are slightlyzygomorphic Tillandsia makoyana has been collectedat varying elevations ranging from sea level to ca2000 m in central Mexico and is variable in plant sizeand inflorescence length However in most individ-uals of the species the peduncle of the inflorescence isabout as long to slightly longer than the leaves

As a result of this newly gained understanding ofrelevant species in the Tillandsia utriculata complexparticularly the identity of T makoyana during thecourse of this study two distinctive new entitiesbecame apparent The new taxa herein described hadbeen previously collected but had until now remained

misidentified in herbaria variously as T limbata Tmakoyana andor T dasyliriifolia

Tillandsia aesii I Ramırez amp Carnevali sp novTYPE Mexico Jalisco mpio La Huertaalrededores de la Est Biol Chamela ca19u309N 105u039W 0ndash10 m 12 Mar 2002 GCarnevali amp I M Ramırez 6973 (holotype CICY[3]) Figure 1

Haec species Tillandsiae makoyanae affinis est sed rosulaaperta superficie adaxiali foliorum viride non cinereainflorescentia breviore spicis paucioribus ceraciter vestitisfloribus actinomorphis cylindricis cum sepalis petalisque etstigma viridibus differt

Epiphytic herb 62ndash100 cm high with 1 to 4 basalrosettes on the flowering or fruiting plant or the plants(more commonly) monocarpic rosette with 12 to 18leaves when flowering with open aspect slightlyfunnelform when young foliar blades slightly tostrongly twisted (curved outward and pointing forward)spreading to reflexed foliar sheaths barely impound-ing Foliar sheaths of external leaves almost flat thoseof internal leaves forming a funnelform structurestraight and erect white-lepidote externally somewhatconcave foliar blades widely triangular at baseattenuate and acute long-caudate coriaceous ratherstiff conspicuously nerved abaxially 30ndash32 6 25ndash4 cm light green adaxially white-lepidote abaxiallymargins slightly curving upward into a canaliculatestructure Inflorescence pyramidal a panicle 32ndash39 cm with up to 11 spikes the new inflorescencedeveloping very slowly and taking up to 3 monthsbefore the first flowers open successively floweringbasipetally for several weeks peduncle shorter or aslong as the leaves pink glaucous 21ndash40 cm ca 1 cmdiam covered by tubular acute imbricate bractsinternodes 2ndash4 cm long bracts of the peduncle almostcovering the internodes green and glaucous 35ndash456 2ndash26 cm (longer and wider toward the base)widely triangular acute margins hyaline conspicu-ously nerved when dry slightly surpassing the inter-nodes inflorescence rachis red or pinkish flesh-colored05 cm diam naked flexuous erect spikes 8 to 11rachis green almost obscured by floral bractsdorsoventrally flattened with flowers in 2 rows bractsof the spikes widely triangular obtuse 23 6 14 cmentire slightly pink white-lepidote floral bractsnarrowly triangular nerved acute margins hyalinebarely covering the rachis 14 6 08ndash09 cm greenshorter than the sepals white lepidote adaxiallyFlowers 5ndash65 cm actinomorphic erect 1 to 4 opendaily per inflorescence (1 at a time per spike) eachflower lasting 1 day protandrous odorless basallygeniculate sepals narrowly triangular to wide-elliptic

Volume 17 Number 1 Ramırez amp Carnevali 732007 New Species in the Tillandsia utriculata Complex

Fig

ure

1

Til

lan

dsi

aa

esii

IR

amır

ezamp

Car

nev

ali

mdashA

Hab

itS

cale

bar

51

0cm

mdashB

Upp

erro

wf

rom

left

tori

ght

flow

erf

low

erw

ith

flor

alb

ract

rem

oved

flo

wer

wit

hou

tfl

oral

bra

cts

epal

san

dp

etal

ssh

owin

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ary

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est

igm

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ents

an

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ther

sB

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wfr

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oral

bra

ctth

ree

sep

als

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thre

ep

etal

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bar

54

cmB

ased

onth

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ype

GC

arn

eva

liamp

IM

R

am

ırez

69

73

(CIC

Y)

Mar

gin

sof

sep

als

and

flor

alb

ract

sar

eso

mew

hat

revo

lute

inth

efi

gure

s

74 Novon

obtusely acute 18ndash25 6 07ndash13 cm 2 sepalsbasally connate by 6ndash15 mm apple green slightlydarker and shinier than petals petals narrowly oblongapically rounded 32ndash38 6 04ndash08 cm slightlyoblique constricted ca 15 cm distally from base palegreen to cream-green basally white stamens exsertedin 2 series filaments concolorous with petals plicatelighter at the base 5ndash55 cm anthers brown 8 mmdorsifixed pollen yellow ovary 15 mm pyramidalnectar sweet abundant 31u Brix (total concentration ofsugars) stigma exserted slightly protruding above theanthers at anthesis lobes conduplicate somewhatspiraled 47ndash49 cm basally light green apicallyapple green Fruits and seeds not observed

Habitat distribution and phenology Tillandsiaaesii is found in low dry caducifolious forestsassociated with Cnidoscolus Pohl Plumeria L Pseu-dosmodingium Engler Agave L Opuntia (L) MillerPedilanthus Necker ex Poiteau Cordia elaeagnoidesDC Bursera simaruba Sargent xeromorphic epiphytessuch as Tillandsia ionantha Planchon T intermediaMez T caput-medusae E Morren T concolor L BSmith Encyclia adenocarpon (La Llave amp Lexarza)Schlechter Myrmecophila galeottiana (Reichenbach f)Rolfe Cactaceae such as Pachycereus (A Berger)Britton amp Rose and thorny leguminous trees andshrubs It has been collected at 0ndash600 m of elevationon the Pacific coast of Mexico Flowering specimenswere collected from March through July and fruit-ing specimens were collected from May throughSeptember

Eponymy The name is based on an acronym of theinitials of Dr Adolfo Espejo Serna a Mexican botanistassociated with herbarium UAMIZ of the UniversidadAutonoma Metropolitana in Mexico DF whose workin Bromeliaceae and other monocots has contributedgreatly to the knowledge of the Mexican flora

Tillandsia aesii is phenetically similar and proba-bly phylogenetically closest to T makoyana a speciesthat commonly features a rosette of straight erect (nottwisted and spreading) leaves Tillandsia makoyanafurthermore displays purple or light purple flowersand a slightly zygomorphic corolla as opposed to theactinomorphic green tubular corolla of T aesii Bothspecies have been collected at varying elevationsranging from sea level to ca 2000 m but T aesii hasonly been collected at elevations from sea level to ca600 m and in dryer environments

At least three other vegetatively similar taxa in theTillandsia utriculata complex display pale flowers Thewidespread T utriculata features longer thinner (easyto bend) peduncles and branches and the smaller whitecorollas are gibbous or somewhat inflated with thepetals convergent at apex Tillandsia dasyliriifolia and

T limbata are easily distinguished from T aesii bytheir inflorescences with much longer pedunclesbearing flowers with white slightly zygomorphiccorollas Tillandsia aesii is further distinguished fromthese two taxa by the proportionally smaller rosette ofstiff leaves that are slightly twisted In addition Tdasyliriifolia and T limbata are restricted to the humidlowlands of the Gulf Coast of Mexico (T limbata) or tothe Yucatan Peninsula (T dasyliriifolia)

Paratypes MEXICO Guerrero mpio Huitzuco Ama-titlan 12 km SE del Huitzuco 4 June 1985 J C Soto Nunezamp Aureoles C 8786 (MEXU) Jalisco mpio La Huertaalred Est Biol Chamela 3 Nov 1997 G Carnevali amp GSalazar 4767 (CICY) camino por la base SE del Cerro de laTambora 16 Jan 1985 P Magana Rueda amp E Lott 10(MEXU) Chamela Field Stat 100 km S of Puerto Vallarta26 Sep 1988 A Gentry 63981 (MO) Oaxaca mpio MatıasRomero carr entre La Ventosa y el entronque a Juchitan10 km N de La Ventosa 4 May 1997 G Carnevali GCampos M Gomez amp F May 4438 (CICY) mpio AsuncionTlacolulita dist Yautepec ca 9 km despues de Reformasobre camino a TehuantepecndashOaxaca 2 Sep 2002 IRamırez A Espejo A Lopez J Ceja y A Mendoza 998(CICY) mpio Asuncion-Ixtaltepec distrito Juchitan 1 kmdespues de Santiago de Ixtaltepec 2 Sep 2002 I RamırezA Espejo A Lopez J Ceja y A Mendoza 979 (CICY) 28 kmNW of La Ventosa along Trans-Isthmian hwy (rte 185) 17July 1958 R M King 630 (MICH TEX US) flat grazed area55 km NE of Juchitan along Pan-Amer hwy (rte 190) 2July 1958 R M King 360 (MEXU MICH TEX US) flatgrazed area 4 km N-NE of Tehuantepec along the Trans-Isthmian hwy (rte 185 amp 190) 5 July 1959 R M King 1352(LL) flat grazed area 16 km N-NE of Tehuantepec 7 July1959 R M King 1389 (TEX) mtns along rte 190 ca74 miles SE of Oaxaca 22 June 1960 R M King 2975(TEX US) Tehuantepec dry open savannah 2 Apr 1957M B Foster amp O C Van Hyning 2939 (US [2]) on rte 1856 km N of Union Hidalgo 10 July 1958 W G Williams Jr ampD Francoeur 21 (MICH) on rte 185 15 km N of UnionHidalgo at edge of steep drop into a small valley 11 July1958 Walter G Williams Jr amp Don Francoeur 24 (MICH)mpio Nejapa de Madero distrito Yautepec 24 km al N-NEde Rıo Hondo brecha a Asuncion Lachixonase 16 May1995 A Salinas T amp E Martınez-Correa 8155 (MEXU)

Tillandsia pinicola I Ramırez amp Carnevali sp novTYPE Mexico Yucatan Merida floreciendo encultivo 13 Jan 2005 I Ramırez amp G Carnevali1274 (holotype CICY [2] isotype MO) [Oaxacampio Santa Marıa Chimalapa Dist Juchitan ca20 km despues de el Mezquite 7 km despues deLazaro Cardenas rumbo a Santa Marıa Chima-lapa 16u469100N 94u509380W ca 385 mcolectada originalmente por I Ramırez AEspejo A Lopez-Ferrari A Mendoza amp J Cejael 2 Sep 2002] Figure 2

Haec species Tillandsiae makoyanae affinis est sed plantaambito parviore foliorum vaginis latissimis foliorum laminisproportione brevioribus apice acicularibus abrupte attenuatis(vs paulo apicaliter attenuatis) et habitatio pinicola abhorret

Volume 17 Number 1 Ramırez amp Carnevali 752007 New Species in the Tillandsia utriculata Complex

Fig

ure

2

Til

lan

dsia

pin

icol

aI

Ram

ırez

ampC

arn

eval

imdash

A

Hab

it

Sca

leb

ar5

10

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B

Up

per

row

fr

omle

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righ

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ower

fl

ower

wit

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mov

ed

flow

erw

ith

out

flor

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ract

se

pal

san

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ary

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igm

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ents

an

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otto

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wfr

omle

ftto

righ

tfl

oral

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ct

two

sep

als

and

two

pet

als

Sca

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ar5

3cm

B

ased

onth

eh

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ype

IR

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ırez

ampG

C

arn

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27

4(C

ICY

)M

argi

ns

ofse

pal

san

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oral

bra

cts

are

som

ewh

atre

volu

tein

the

figu

res

76 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

plant architecture In many cases the taxa thatcompose these species complexes are allopatric orparapatric but because herbarium data are scarcethey fail to reveal the often subtle ecological andgeographical patterns which can only be disclosedwith extensive fieldwork and an abundant herbariumrecord

A brief description of the Tillandsia utriculatacomplex was provided by Ramırez et al (2004)Described species in the T utriculata complexinclude T makoyana Baker T limbata Schlechten-dal T dasyliriifolia Baker T pringlei S Watson(treated by Gardner (1984) as Tillandsia utriculatasubsp pringlei (S Watson) C S Gardner) Tutriculata T calcicola L B Smith amp Proctor Tcucaensis Wittman T pulvinata Baker T geniculataBaker and a few other related species some of whichare probably still undescribed A few additional taxasuch as T flexuosa Swartz T karwinskyana Schultesf and T albida Mez amp Purpus have inflorescencesand flowers that recall those of this complex but theplants are strikingly different Phylogenetic analysesare required to better ascertain the position of thesetaxa Tillandsia utriculata is the type species of thegenus

As a result of extensive fieldwork in Mexico as wellas cultivation of plants of the group for several yearsthe delimitation of taxa in the Tillandsia utriculatacomplex in Mexican populations is now betterunderstood Species whose identities and circum-scriptions have come into focus with the study of thiscomplex include T dasyliriifolia (Ramırez et al2004) T limbata Schlechtendal (Espejo et al 2005)and T makoyana The type of the latter species isa watercolor painting by Edouard Morren housed inthe Kew library without collection data the name hasbeen applied in herbaria and in the literature (egSmith amp Downs 1977) to several species of the Tutriculata complex that display purple corollas Ourinterpretation of Morrenrsquos plate is that it is conspecificwith a species that commonly features a rosette ofstraight erect (not twisted nor spreading) gray leavesand purple or light purple corollas that are slightlyzygomorphic Tillandsia makoyana has been collectedat varying elevations ranging from sea level to ca2000 m in central Mexico and is variable in plant sizeand inflorescence length However in most individ-uals of the species the peduncle of the inflorescence isabout as long to slightly longer than the leaves

As a result of this newly gained understanding ofrelevant species in the Tillandsia utriculata complexparticularly the identity of T makoyana during thecourse of this study two distinctive new entitiesbecame apparent The new taxa herein described hadbeen previously collected but had until now remained

misidentified in herbaria variously as T limbata Tmakoyana andor T dasyliriifolia

Tillandsia aesii I Ramırez amp Carnevali sp novTYPE Mexico Jalisco mpio La Huertaalrededores de la Est Biol Chamela ca19u309N 105u039W 0ndash10 m 12 Mar 2002 GCarnevali amp I M Ramırez 6973 (holotype CICY[3]) Figure 1

Haec species Tillandsiae makoyanae affinis est sed rosulaaperta superficie adaxiali foliorum viride non cinereainflorescentia breviore spicis paucioribus ceraciter vestitisfloribus actinomorphis cylindricis cum sepalis petalisque etstigma viridibus differt

Epiphytic herb 62ndash100 cm high with 1 to 4 basalrosettes on the flowering or fruiting plant or the plants(more commonly) monocarpic rosette with 12 to 18leaves when flowering with open aspect slightlyfunnelform when young foliar blades slightly tostrongly twisted (curved outward and pointing forward)spreading to reflexed foliar sheaths barely impound-ing Foliar sheaths of external leaves almost flat thoseof internal leaves forming a funnelform structurestraight and erect white-lepidote externally somewhatconcave foliar blades widely triangular at baseattenuate and acute long-caudate coriaceous ratherstiff conspicuously nerved abaxially 30ndash32 6 25ndash4 cm light green adaxially white-lepidote abaxiallymargins slightly curving upward into a canaliculatestructure Inflorescence pyramidal a panicle 32ndash39 cm with up to 11 spikes the new inflorescencedeveloping very slowly and taking up to 3 monthsbefore the first flowers open successively floweringbasipetally for several weeks peduncle shorter or aslong as the leaves pink glaucous 21ndash40 cm ca 1 cmdiam covered by tubular acute imbricate bractsinternodes 2ndash4 cm long bracts of the peduncle almostcovering the internodes green and glaucous 35ndash456 2ndash26 cm (longer and wider toward the base)widely triangular acute margins hyaline conspicu-ously nerved when dry slightly surpassing the inter-nodes inflorescence rachis red or pinkish flesh-colored05 cm diam naked flexuous erect spikes 8 to 11rachis green almost obscured by floral bractsdorsoventrally flattened with flowers in 2 rows bractsof the spikes widely triangular obtuse 23 6 14 cmentire slightly pink white-lepidote floral bractsnarrowly triangular nerved acute margins hyalinebarely covering the rachis 14 6 08ndash09 cm greenshorter than the sepals white lepidote adaxiallyFlowers 5ndash65 cm actinomorphic erect 1 to 4 opendaily per inflorescence (1 at a time per spike) eachflower lasting 1 day protandrous odorless basallygeniculate sepals narrowly triangular to wide-elliptic

Volume 17 Number 1 Ramırez amp Carnevali 732007 New Species in the Tillandsia utriculata Complex

Fig

ure

1

Til

lan

dsi

aa

esii

IR

amır

ezamp

Car

nev

ali

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Hab

itS

cale

bar

51

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Upp

erro

wf

rom

left

tori

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flow

erf

low

erw

ith

flor

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ract

rem

oved

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wer

wit

hou

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tfl

oral

bra

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als

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thre

ep

etal

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54

cmB

ased

onth

eh

olot

ype

GC

arn

eva

liamp

IM

R

am

ırez

69

73

(CIC

Y)

Mar

gin

sof

sep

als

and

flor

alb

ract

sar

eso

mew

hat

revo

lute

inth

efi

gure

s

74 Novon

obtusely acute 18ndash25 6 07ndash13 cm 2 sepalsbasally connate by 6ndash15 mm apple green slightlydarker and shinier than petals petals narrowly oblongapically rounded 32ndash38 6 04ndash08 cm slightlyoblique constricted ca 15 cm distally from base palegreen to cream-green basally white stamens exsertedin 2 series filaments concolorous with petals plicatelighter at the base 5ndash55 cm anthers brown 8 mmdorsifixed pollen yellow ovary 15 mm pyramidalnectar sweet abundant 31u Brix (total concentration ofsugars) stigma exserted slightly protruding above theanthers at anthesis lobes conduplicate somewhatspiraled 47ndash49 cm basally light green apicallyapple green Fruits and seeds not observed

Habitat distribution and phenology Tillandsiaaesii is found in low dry caducifolious forestsassociated with Cnidoscolus Pohl Plumeria L Pseu-dosmodingium Engler Agave L Opuntia (L) MillerPedilanthus Necker ex Poiteau Cordia elaeagnoidesDC Bursera simaruba Sargent xeromorphic epiphytessuch as Tillandsia ionantha Planchon T intermediaMez T caput-medusae E Morren T concolor L BSmith Encyclia adenocarpon (La Llave amp Lexarza)Schlechter Myrmecophila galeottiana (Reichenbach f)Rolfe Cactaceae such as Pachycereus (A Berger)Britton amp Rose and thorny leguminous trees andshrubs It has been collected at 0ndash600 m of elevationon the Pacific coast of Mexico Flowering specimenswere collected from March through July and fruit-ing specimens were collected from May throughSeptember

Eponymy The name is based on an acronym of theinitials of Dr Adolfo Espejo Serna a Mexican botanistassociated with herbarium UAMIZ of the UniversidadAutonoma Metropolitana in Mexico DF whose workin Bromeliaceae and other monocots has contributedgreatly to the knowledge of the Mexican flora

Tillandsia aesii is phenetically similar and proba-bly phylogenetically closest to T makoyana a speciesthat commonly features a rosette of straight erect (nottwisted and spreading) leaves Tillandsia makoyanafurthermore displays purple or light purple flowersand a slightly zygomorphic corolla as opposed to theactinomorphic green tubular corolla of T aesii Bothspecies have been collected at varying elevationsranging from sea level to ca 2000 m but T aesii hasonly been collected at elevations from sea level to ca600 m and in dryer environments

At least three other vegetatively similar taxa in theTillandsia utriculata complex display pale flowers Thewidespread T utriculata features longer thinner (easyto bend) peduncles and branches and the smaller whitecorollas are gibbous or somewhat inflated with thepetals convergent at apex Tillandsia dasyliriifolia and

T limbata are easily distinguished from T aesii bytheir inflorescences with much longer pedunclesbearing flowers with white slightly zygomorphiccorollas Tillandsia aesii is further distinguished fromthese two taxa by the proportionally smaller rosette ofstiff leaves that are slightly twisted In addition Tdasyliriifolia and T limbata are restricted to the humidlowlands of the Gulf Coast of Mexico (T limbata) or tothe Yucatan Peninsula (T dasyliriifolia)

Paratypes MEXICO Guerrero mpio Huitzuco Ama-titlan 12 km SE del Huitzuco 4 June 1985 J C Soto Nunezamp Aureoles C 8786 (MEXU) Jalisco mpio La Huertaalred Est Biol Chamela 3 Nov 1997 G Carnevali amp GSalazar 4767 (CICY) camino por la base SE del Cerro de laTambora 16 Jan 1985 P Magana Rueda amp E Lott 10(MEXU) Chamela Field Stat 100 km S of Puerto Vallarta26 Sep 1988 A Gentry 63981 (MO) Oaxaca mpio MatıasRomero carr entre La Ventosa y el entronque a Juchitan10 km N de La Ventosa 4 May 1997 G Carnevali GCampos M Gomez amp F May 4438 (CICY) mpio AsuncionTlacolulita dist Yautepec ca 9 km despues de Reformasobre camino a TehuantepecndashOaxaca 2 Sep 2002 IRamırez A Espejo A Lopez J Ceja y A Mendoza 998(CICY) mpio Asuncion-Ixtaltepec distrito Juchitan 1 kmdespues de Santiago de Ixtaltepec 2 Sep 2002 I RamırezA Espejo A Lopez J Ceja y A Mendoza 979 (CICY) 28 kmNW of La Ventosa along Trans-Isthmian hwy (rte 185) 17July 1958 R M King 630 (MICH TEX US) flat grazed area55 km NE of Juchitan along Pan-Amer hwy (rte 190) 2July 1958 R M King 360 (MEXU MICH TEX US) flatgrazed area 4 km N-NE of Tehuantepec along the Trans-Isthmian hwy (rte 185 amp 190) 5 July 1959 R M King 1352(LL) flat grazed area 16 km N-NE of Tehuantepec 7 July1959 R M King 1389 (TEX) mtns along rte 190 ca74 miles SE of Oaxaca 22 June 1960 R M King 2975(TEX US) Tehuantepec dry open savannah 2 Apr 1957M B Foster amp O C Van Hyning 2939 (US [2]) on rte 1856 km N of Union Hidalgo 10 July 1958 W G Williams Jr ampD Francoeur 21 (MICH) on rte 185 15 km N of UnionHidalgo at edge of steep drop into a small valley 11 July1958 Walter G Williams Jr amp Don Francoeur 24 (MICH)mpio Nejapa de Madero distrito Yautepec 24 km al N-NEde Rıo Hondo brecha a Asuncion Lachixonase 16 May1995 A Salinas T amp E Martınez-Correa 8155 (MEXU)

Tillandsia pinicola I Ramırez amp Carnevali sp novTYPE Mexico Yucatan Merida floreciendo encultivo 13 Jan 2005 I Ramırez amp G Carnevali1274 (holotype CICY [2] isotype MO) [Oaxacampio Santa Marıa Chimalapa Dist Juchitan ca20 km despues de el Mezquite 7 km despues deLazaro Cardenas rumbo a Santa Marıa Chima-lapa 16u469100N 94u509380W ca 385 mcolectada originalmente por I Ramırez AEspejo A Lopez-Ferrari A Mendoza amp J Cejael 2 Sep 2002] Figure 2

Haec species Tillandsiae makoyanae affinis est sed plantaambito parviore foliorum vaginis latissimis foliorum laminisproportione brevioribus apice acicularibus abrupte attenuatis(vs paulo apicaliter attenuatis) et habitatio pinicola abhorret

Volume 17 Number 1 Ramırez amp Carnevali 752007 New Species in the Tillandsia utriculata Complex

Fig

ure

2

Til

lan

dsia

pin

icol

aI

Ram

ırez

ampC

arn

eval

imdash

A

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it

Sca

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ower

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ower

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sep

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als

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onth

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76 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

Fig

ure

1

Til

lan

dsi

aa

esii

IR

amır

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Car

nev

ali

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Hab

itS

cale

bar

51

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mdashB

Upp

erro

wf

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left

tori

ght

flow

erf

low

erw

ith

flor

alb

ract

rem

oved

flo

wer

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hou

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an

dan

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als

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thre

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bar

54

cmB

ased

onth

eh

olot

ype

GC

arn

eva

liamp

IM

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am

ırez

69

73

(CIC

Y)

Mar

gin

sof

sep

als

and

flor

alb

ract

sar

eso

mew

hat

revo

lute

inth

efi

gure

s

74 Novon

obtusely acute 18ndash25 6 07ndash13 cm 2 sepalsbasally connate by 6ndash15 mm apple green slightlydarker and shinier than petals petals narrowly oblongapically rounded 32ndash38 6 04ndash08 cm slightlyoblique constricted ca 15 cm distally from base palegreen to cream-green basally white stamens exsertedin 2 series filaments concolorous with petals plicatelighter at the base 5ndash55 cm anthers brown 8 mmdorsifixed pollen yellow ovary 15 mm pyramidalnectar sweet abundant 31u Brix (total concentration ofsugars) stigma exserted slightly protruding above theanthers at anthesis lobes conduplicate somewhatspiraled 47ndash49 cm basally light green apicallyapple green Fruits and seeds not observed

Habitat distribution and phenology Tillandsiaaesii is found in low dry caducifolious forestsassociated with Cnidoscolus Pohl Plumeria L Pseu-dosmodingium Engler Agave L Opuntia (L) MillerPedilanthus Necker ex Poiteau Cordia elaeagnoidesDC Bursera simaruba Sargent xeromorphic epiphytessuch as Tillandsia ionantha Planchon T intermediaMez T caput-medusae E Morren T concolor L BSmith Encyclia adenocarpon (La Llave amp Lexarza)Schlechter Myrmecophila galeottiana (Reichenbach f)Rolfe Cactaceae such as Pachycereus (A Berger)Britton amp Rose and thorny leguminous trees andshrubs It has been collected at 0ndash600 m of elevationon the Pacific coast of Mexico Flowering specimenswere collected from March through July and fruit-ing specimens were collected from May throughSeptember

Eponymy The name is based on an acronym of theinitials of Dr Adolfo Espejo Serna a Mexican botanistassociated with herbarium UAMIZ of the UniversidadAutonoma Metropolitana in Mexico DF whose workin Bromeliaceae and other monocots has contributedgreatly to the knowledge of the Mexican flora

Tillandsia aesii is phenetically similar and proba-bly phylogenetically closest to T makoyana a speciesthat commonly features a rosette of straight erect (nottwisted and spreading) leaves Tillandsia makoyanafurthermore displays purple or light purple flowersand a slightly zygomorphic corolla as opposed to theactinomorphic green tubular corolla of T aesii Bothspecies have been collected at varying elevationsranging from sea level to ca 2000 m but T aesii hasonly been collected at elevations from sea level to ca600 m and in dryer environments

At least three other vegetatively similar taxa in theTillandsia utriculata complex display pale flowers Thewidespread T utriculata features longer thinner (easyto bend) peduncles and branches and the smaller whitecorollas are gibbous or somewhat inflated with thepetals convergent at apex Tillandsia dasyliriifolia and

T limbata are easily distinguished from T aesii bytheir inflorescences with much longer pedunclesbearing flowers with white slightly zygomorphiccorollas Tillandsia aesii is further distinguished fromthese two taxa by the proportionally smaller rosette ofstiff leaves that are slightly twisted In addition Tdasyliriifolia and T limbata are restricted to the humidlowlands of the Gulf Coast of Mexico (T limbata) or tothe Yucatan Peninsula (T dasyliriifolia)

Paratypes MEXICO Guerrero mpio Huitzuco Ama-titlan 12 km SE del Huitzuco 4 June 1985 J C Soto Nunezamp Aureoles C 8786 (MEXU) Jalisco mpio La Huertaalred Est Biol Chamela 3 Nov 1997 G Carnevali amp GSalazar 4767 (CICY) camino por la base SE del Cerro de laTambora 16 Jan 1985 P Magana Rueda amp E Lott 10(MEXU) Chamela Field Stat 100 km S of Puerto Vallarta26 Sep 1988 A Gentry 63981 (MO) Oaxaca mpio MatıasRomero carr entre La Ventosa y el entronque a Juchitan10 km N de La Ventosa 4 May 1997 G Carnevali GCampos M Gomez amp F May 4438 (CICY) mpio AsuncionTlacolulita dist Yautepec ca 9 km despues de Reformasobre camino a TehuantepecndashOaxaca 2 Sep 2002 IRamırez A Espejo A Lopez J Ceja y A Mendoza 998(CICY) mpio Asuncion-Ixtaltepec distrito Juchitan 1 kmdespues de Santiago de Ixtaltepec 2 Sep 2002 I RamırezA Espejo A Lopez J Ceja y A Mendoza 979 (CICY) 28 kmNW of La Ventosa along Trans-Isthmian hwy (rte 185) 17July 1958 R M King 630 (MICH TEX US) flat grazed area55 km NE of Juchitan along Pan-Amer hwy (rte 190) 2July 1958 R M King 360 (MEXU MICH TEX US) flatgrazed area 4 km N-NE of Tehuantepec along the Trans-Isthmian hwy (rte 185 amp 190) 5 July 1959 R M King 1352(LL) flat grazed area 16 km N-NE of Tehuantepec 7 July1959 R M King 1389 (TEX) mtns along rte 190 ca74 miles SE of Oaxaca 22 June 1960 R M King 2975(TEX US) Tehuantepec dry open savannah 2 Apr 1957M B Foster amp O C Van Hyning 2939 (US [2]) on rte 1856 km N of Union Hidalgo 10 July 1958 W G Williams Jr ampD Francoeur 21 (MICH) on rte 185 15 km N of UnionHidalgo at edge of steep drop into a small valley 11 July1958 Walter G Williams Jr amp Don Francoeur 24 (MICH)mpio Nejapa de Madero distrito Yautepec 24 km al N-NEde Rıo Hondo brecha a Asuncion Lachixonase 16 May1995 A Salinas T amp E Martınez-Correa 8155 (MEXU)

Tillandsia pinicola I Ramırez amp Carnevali sp novTYPE Mexico Yucatan Merida floreciendo encultivo 13 Jan 2005 I Ramırez amp G Carnevali1274 (holotype CICY [2] isotype MO) [Oaxacampio Santa Marıa Chimalapa Dist Juchitan ca20 km despues de el Mezquite 7 km despues deLazaro Cardenas rumbo a Santa Marıa Chima-lapa 16u469100N 94u509380W ca 385 mcolectada originalmente por I Ramırez AEspejo A Lopez-Ferrari A Mendoza amp J Cejael 2 Sep 2002] Figure 2

Haec species Tillandsiae makoyanae affinis est sed plantaambito parviore foliorum vaginis latissimis foliorum laminisproportione brevioribus apice acicularibus abrupte attenuatis(vs paulo apicaliter attenuatis) et habitatio pinicola abhorret

Volume 17 Number 1 Ramırez amp Carnevali 752007 New Species in the Tillandsia utriculata Complex

Fig

ure

2

Til

lan

dsia

pin

icol

aI

Ram

ırez

ampC

arn

eval

imdash

A

Hab

it

Sca

leb

ar5

10

cmmdash

B

Up

per

row

fr

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righ

tfl

ower

fl

ower

wit

hfl

oral

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ed

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erw

ith

out

flor

alb

ract

se

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owin

gth

eov

ary

styl

est

igm

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ents

an

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mro

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omle

ftto

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tfl

oral

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ct

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sep

als

and

two

pet

als

Sca

leb

ar5

3cm

B

ased

onth

eh

olot

ype

IR

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ırez

ampG

C

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ICY

)M

argi

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tein

the

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res

76 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

obtusely acute 18ndash25 6 07ndash13 cm 2 sepalsbasally connate by 6ndash15 mm apple green slightlydarker and shinier than petals petals narrowly oblongapically rounded 32ndash38 6 04ndash08 cm slightlyoblique constricted ca 15 cm distally from base palegreen to cream-green basally white stamens exsertedin 2 series filaments concolorous with petals plicatelighter at the base 5ndash55 cm anthers brown 8 mmdorsifixed pollen yellow ovary 15 mm pyramidalnectar sweet abundant 31u Brix (total concentration ofsugars) stigma exserted slightly protruding above theanthers at anthesis lobes conduplicate somewhatspiraled 47ndash49 cm basally light green apicallyapple green Fruits and seeds not observed

Habitat distribution and phenology Tillandsiaaesii is found in low dry caducifolious forestsassociated with Cnidoscolus Pohl Plumeria L Pseu-dosmodingium Engler Agave L Opuntia (L) MillerPedilanthus Necker ex Poiteau Cordia elaeagnoidesDC Bursera simaruba Sargent xeromorphic epiphytessuch as Tillandsia ionantha Planchon T intermediaMez T caput-medusae E Morren T concolor L BSmith Encyclia adenocarpon (La Llave amp Lexarza)Schlechter Myrmecophila galeottiana (Reichenbach f)Rolfe Cactaceae such as Pachycereus (A Berger)Britton amp Rose and thorny leguminous trees andshrubs It has been collected at 0ndash600 m of elevationon the Pacific coast of Mexico Flowering specimenswere collected from March through July and fruit-ing specimens were collected from May throughSeptember

Eponymy The name is based on an acronym of theinitials of Dr Adolfo Espejo Serna a Mexican botanistassociated with herbarium UAMIZ of the UniversidadAutonoma Metropolitana in Mexico DF whose workin Bromeliaceae and other monocots has contributedgreatly to the knowledge of the Mexican flora

Tillandsia aesii is phenetically similar and proba-bly phylogenetically closest to T makoyana a speciesthat commonly features a rosette of straight erect (nottwisted and spreading) leaves Tillandsia makoyanafurthermore displays purple or light purple flowersand a slightly zygomorphic corolla as opposed to theactinomorphic green tubular corolla of T aesii Bothspecies have been collected at varying elevationsranging from sea level to ca 2000 m but T aesii hasonly been collected at elevations from sea level to ca600 m and in dryer environments

At least three other vegetatively similar taxa in theTillandsia utriculata complex display pale flowers Thewidespread T utriculata features longer thinner (easyto bend) peduncles and branches and the smaller whitecorollas are gibbous or somewhat inflated with thepetals convergent at apex Tillandsia dasyliriifolia and

T limbata are easily distinguished from T aesii bytheir inflorescences with much longer pedunclesbearing flowers with white slightly zygomorphiccorollas Tillandsia aesii is further distinguished fromthese two taxa by the proportionally smaller rosette ofstiff leaves that are slightly twisted In addition Tdasyliriifolia and T limbata are restricted to the humidlowlands of the Gulf Coast of Mexico (T limbata) or tothe Yucatan Peninsula (T dasyliriifolia)

Paratypes MEXICO Guerrero mpio Huitzuco Ama-titlan 12 km SE del Huitzuco 4 June 1985 J C Soto Nunezamp Aureoles C 8786 (MEXU) Jalisco mpio La Huertaalred Est Biol Chamela 3 Nov 1997 G Carnevali amp GSalazar 4767 (CICY) camino por la base SE del Cerro de laTambora 16 Jan 1985 P Magana Rueda amp E Lott 10(MEXU) Chamela Field Stat 100 km S of Puerto Vallarta26 Sep 1988 A Gentry 63981 (MO) Oaxaca mpio MatıasRomero carr entre La Ventosa y el entronque a Juchitan10 km N de La Ventosa 4 May 1997 G Carnevali GCampos M Gomez amp F May 4438 (CICY) mpio AsuncionTlacolulita dist Yautepec ca 9 km despues de Reformasobre camino a TehuantepecndashOaxaca 2 Sep 2002 IRamırez A Espejo A Lopez J Ceja y A Mendoza 998(CICY) mpio Asuncion-Ixtaltepec distrito Juchitan 1 kmdespues de Santiago de Ixtaltepec 2 Sep 2002 I RamırezA Espejo A Lopez J Ceja y A Mendoza 979 (CICY) 28 kmNW of La Ventosa along Trans-Isthmian hwy (rte 185) 17July 1958 R M King 630 (MICH TEX US) flat grazed area55 km NE of Juchitan along Pan-Amer hwy (rte 190) 2July 1958 R M King 360 (MEXU MICH TEX US) flatgrazed area 4 km N-NE of Tehuantepec along the Trans-Isthmian hwy (rte 185 amp 190) 5 July 1959 R M King 1352(LL) flat grazed area 16 km N-NE of Tehuantepec 7 July1959 R M King 1389 (TEX) mtns along rte 190 ca74 miles SE of Oaxaca 22 June 1960 R M King 2975(TEX US) Tehuantepec dry open savannah 2 Apr 1957M B Foster amp O C Van Hyning 2939 (US [2]) on rte 1856 km N of Union Hidalgo 10 July 1958 W G Williams Jr ampD Francoeur 21 (MICH) on rte 185 15 km N of UnionHidalgo at edge of steep drop into a small valley 11 July1958 Walter G Williams Jr amp Don Francoeur 24 (MICH)mpio Nejapa de Madero distrito Yautepec 24 km al N-NEde Rıo Hondo brecha a Asuncion Lachixonase 16 May1995 A Salinas T amp E Martınez-Correa 8155 (MEXU)

Tillandsia pinicola I Ramırez amp Carnevali sp novTYPE Mexico Yucatan Merida floreciendo encultivo 13 Jan 2005 I Ramırez amp G Carnevali1274 (holotype CICY [2] isotype MO) [Oaxacampio Santa Marıa Chimalapa Dist Juchitan ca20 km despues de el Mezquite 7 km despues deLazaro Cardenas rumbo a Santa Marıa Chima-lapa 16u469100N 94u509380W ca 385 mcolectada originalmente por I Ramırez AEspejo A Lopez-Ferrari A Mendoza amp J Cejael 2 Sep 2002] Figure 2

Haec species Tillandsiae makoyanae affinis est sed plantaambito parviore foliorum vaginis latissimis foliorum laminisproportione brevioribus apice acicularibus abrupte attenuatis(vs paulo apicaliter attenuatis) et habitatio pinicola abhorret

Volume 17 Number 1 Ramırez amp Carnevali 752007 New Species in the Tillandsia utriculata Complex

Fig

ure

2

Til

lan

dsia

pin

icol

aI

Ram

ırez

ampC

arn

eval

imdash

A

Hab

it

Sca

leb

ar5

10

cmmdash

B

Up

per

row

fr

omle

ftto

righ

tfl

ower

fl

ower

wit

hfl

oral

bra

ctre

mov

ed

flow

erw

ith

out

flor

alb

ract

se

pal

san

dp

etal

ssh

owin

gth

eov

ary

styl

est

igm

afi

lam

ents

an

dan

ther

sB

otto

mro

wfr

omle

ftto

righ

tfl

oral

bra

ct

two

sep

als

and

two

pet

als

Sca

leb

ar5

3cm

B

ased

onth

eh

olot

ype

IR

am

ırez

ampG

C

arn

eva

li1

27

4(C

ICY

)M

argi

ns

ofse

pal

san

dfl

oral

bra

cts

are

som

ewh

atre

volu

tein

the

figu

res

76 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

Fig

ure

2

Til

lan

dsia

pin

icol

aI

Ram

ırez

ampC

arn

eval

imdash

A

Hab

it

Sca

leb

ar5

10

cmmdash

B

Up

per

row

fr

omle

ftto

righ

tfl

ower

fl

ower

wit

hfl

oral

bra

ctre

mov

ed

flow

erw

ith

out

flor

alb

ract

se

pal

san

dp

etal

ssh

owin

gth

eov

ary

styl

est

igm

afi

lam

ents

an

dan

ther

sB

otto

mro

wfr

omle

ftto

righ

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76 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

Epiphytic herb 50ndash60 cm high the rosettes appar-ently monocarpic rosette with 20 to 30 leaves whenflowering leaves erect with divergent apices withfunnelform rosette apices erect when young or fruitingand then the rosette shape subcylindrical foliar sheathsbarely impounding Foliar sheaths with incurvedmargins 6ndash10 6 6ndash8 cm elliptical green adaxiallydark castaneous upon drying foliar blades narrowlytriangular abruptly attenuating into an acicular apexmargins incurved coriaceous rather stiff conspicuouslynerved abaxially 15ndash18 cm long 38ndash45 cm wide atthe base concolorous dull green slightly white-lepidoteabaxially Inflorescence an erect panicle 40ndash60 cmovoid-ellipsoid some of the lowermost branches 2-pinnate 50ndash60 cm long from the base of the rosette with11 to 16 spikes the new inflorescence developing slowlyand taking up to 2 months before the first flowers opensuccessively flowering basipetally and producing newflowers for 2 to 3 weeks peduncle 10ndash15 cm shorter oras long as the leaves bright crimson red glabrous ca1 cm diam covered by tubular acute imbricatingbracts internodes 2ndash3 cm long bracts of the peduncleslightly shorter than the internodes and not totallyenveloping them crimson with green tinges glabrous35ndash45 6 2ndash26 cm (longer and wider toward thebase) widely triangular acute margins hyaline con-spicuously nerved when dry slightly surpassing theinternodes inflorescence rachis crimson red of smallerdiameter than the peduncle naked flexuous thebranches diverging at angles of 45undash60u to the in-florescence rachis basal spikes 15ndash17 cm apical ones9ndash12 cm sterile portion of basal ones 65ndash75 cm of theapical ones 18ndash25 cm rachis green with dark redtinges almost entirely covered by floral bracts dorso-ventrally flat with flowers in 2 rows (5 to)10 flowers perbranch bracts of the spikes 1ndash18 cm (longer ones atbase) widely triangular obtuse pale crimson with slightgreenish tinges glabrous similar in shape to the floralbracts floral bracts 16ndash18 6 9ndash10 mm ovate-ellipticacute conspicuously nerved margin hyaline shorterthan sepals barely covering the rachis those subtendingmature flowers dull crimson apple green toward dorsalportion more immature flowers apple green with crimsonconfined to the margins and apex glabrous shinyFlowers 33ndash35 cm actinomorphic forming an acuteangle with the rachis up to 7 flowers open daily perinflorescence (1 at a time per spike) each flower lasting1 day protandrous odorless basally very slightlygeniculate sepals spathulate-elliptic broadly acute15 cm 6 6 mm the 2 adaxial ones basally connatefor 4ndash6 mm apple green apically and marginally dullred-maroon adaxially sparsely lepidote more so towardapex petals narrowly oblong apically rounded 28ndash3605 cm slightly oblique constricted about 15 cmdistally from base dull purple paler toward the base

the portion hidden by the sepals hyaline exserted

portion 18ndash2 cm long stamens exserted in 2 series of

unequal length filaments pale cream-green plicate

upon drying lighter at the base 37ndash38 cm flatter

toward the base anthers brown or black-brown 45 mm

8ndash14 mm thick dorsifixed pollen yellow ovary 86 ca

3 mm ellipsoid-pyramidal nectar sweet abundant

stigma exserted slightly protruding above the anthers

at anthesis lobes conduplicate not spiraled style

44 cm basally light green apically apple green Fruits

and seeds not observed

Habitat distribution and phenology Tillandsia

pinicola grows on pine trees mostly Pinus oocarpa

Schiede ex Schlechtendal and is associated with

Catopsis berteroniana (Schultes amp Schultes f) Mez T

streptophylla Scheidweiler ex E Morren and T aff

fasciculata Swartz It occurs at 300ndash400 m in

elevation and is known only from Mexico (Oaxaca)

It has been collected in flower during January and in

fruit in September

Tillandsia pinicola is one of the most distinctive

members of the T utriculata complex by its small size

erect green leaves that are relatively short and with

wide sheaths and its inflorescence with short

peduncles and purple flowers It is very similar to T

makoyana and the new species herein described as T

aesii but it differs from them by the overall smaller

size the foliar sheaths wide and abruptly attenuating

into a narrow foliar blade that ends in an acicular

apex leaves strongly nerved when alive or dry leaves

dark green on both sides but lightly white-lepidote on

the abaxial surface the inflorescence with a short

peduncle that is usually shorter than the leaves much

shorter flowers and floral bracts and branches that are

both fewer in number and shorter

Paratypes MEXICO Oaxaca mpio Santa Marıa Chi-malapa distrito Juchitan ca 20 km despues de El Mezquite7 km despues de Lazaro Cardenas rumbo a Santa MarıaChimalapa 2 Sep 2002 I Ramırez A Espejo A Lopez-Ferrari A Mendoza amp J Ceja 987 (CICY UAMIZ) mpioLazaro Cardenas dist Juchitan 36 km al E de LazaroCardenas hacia Santa Marıa Chimalapa entrando por elMezquite 22 Jan 1988 R Torres C amp E Martınez 11225(IEB MEXU)

Acknowledgments To the Elizabeth Bascom Fel-

lowship for Latinoamerican Women and the Missouri

Botanical Garden for a scholarship granted to the first

author to the Bromeliad Society for financial support to

the first author for field equipment to the Marie Selby

Botanical Gardens for financial support to the first

author while working on the Bromeliaceae of Mexico

project to Adolfo Espejo Ana Rosa Lopez-Ferrari

Aniceto Mendoza and Jacqueline Ceja for field support

in Chiapas Guerrero Jalisco Michoacan Morelos and

Volume 17 Number 1 Ramırez amp Carnevali 772007 New Species in the Tillandsia utriculata Complex

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon

Oaxaca to Francisco Chi May Filogonio May SilviaHernandez-Aguilar Lorena Can-Itza and Jose LuisTapia for herbarium and field assistance in the YucatanPeninsula when collecting preparing herbarium speci-mens and searching for bibliographical material ToFrancisco Lorea-Hernandez Curator of HerbariumXAL and Carlos Duran for field assistance in Veracruzand Puebla Our gratitude to Demetria Mondragon formaterial collected in Chiapas and Oaxaca that allowsa better understanding of the species in this complexTo Derek Butcher and Walter Till for useful commen-taries and iconographic material of species in thiscomplex To William Cetzal Ix for his help inelaborating the plates of the two new species We aregrateful to the following herbaria for loans CIQR FGH LL MEXU MICH MO TEX UAMIZ UCAMUS and XAL

Literature Cited

Barfuss M H J R Samuel W Till amp T F Stuessy 2005Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) based on DNA sequence data from sevenplastid regions Amer J Bot 92 337ndash351

Espejo S A A R Lopez-Ferrari amp I Ramırez M 2005Bromeliaceae Pp 1ndash305 in Flora de Veracruz Fasc 136Instituto de Ecologıa A C Xalapa Veracruz Mexico

Gardner S 1984 Natural hybridization in Tillandsiasubgenus Tillandsia Selbyana 7 380ndash393

mdashmdashmdash 1986 Preliminary classification of Tillandsia basedon floral characters Selbyana 9 130ndash146

Luther H 1995 An annotated checklist of the Bromeliaceaeof Costa Rica Selbyana 16 230ndash234

Mez C 1896 Bromeliaceae In C de Candolle MongraphiaePhanerogamarum Vol 9 Sumptibus Masson Paris

Morales J F 2003 Bromeliaceae Pp 297ndash375 in B EHammel M H Grayum C Herrera amp N Zamora (editors)Manual de Plantas de Costa Rica Volumen II MonogrSyst Bot Missouri Bot Gard 92

Ramırez I G Carnevali amp F Chi 2004 Portraits ofBromeliaceae from the Mexican Yucatan PeninsulamdashIVTillandsia dasyliriifolia Baker Taxonomy and reproduc-tive biology J Bromeliad Soc 54 112ndash121

Spencer M A amp L B Smith 1993 Racinaea a new genus ofBromeliaceae (Tillandsioideae) Phytologia 74 151ndash160

Smith L B amp R J Downs 1977 Tillandsioideae(Bromeliaceae) Pp 663ndash1492 in Flora Neotropica Mono-graph 14 Part 2 Hafner Press New York

Terry R G G K Brow amp R G Olmstead 1997Phylogenetic relationships in subfamily Tillandsioideae(Bromeliaceae) using ndhF sequences Syst Bot 22333ndash345

78 Novon