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Pachyphytum huastecanum (), a New Species from the Huasteca Canyon, Nuevo León, Mexico

Article in Cactus and Succulent Journal · June 2020 DOI: 10.2985/015.092.0207

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Pachyphytum huastecanum (Crassulaceae), a new species from the Huasteca Canyon, Nuevo León, Mexico

Abstract: Pachyphytum huastecanum is described as a new species discovered in the Cañón de la Huasteca in the Sierra Madre Oriental in the Mexican state of Nuevo León. This species is placed in section Diotostemon, where it is morphologically related to , from which it differs by its shorter stems, compact rosettes, flat, olive-green colored leaves, shorter pedicels, its campanulate flowers and obovate petals. Addition- ally, two DNA diagnostic nucleotides were found at the ITS2 nuclear genome region which differentiate the new species from other 19 Pachyphytum species. Resumen: Pachyphytum huastecanum es descrito como nueva especie descubierta en el Cañón de la Huasteca en la Sierra Madre Oriental en el estado mexicano de Nuevo León. Esta especie es ubicada en la sección Diot- ostemon, en donde está morfológicamente relacionada a Pachyphytum hookeri, de la cual difiere por sus tallos más cortos, rosetas compactas, hojas planas y de color verde oliva, pedicelos más cortos, sus flores campanuladas y por sus pétalos obovados. Adicionalmente, dos nucleótidos diagnósticos de ADN fueron encontrados en la región genómica nuclear ITS2, los cuales lo diferencian de otras 19 especies de Pachyphytum. Key words: Huasteca Canyon, ITS2, matK, Pachyphytum hookeri, rbcL

Introduction specifically to the greenhouses known as National Laboratories of Biodiversity (Laboratorio Nacional de In March 2016, Michael Bechtold and Wolf- Biodiversidad (LANABIO)), where almost all Pachy- gang Metorn, German enthusiasts of the Agavace- phytum species are also sheltered. ae family, visited El Pajonal in the Huasteca Can- The produced flowers in March 2018 and yon to search for Agave albopilosa I. Cabral, Vil- after a thorough comparison with related Pachyphy- lareal & A.E. Estrada. Looking for more plants tum species, morphological diagnostic characters of this rare species they wandered along the for this new species were found. At the same time, Arroyo Potreritos where they found a species of DNA sequences of rbcL and matK of the chloroplast Pachyphytum Link, Klotzsch & Otto. They sent pic- genome, and ITS2 region from the nuclear genome tures of the plants to Julia Etter and Martin Kristen, were obtained for one Pachyphytum huastecanum speci- editors of the Crassulaceae database website (www. men and then compared with sequences of P. brachetii crassulaceae.com). The specimens resembled P. hookeri J.Reyes, O.González & A.Gutiérrez, P. bracteosum Link, A. Berger, however, that species occurs about 370 kms Klotzsch & Otto, P. brevifolium Rose, P. caesium Kim- south of El Pajonal as the crow flies. It was not until nach & Moran, P. compactum Rose, P. contrerasii Pérez- November 2016 when Julia Etter and Martin Kris- Calix, I.García & Cházaro, P. fittkaui Moran, P. garci- ten, searching for species of the Crassulaceae family, ae Pérez-Calix & Glass, P. glutinicaule Moran, P. h o o k - and particularly those belonging to the Echeve- eri, P. kimnachii Moran, P. longifolium Rose, P. machu- ria, went to the Huasteca Canyon to collect this new cae I.García, Glass & Cházaro, P. oviferum J.A.Purpus, species of the genus Pachyphytum, along with P. w e r - P. rzedowskii I.García, Pérez-Calix & J.Meyrán, P. salt- dermannii Poelln., the northernmost known to date. ense Brachet, Reyes & Mondragón, P. viride E.Walther, Plants were not flowering at this time of the year and P. viscidum J.Reyes & L.E.Cruz-López (described in living plant material was sent to the facilities of the 2019) and P. werdermannii. The sequences of the afore- Botanical Garden of the Biology Institute of Uni- mentioned 19 Pachyphytum species were previously versidad Nacional Autónoma de México (UNAM), generated to build a DNA barcoding library in collabo- 1Jardín Botánico, Instituto de Biología, UNAM, Apartado ration with the University of Guelph and Barcode Of Postal 70-614, Ciudad Universitaria, 04510, Ciudad de Life Data Systems (www.boldsystems.org) in 2008. At México, México. email: [email protected] that time, P. confusum Pérez-Calix, Guadián-Marín & 2www.crassulaceae.com. email: [email protected] 3 I. García, P. neglectum J.Meyrán, and P. rogeliocardenasii Laboratorio de Sistemática Molecular, Instituto de Biología, UNAM, Ciudad Universitaria, 04510, Ciudad de México, Pérez-Calix & R.Torres had not yet been described and México. email: [email protected] therefore were not sampled.

2020 VOLUME 92 NUMBER 2 Author Copy 141 2. Stems of Pachyphytum huastecanum. Photo by Julia Etter & Martin Kristen.

1. Pachyphytum huastecanum flowering in habitat. Photo by Julia Etter & Martin Kristen.

A table with the diagnostic nucleotides of Pachy- phytum species accompanied the description of P. viscidum in a character-based approach (Reyes et al, 2019) and is updated here to include P. huastecanum. For details of DNA extraction, sequencing and meth- ods for the analysis of the sequences see Reyes et al (2019). 3. Pachyphytum huastecanum rosettes. Photo by Julia Pachyphytum huastecanum J. Reyes, Etter & Martin Kristen. Etter & Kristen sp. nov. bluish-green colored, somewhat farinose, margin Pachyphytum huastecanum, P. hookero entire, apex acute, hyaline and reddish (Fig. 4). Axil- (Salm-Dyck) Berger affiniis sed caulibus brevibus, lary floral stem close to the apex of the branches, foliis oblanceolatis, applanatis, (vs. caulibus longis, 30–40 cm long, ca. 5 mm thick at the base, faint green foliis oblanceolatis fusiformibus), rosulis laxis (vs. to reddish, ascending to pendant. Inflorescence a cin- compactis) pedicellis usque ad 1 mm longis (vs. pedi- cinnus, pink colored, with 13–20 flowers; bracts read- cellis usque ad 37 mm), petalis roseo-purpureis (vs. ily deciduous, ascending, oblanceolate, spurred at the luteo-rubentibus), squama rosea (vs. lutea) differt. base, 10–15 mm long, ca. 5 mm wide, green to pink- green; floral bracts 5–10 mm long and 3–5 mm wide, Plant perennial, suffrutescent, erect, succulent, lanceolate, green to pink-green, base spurred, some- roots fibrous (Fig. 1). Stems much branched near the what pruinose (Fig. 5). Pedicels 5–10 mm long, ca. base, 5–10 cm long and 1–1.5 cm thick at the base 1 mm thick, faint rose colored, farinaceous. Calyx 5 (Fig. 2). Rosette dense, 8–10 cm in diameter (Fig. 3). unequal sepals, joined at the base, segments oblong to Leaves ascending, 4–6 cm long and 1–1.5 cm wide, lanceolate, the smaller ones 4.5–6 mm long and 2–3 oblanceolate, flat to slightly concave, olive-green to mm wide, lanceolate, the larger ones 6–6.5 mm long

142 Author Copy CACTUS AND SUCCULENT JOURNAL 4. Pachyphytum huastecanum leaves. Photo by Jeronimo Reyes.

6. General view of Pachyphytum huastecanum habitat. Photo by Julia Etter & Martin Kristen.

5. Pachyphytum huastecanum inflorescence. Photo by Julia Etter & Martin Kristen.

and 3.5–4.5 mm wide, oblong, pale green, apex acute. Corolla campanulate, petals 5, erect, 10–12 mm long and 4–5 mm wide, apex mucronate, faint rose colored near the base and purplish-pink from the thickest part to the apex, appendices flattened, amorphous, faint rose color. Stamens 10, 5 antisepalous filaments 6.5–7 mm long, 5 antipetalous filaments 3.5–4 mm long, anthers yellow. Nectary scales oblong, ca. 1 mm long, yellowish. Ovary ovoid ca. 6 mm long and 4 mm wide, faint yellow to greenish, style ca. 2 mm long, purple colored, stigma greenish. Fruit, a polyfollicle. Type: Mexico, Nuevo León: municipality of Santa Catarina, Arroyo Potreritos, near El Pajonal, Cumbres de Monterrey National Park, 1690 m. 7 November 7. Pachyphytum huastecanum growing on rocks. 2016. J. Etter & M. Kristen 4185 (Holotype: MEXU!). Photo by Julia Etter & Martin Kristen.

2020 VOLUME 92 NUMBER 2 Author Copy 143 9. Detail of Pachyphytum huastecanum flowers. Photo by Jeronimo Reyes.

8. Detail of Pachyphytum huastecanum flowers. Photo by Jeronimo Reyes.

Phenology: Plants flower from March to April in habitat and in the greenhouse. Distribution and Habitat: Pachyphytum huasteca- num is a micro-endemic species. Because of the rug- ged topography of the area and the resulting difficulty for thorough exploration, it is very possible that its distribution is much wider (Figs. 6 & 7). Associated species are simulans Rose, palmeri S. Watson, Sedum calcicola B.L. Rob. & Greenm., Sedum nanifolium Fröd., Agave aff. ovatifolia G.D. Starr & Villarreal, Opuntia cantabrigiensis Lynch, Echinocere- us viereckii Werderm., Dasylirion berlandieri S.Watson, Tillandsia usneoides (L.) L. Close to the site in the Arroyo Potreritos flows a seasonal creek in a forest of Pachyphytum huastecanum petal interior surface. Quercus sp and Juniperus sp. 10. Photo by Jeronimo Reyes. Etymology: The specific epithet is a reference to the Huasteca region in the state of Nuevo León, Mexico. sepals, these shorter than the corolla (Figs. 8 & 9), and Discussion by the absence of red spots on the inner face of the petals (Fig. 10). Because of leaf shape and the pinkish Pachyphytum huastecanum is placed in sect. Diot- petals, the new species is morphologically related to P. ostemon (Salm-Dyck) E. Walther by having sub-equal fittkaui, P. glutinicaule, and P. hookeri. From P. hookeri,

144 Author Copy CACTUS AND SUCCULENT JOURNAL Table 1. Morphological comparison between Pachyphytum huastecanum and related Pachyphytum species. Character/Species P. huastecanum P. fittkaui P. glutinicaule P. hookeri Habit Usually solitary Usually solitary Usually solitary Usually cespitose Stem length Up to 10 cm (50–70 cm) up to 1 m 30–60 cm Up to 30 cm Rosette Compact Lax Lax Lax Rosette diameter 8–10 cm 10–20 cm 7–12 cm 7–12 cm Obovate, obovate- Lanceolate- Oblanceolate Leaves shape Elliptic-oblanceolate spatulate or oblanceolate to flattened oblanceolate fusiform 4–6 cm long, 1–1.5 cm 3–9.5 cm long, 2–4 cm 3–6.5 cm long, 2–3.5 2.5–5 cm long, 0.6–1.8 Leaves dimensions wide wide cm wide wide Olive-green to bluish- Glaucous, purple Green or glaucous, Leaves color Deep green green tinged reddish at tips Floral stem length 30–40 cm Up to 35 cm Up to 40 cm 5–35 cm Pedicel length 5–10 mm 2–6 mm 4–15 mm 4–37 mm The larger ones 6–6.5 The larger ones 4–8 Sepal length mm, the smaller ones 11–23 mm 10–16 mm mm, the smaller ones 4.5–6 mm 3–7 mm Oblong to oblong- Petal shape Obovate-oblanceolate Deltoid-lanceolate Oblong-oblanceolate oblanceolate, 5.5–15 mm Petal length 10–12 mm 12–17 mm 12–17 mm 5.5–15 mm Scale (appendages) Yellow Pink Deep pink Pink color Ovaries Yellowish White White or cream Yellow Faint yellow to Nectary scale color Yellowish Yellowish Yellow greenish Flowering time March to April February and June March to June Aguascalientes, Guanajuato, San Luis Distribution Nuevo León Hidalgo San Luis Potosí, Potosí Guanajuato Table 2. Molecular diagnostic sites summary for Pachyphytum huastecanum and another 19 Pachyphytum species. Loci rbcL matK ITS2 Taxa/Site 376 50 114 159 198 357 591 624 636 754 820 144 271 296 298 388 439 440 P. brachetii J.Reyes, C C G T T C C G C A A C A G C T C C O.González & A.Gut. P. bracteosum Link, Klotzsch C C G T A C C G C A A A A G C C C C & Otto P. brevifolium Rose C C G T A C C A C A A C A G C T C C P. caesium Kimnach & Moran C C G T A C C G C A A C A G C T C C P. compactum Rose C G G T A C T G C A A C A G C T C C P. contrerasii Pérez-Calix, C C G T A C C G C A A C A G C T C C I.García & Cházaro P. fittkaui Moran C C G T A C C G C A A C A G C T C C P. garciae Pérez-Calix & Glass C C G T A C C G C A A C A G C T C C P. glutinicaule Moran C C A G A C C G C A - C A G C T C C P. hookeri A.Berger G C G T A C C G C A A C A G C T C C P. huastecanum J.Reyes, Etter C C G T A C C G C A A C G C C ? ? ? & Kristen P. kimnachii Moran C C G T A C C G C A A C A G C T C C P. longifolium Rose C C G T A C C G C A A C A G C T C C P. machucae I.García, Glass & C C G T A C C G C A A C A G C T C C Cházaro P. oviferum J.A.Purpus C C G T A C C G T A A C A G C T C C P. rzedowskii I.García, Pérez- C C G T A C C G C A A C A G C T C A Calix & J.Meyrán P. saltense Brachet, Reyes & C C G T A C C G C A A C A G C T C C Mondragón P. viride E.Walther C C G T A A C G C A A C A G C T C C P. viscidum J.Reyes & L.E.Cruz- C C G T A C C G C G A C A G T T T C López P. werdermannii Poelln. C C G T A C C G C A A C A G C T C C

2020 VOLUME 92 NUMBER 2 Author Copy 145 Map 1. Distribution of Pachyphytum huastecanum in the municipality of Santa Catarina.

the new species can be differentiated by its thinner the rest of the Pachyphytum species sampled (Tables and flat vs. cylindric-fusiform leaves, the color of the 2 & 3). Additionally, the new species is geograph- leaves (olive-green vs. green, bluish-green to bluish- ically isolated by the mountain ranges of the Sierra gray), pale green and wider vs green to red sepals, by Madre Oriental becoming the northernmost species its strongly campanulate, wider vs tubular to slightly known to date, and is only known in the municipal- campanulate flowers, and by its shorter (10–12 mm vs ity of Santa Catarina (Map 1). Specifically, it is found 8–15 mm) obovate vs. oblanceolate petals. Pachyphy- in the Parque Nacional Cumbres de Monterrey which tum huastecanum can be differentiated from P. glutini- encompasses more than 183,000 hectares, meaning caule because its stems are not glutinous and by hav- that the species is legally protected as it occurs in a ing olive-green vs. grayish-blue colored leaves. Finally, National Protected Area (Áreas Nacionales Protegi- the new species can be distinguished from P. fittkaui das) (Comisión Nacional de Áreas Naturales Prote- by its shorter stems and leaves (Table 1), its shorter gidas, 2006). and wider sepals vs. narrowed as long as the corolla sepals (sepals to petals ratio: Moran, 1971), and by its Acknowledgments obovate vs. deltoid-lanceolate petals. From a DNA barcoding character-based perspec- The authors would like to thank the Sociedad tive (DeSalle et al., 2005; Goldstein & DeSalle, 2011), Mexicana de Cactología, A.C. for the arrangement P. huastecanum can be distinguished from other 19 of the federal collecting permit: Oficio Núm. SGPA/ Pachyphytum species — including P. fittkaui, P. glutini- DGVS/02365/17, as well as the National Laborato- caule, and P. hookeri — by the presence of the nucleo- ry of Biodiversity (Laboratorio Nacional de Biodiver- tides G (Guanine) and C (Cytosine) at the sites 271 sidad (LANABIO) of the National Council of Sci- and 296, respectively, of the ITS2 genome region, in ences and Technologies ((Consejo Nacional de Cien- contrast to the A and G present in the sequences of cia y Tecnología (CONACYT)). We also express our

146 Author Copy CACTUS AND SUCCULENT JOURNAL Table 3. GenBank accession numbers for the taxa sampled. Taxa/locus rbcL matK ITS2 BOLD ID P. brachetii J.Reyes, O.González & A.Gut. MG220464.1 MG220520.1 MG217180.1 JE-4350 P. bracteosum Link, Klotzsch & Otto MG220447.1 MG220503.1 MG217163.1 JE-6275 P. brevifolium Rose MG220451.1 MG220507.1 MG217167.1 JE-5048 P. caesium Kimnach & Moran MG220475.1 MG220531.1 MG217191.1 JE-5185 P. compactum Rose MG521706.1 MG521586.1 MG521432.1 JE-5273 P. contrerasii Pérez-Calix, I.García & Cházaro MG521760.1 MG521636.1 MG521484.1 JE-5627 P. fittkaui Moran MG521807.1 MG521675.1 MG521528.1 JE-5237 P. garciae Pérez-Calix & Glass MG521764.1 MG521639.1 MG521488.1 JE-5885 P. glutinicaule Moran MG521736.1 MG521613.1 MG521461.1 JE-5653 P. hookeri A.Berger MG521819.1 MG521687.1 MG521539.1 JE-5036 P. huastecanum J.Reyes, Etter & Kristen Pending Pending Pending PH-01 P. kimnachii Moran MG521809.1 MG521677.1 MG521530.1 JE-5305 P. longifolium Rose MG521693.1 MG521573.1 MG521419.1 JE-6276 P. machucae I.García, Glass & Cházaro MG521768.1 MG521643.1 MG521492.1 JE-5071 P. oviferum J.A.Purpus MG521718.1 MG521597.1 MG521444.1 JE-6038 P. rzedowskii I.García, Pérez-Calix & J.Meyrán MG521770.1 MG521645.1 MG521494.1 JE-5562 P. saltense Brachet, J.Reyes & Mondragón MG521799.1 MG521670.1 MG521521.1 JE-3644 P. viscidum J.Reyes & L.E.Cruz-López MG521795.1 MG521666.1 MG521518.1 JE-5043 P. viride E.Walther MG521815.1 MG521683.1 MG521535.1 JE-5873 P. werdermannii Poelln. MG521785.1 MG521657.1 MG521509.1 JE-5451

Additional specimens examined (all from México): P. brachetii. Hidalgo: mpio. Actopan, 5 km al sur de Actopan, J. Reyes 4350 (MEXU). P. bracteosum. Hidalgo: Laguna de Meztitlán, J. Reyes 6275 (MEXU). P. brevi- folium. Guanajuato: Los Picones, J. Reyes 5048 (MEXU). P. caesium. Aguascalientes: Barranca Montoro, J. Reyes 5185 (MEXU). P. compactum. Querétaro: La Cañada, J. Reyes 5273 (MEXU). P. contrerasii. Jalisco: Mirador Dr. Atl, J. Reyes 5627 (MEXU). P. fittkaui. Guanajuato: Los Jofres, J. Reyes 5237 (MEXU). P. garciae. Querétaro: El Zapote, J. Reyes 5885 (MEXU). P. glutinicaule. Hidalgo: mpio. Tasquillo, Puerto Juárez, J. Reyes 5653 (MEXU). P. hookeri. Guanajuato: Cerro Gordo, J. Reyes 5036 (MEXU). P. kimnachii. San Luis Potosí: Cerro Agujón, J. Reyes 5305 (MEXU). P. longifolium. Hidalgo: entre Gilo y Almolón, J. Reyes 6276 (MEXU). P. machucae. Micho- acán: Cañada de Agua, J. Reyes 5071 (MEXU). P. oviferum. San Luis Potosí: El Cajón, J. Reyes 6038 (MEXU). P. rzedowskii. Michoacán: 1.5 km al noroeste de Puente Grande, J. Reyes 5562 (MEXU). P. saltense. Zacatecas: El Salto, J. Reyes 3644 (MEXU). P. viscidum. (paratypes). Aguascalientes: mpio. Tepezala, in cliffs above RMO Tepezala, 2421 m, June 16, 2012, Julia Etter & Martin Kristen 3619 (MEXU). Jalisco: mpio. Ojuelos de Jalisco, 3 km NW of Matanzas, between Lagos de Moreno and Ojuelos along highway 80, in cliffs behind the cem- etery, 2320 m, March 27, 2012 (flowering), Julia Etter & Martin Kristen 3494 (MEXU). Zacatecas: mpio. Villa García, 4.5 km from Villa García along dirt road to Loreto, 2150 m, May 15, 2012, Julia Etter & Martin Kristen 3531 (MEXU). P. viride. Guanajuato: entrada al Pueblo de Coconoxtle, J. Reyes 5873 (MEXU). P. werdermannii. Tamaulipas: Microondas-Las Mulas, Julia Etter & Martin Kristen 2000.

Meyrán, G.J. & Chávez, L.L (2003). Las Crasuláceas de México. Sociedad gratitude to Omar González Zorzano for the Latin Mexicana de Cactología, A.C., México, Ciudad de México. 234 pp. diagnosis. Julia Etter and Martin Kristen would also Moran R (1971). Pachyphytum fittkaui, a new species from Guanajuato, Mexico. Cactus & Succulent Journal of America 43: 26–32. like to thank Michael Bechtold and Wolfgang Metorn Reyes S, J., de la Cruz-López, L. E., Kuzmina, M., & Vergara-Silva, F (2019). for their patient companionship on several field trips. Morphological and molecular diagnostic characters reveal a new species of Pachyphytum (Crassulaceae). Haseltonia 26: 14–22. Literature cited Comisión Nacional de Áreas Naturales Protegidas (2006). Programa de Conservación y Manejo: Parque Nacional Cumbre de Monterrey. 178 pp. DeSalle R, Egan, M. G. & Siddall, M (2005). The unholy trinity: , species delimitation and DNA bar­coding. Philosophical Transactions of the Royal Society B 360: 1905–1916. Goldstein, P. Z. & DeSalle, R (2011). Integrating DNA barcode data and taxonomic practice: determination, discovery, and description. Bioessays 33: 135–147.

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