vázquez-garcía et al. 2013. echeveria yalmanantlanensis

8/11/2019 Vzquez-Garca Et Al. 2013. Echeveria Yalmanantlanensis

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Echeveria yalmanantlanensis (Crassulaceae): A new speciesfrom Cerro Grande, Sierra de Manantln, western Mexico

J. ANTONIOVZQUEZ-GARCA1

(

DAVIDJIMENO-S.

2

(

RAMNCUEVAS-G.

3

MIGUELCHZARO-B.4( ANDMIGUELA. MUIZ-CASTRO1

1Instituto de Botnica, Departamento de Botnica y Zoologa, Universidad de Guadalajara-CUCBA,Zapopan, C.P. 45110 Jalisco, Mexico; e-mails: [email protected];[email protected] de Biologa, Universidad Veracruzana, Xalapa, Veracruz C.P. 91090, Mexico; e-mail:[email protected] de Ecologa y Recursos Naturales, Universidad de Guadalajara-CUCSUR, Autlnde Navarro, Jalisco C.P. 48900, Mexico; e-mail: [email protected] de Geografa y Ordenamiento Territorial, Universidad de Guadalajara-CUCSH,

Guadalajara, Jalisco C.P. 45101, Mexico; e-mail: [email protected]

Abstract. Echeveria yalmanantlanensis, a new species from eastern Sierra de Ma-nantln, Biosphere Reserve, on the Jalisco-Colima border in western Mexico, is de-scribed and illustrated, and a table for comparison of morphologically related taxa is

provided. This species belongs to series Valvatae, differing from the other members ofthe series by its acaulescent habit, shape and color of leaves and bracts, size of scape,

bract pattern on scape, size and color of the corolla, habitat, and geographic distrib-ution. There is support for the hypotheses that this species is endemic to the calcar-eous massif Cerro Grande and is already in danger of extinction.

Key Words:Allopatric, Endangered, Endemic, Echeveriaseries Valvatae.

Resumen. Se describe e ilustra Echeveria yalmanantlanensis, una especie nueva dela Sierra de Manantln oriental, en los lmites de los estados de Colima y Jalisco,occidente de Mxico; se provee un cuadro comparativo para la identicacin de ta-xones relacionados morfolgicamente. Esta especie pertenece a la serie Valvatae, di-ere de los otros dos miembros de la serie por su hbito acaulescente, forma y colorde sus hojas y brcteas, tamao del escapo, patrn de brcteas en el escapo, tamao dela corola, color de spalos y de corola, distribucin geogrca y hbitat. Se apoyan lashiptesis de que esta especie es endmica del macizo calcreo Cerro Grande y de quese encuentra en peligro de extincin.

EcheveriaDC. comprises some 140 knownspecies, and 95 % of these are found inMexico, the richest center of diversity andendemism for the genus (Uhl, 1992; Thiede,1995; Meyrn & Lpez-Chvez,2003). Thereare about ten species ofEcheveriain the stateof Jalisco (Pilbeam,2008), mostly distributedin temperate mountainous rocky areas, withonly two of these occurring in tropical dry

forest: E. pringlei (Watson) Rose and E.elegans var. tuxpanensis Walther. Eventhough the Sierra de Manantln Biosphere

Reserve in Western Mexico has been exten-sively explored for decades, no Echeveriaspecies has ever been recorded from thisgeographical area (Vzquez-Garca et al.,1995; Vzquez-Garca & Givnish, 1998;Cuevas & Jardel, 2004). However, during a

botanical excursion to Cerro Grande, Sierrade Manantln, in 2002, led by MiguelChzaro and including Ignacio Contreras

and Antonio Machuca, two undeterminedspecies of this genus were found. One ofthese species, a short-statured, pink-owered

Brittonia 65(3): 273Y279 (2013), DOI 10.1007/s12228-012-9274-9ISSN: 0007-196X (print) ISSN: 1938-436X (electronic) 2013, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

Published online: 9 July 2013


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Echeveria morphologically similar to E.fulgens Lemaire, was found at higher eleva-tions (22002300 m) in oak forest withnumerous individuals in each of several

populations. This taxon remains to be

studied. The other one, an extremely rareand unusual ivory-white owered tropical

Echeveria, was found by Mr. Contreras ina small population with few individuals ona single limestone outcrop at 1550 maltitude. Soon after, Mr. Chzaro suggestedthat the taxon was an undescribed speciesof series Valvatae Moran, a series consist-ing of two species: Echeveria valvataMoran and E. calycosa Moran (Moran,1963, 1967). In this paper, we describe a

third known species of series Valvatae andargue that it is a narrow endemic in danger ofextinction.

Echeveria yalmanantlanensis A.Vzquez& Chzaro, sp. nov. Type: Mexico. Col-ima-Jalisco: Municipio de Comala-Toli-mn, camino a Campo Cuatro, CerroGrande, on a rock of a limestone slope,1550 m, 26 Sep 2011 (, fr), J. AntonioVzquez-Garca, M. Chzaro B. & J.

Padilla-Lepe 9175 (holotype: IBUG; iso-type: NY). (Figs.1,2)

Planta perennis glabra; rosula solitaria, sessilis, 822(35) cm lata; folia oblonga, oblong-oblanceolata, vixglauca, 613(20) cm longa, 1.33.6 cm lata, 4 mm crassa;rami oriferi 13, erecti, 5.15.8 dm alti, foliis 1420instructi; cincinnus solitarius, 813(16) cm longus, 18orus; pedicelli 1.53.23 mm, breves; calyx 713 mmlongus, segmentis erectis, inaequalibus; corolla eburnea,glauca, 10 mm longa, segmentis valvatis, triangulo-lanceo-latis, acutis, 0.5-1 mm connatis.

Perennial with a single herbaceous rosette,56822(35) cm, acaulescent, lax, gla-

brous; rootsbrous; leaves 918, 613(20)1.33.6 cm, 4 mm thick, the blades oblanceo-late to oblong-oblanceolate, acuminate atthe apex, slightly keeled at the base,ventrally attened and dorsally convex,grayish green to green, the surface waxy

when young, the margin straight, hyalineand pinkish; scapes 13, 3846(54) cmfrom base to rst ower, 5158(70) cm in

total length, lateral and ascendant, pale yellow-ish green; cincinnus solitary, 813(16)1.52.5 cm, circinate before anthesis, becomingerect after anthesis, grayish white to gray-ish green and waxy; scape bracts 1420,

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5.90.5

1.28 cm, spirally arranged,erect, oblong-oblanceolate to narrowly tri-angular, ventrally attened and dorsallyconvex, acute to acute-rounded, usuallywith two spurs at the base, grayish green;cincinnus bracts 22, 82237.5 mm,2 mm thick, ovate to oblanceolate, acumi-nate at the apex, slightly curved inwardand adaxially concave, tightly imbricateand succulent before anthesis, sparselyarranged in zigzag and dried soon after

anthesis, with two spurs at the base,grayish green at the apex, pinkish towardthe base; bractlets 22, 358 mm, triangu-lar, with two spurs at the base, above the rightside of each bract before anthesis, below theright side of a bract after anthesis, eshy beforeduring and after anthesis, ivory to yellowish;

pedicels 1.53.2 3 mm, grayish green; owers1822(44), 1314 mm long, 11 mm wide atthe base, 16 mm wide at the apex, in a zigzag

pattern; sepals 5, unequal, equal or larger than

the corolla, larger ones 15

176 mm, smallerones 6.4142.84 mm, triangular, barelyimbricate if at all or rarely spaced at the base,erect and appressed before anthesis, ascendantafter anthesis, lanceolate, acuminate, spreadingafter anthesis, inconspicuously bicolored, gray-ish white before and during anthesis, becominggreen after anthesis and pinkish at the base;corolla 1213810 mm, pentagonal, conical,slightly urceolate, forming a pale yellow tube,ivory-white to pale yellow; petals fully valvateat aestivation, partially valvate in the lower

portion (2/3 to 1/2) during anthesis, slightlykeeled, triangular-lanceolate, spreading out-ward at the upper third, connate at the base0.51 mm, inconspicuously bicolored, ivory-white to glaucous pale yellow, pinkish at theapex after anthesis; stamens 10, at two levels,the epipetalous ones 55.1 mm long, theinsertion point 1 mm above the base, theantipetalous ones 6.36.4 mm long, the inser-tion point at the base, between corolla andcarpels, the laments ivory-white to pale

yellowish; anthers 0.9 0.5 mm, the epi-petalous ones yellow, the antipetalous onesbrownish; nectar glands 0.511.32 m,

274 BRITTONIA [VOL 65


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oblong, transversely narrow, hyaline pale

amber; carpels 7.53 mm; styles 2.6 mmlong, the lower half mostly white, green at theapex; follicles erect, 7.53 mm, erect, reddish;

seeds numerous, 0.390.900.160.31 mm,

elliptic, narrow and acute at the apex andbase, reticulate and wrinkled, pale brownto hyaline.

FIG. 1. Echeveria yalmanantlanensis.A. Habit. B. Cincinnus detail. C. Flower in cross section, seen from above.D.Flower in lateral view.E. Flower dissected.F.Carpels.G. Leaf and outlines of cross section at two levels. H. Scape

bracts before rst ower and outlines of cross section at one level. I. Bracts at cincinnus. J. Bractlets adjacent tobracts.K. Seed (A, B, DJ fromVzquez-Garca 7830b, IBUG; C from photograph by M. Chzaro at Jess Trujillossucculent home garden (August 2004); K scanned from Vzquez-Garca 7830b, IBUG.)

275VZQUEZ-GARCA ET AL.: ECHEVERIA(CRASSULACEAE)2013]


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Distribution and ecology.Echeveria yal-manantlanensis is known only from the typelocality, growing on a single rock, at 1550 mon steep karstic topography with a thin(ca. 2 cm) layer of soil, closely sharing thehabitat with the rupiculous Tillandsia capi-tata Griseb., under tropical dry forest with

Jatropha bartlettii Wilbur, Bursera mac-vaughiana Cuevas & Rzed., and Agaveattenuata Salm-Dyck. The climate of the

region is semi-warm sub-humid (A)Ca (w1)(w) of Kppen (Garca,1973, Vzquez-Garcaet al.,1995).

Phenology.Flowering from late July toearly October; fruiting from September toDecember.

Etymology.The specic epithet honors

both plant ecologist Yalma Luisa Vargas-Rodrguez, from the Department of Biologi-cal Sciences, Louisiana State University, and

FIG

. 2. Echeveria yalmanantlanensis. A. Rosette. B. Cincinnus, lateral view. C. Cincinnus, front view, with fullvalvate petals at aestivation and half valvate petals at anthesis. D, E. Flower in longitudinal section, showing sepals,petals, carpels, and stamens.



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he Sierra de Manantln, western Mexico,where she has conducted relevant ecolog-ical plant community sampling and analy-sis of both maple forest and tropical dryforest (Vargas-Rodrguez et al., 2005,

2010).

Additional specimens examined: MEXICO. Col-ima-Jalisco: Municipios Comala-Tolimn border,camino de Campo Cuatro a La Ailera, Cerro Grande,tropical dry forest, on a rock of a limestone slope,1500 m, 18 Jul 2004 (), Vzquez-Garca & Contreras7830a, 7830b(IBUG).

Echeveria yalmanantlanensis, except forits inconspicuously bicolored sepals, shareswithEcheveria calycosaandEcheveria valvatamost of the characters of series Valvatae(Moran, 1963; Meyrn & Lpez-Chvez,2003), such as single cincinnus, closely imbri-cate bracts, erect or ascending sepals equal to orlarger than the corolla, reticulate seeds, pentag-onal corollas with segments free nearly to the

base and valvate petals, the latter an unusualcharacter within the genus and restricted toseries Valvatae. However, Echeveria yalma-nantlanensis differs from these other twospecies in its acaulescent habit, shape and colorof leaves and bracts, size of scape, bract patternon scape, size and color of the corolla, habitat,and geographic distribution (TableI). Echeve-ria yalmanantlanensisis now the third knownmember of series Valvatae, this species is theonly member of the series with ivory-white to

pale yellow corollas, and the only one in theseries inhabiting tropical dry forest at 1500 m inelevation. Echeveria yalmanantlanensis alsorepresents the rst member of the series forthe states Colima-Jalisco and is the rst record

of Echeveria for the Sierra de ManantlnBiosphere Reserve (Vzquez-Garca et al.,1995; Vzquez-Garca et al., 1998; Cuevas &Jardel,2004).

Field studies indicate that Echeveria yal-manantlanensis is endemic to the calcareousmassif Cerro Grande. Several major efforts

TABLEIDIFFERENCES BETWEENECHEVERIA YALMANANTLANENSISAND OTHER MEMBERS OF SERIES VALVATAE.

E. calycosa E. valvata E. yalmanantlanensis

Rosette diameter 510(15) cm 815(20) cm 822(35) cmRosette habit Shortly caulescent Shortly caulescent Acaulescent Blade shape Spatulate, rounded at

the apex and weaklymucronate

Cuneate-spatulate to narrowlyobovate or somewhat rhombic

Oblanceolate to oblong-oblanceolate, acuminateat the apex

Leaf color Pale green Pale green Grayish greenLeaf size 2.551.52.5 cm 41023.5 cm 613(20)1.33.6 cmScape length to

rstower521 cm 1035 cm 3846(54) cm

Distribution ofscape bracts

Absent in the lower18 cm

Absent in the lower 110 cm Present throughout

Corolla length 7.5

10 mm 7

11 mm 12

13 mmColor of sepals Conspicuously bicolored,pinkish at the base andpale green at the apex

Conspicuously bicolored, green,pinkish at the base, becominggreen

Inconspicuously bicolored,grayish white duringanthesis, becominggreen with pinkishat the base

Corolla color Conspicuously bicolored,yellow at the apex, pinkishto reddish at the base,orange in appearance

Conspicuously bicolored,glaucous raspberry red,appearing pinkish, palerat the base

Inconspicuously bicolored,ivory-white to paleglaucous yellow

becoming pinkish atthe apex after anthesis

Distribution Southern Uruapan,Michoacn (Tzararacua,Ro Cupatitzio, Tributario

La Tinaja)

Southwestern and northernState of Mexico (Jilotepec,

Nachititla, Piedra Grande,

Temascaltepec, Valle de Bravo)

Colima-Jalisco border(Cerro Grande)

Habitat Pine oak forest Oak forest Tropical dry forest Elevation 1200 m 14501800 m 1550 m



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have failed to locate additional populations,including moderate exploration through oris-tic and ecological studies in tropical dry forestsof both the calcareous Cerro Grande (Vzquez-Garca et al., 1995; Vzquez-Garca &

Givnish1998,2000) and the adjacent volcanicareas (perhaps substrate barriers); Sierra deManantln central (Vzquez-Garca et al.,1995; Cuevas & Jardel, 2004; Vargas-Rodrguez, et al., 2005; Luqun-Snchez,2003) and Nevado de Colima (CONANP-SEMARNAT,2006). Since its discovery, near-ly a decade ago, six guided attempts by at leastsix experienced botanists were conducted inorder to locate additional populations in thevicinity of Campo Cuatro, Cerro Grande, all

without success. A recent survey, on the agavesof western Mexico (Vzquez-Garca et al.,2007) allowed us to conduct further unsuccess-ful explorations to other potential habitats ofE.

yalmanantlanensis, such as the calcareoushabitats with tropical dry forest in the states ofColima (Coquimatln, Ixtlahuacn, Jalisco;Pihuamo, and Michoacn: Aquila-Coalcomnand Maruata). The other members of series

Valvatae do not overlap in their narrow geo-graphic range (Fig.3).

We suggest that its current pattern andmorphological divergence from other mem-

bers of the series (Table I) perhaps resultedfrom a relatively recent process of allopatricspeciation within the series.

Observational data support the hypothesisthatEcheveria yalmanantlanensisis currentlyendangered with extinction, probably due toovercollection. As far as we know the speciesis conned to a single rock and for nearly adecade has declined in number of individualswith no evident natural extrinsic threat. Wedo not recommend Echeveria yalmanantla-nensisas an ornamental because it is difcult

to cultivate. For reasons unknown, variousspecimens have died at home gardens inGuadalajara and Talpa de Allende after twoand three years of cultivation respectively.Overcollection seems to be a major threat,since the original population size of ca. 14individuals has drastically declined to twoindividuals in situ and two individuals ex situ.Thus, this species must be immediately

FIG. 3. Known populations of Echeveriaseries Valvatae in western and central Mexico.



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included in the Mexican Endangered SpeciesAct, NOM-ECOL-059 (2001) as an endan-gered species.

Further systematic exploration to locateadditional populations of Echeveria yalma-nantlanensisare urgently needed, particularlyin the vicinity of Cerro Jumpiche and Piedrade Juluapan, Cerro Grande, a few kilometerssouthwest of the type locality. Ecological andgenetic studies are recommended. We urgecurrent managers at the Sierra de ManantlnBisosphere Reserve to take legal and physicalaction to prevent Echeveria yalmanantlanen-

sis from vanishing forever and to promoteresearch into the causes that might be drivingthis species to extinction.

Meanwhile, efforts are being made to prop-agate the species at the University of Guadala-

jara-CUCBA, Zapopan, using seed germinationand tissue-culture methods on seeds and scape

bracts. These efforts include the elaboration of aconservation strategy for the species.

Acknowledgments

We thank Ignacio Contreras, AntonioMachuca, Jess Trujillo, Jess Padilla-Lepe

and Yocupitzia Ramrez for assistance witheldwork in the Cerro Grande, Sierra deManantln. We also thank the anonymousreviewers of this manuscript for their assis-tance, Jess G. Gonzlez Gallegos for his aidand suggestions to improve this manuscript,Martn and Rafael Soltero for in-vitro repro-ducing of Echeveria yalmanantlanensis, Ser-vando Carvajal for providing a Latin diagnosis,

Noem Jimnez Reyes for assistance with thepreparation of Fig.1K, and Ramn Rodrguez

Macas, Chairman at Departamento de Botn-ica y Zoologa and Salvador Mena Mungua,Campus Chancellor, of the Universidad deGuadalajara, fornancial support and researchfacilities.

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