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Morphological and Molecular Diagnostic Characters Reveal a New Species of Pachyphytum(Crassulaceae)

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Morphological and Molecular Diagnostic Characters Reveal a New Species of Pachyphytum(Crassulaceae) 14Haseltonia 26: 14–22. 2019 Haseltonia 26: 14–22. 201914 Morphological and molecular diagnostic characters reveal a new species of Pachyphytum (Crassulaceae) Jerónimo Reyes Santiago1, Luis E. de La Cruz-López2*, Maria Kuzmina3, & Francisco Ver- gara-Silva1* 1 Jardín Botánico, Instituto de Biología, Universidad Nacional Autónoma de México. Circuito exterior s/n, Ciudad Universitaria, Copilco, Coyoacán, C.P. 04510, Ciudad de México, Mexico. 2 Posgrado en Ciencias Biológicas, Instituto de Biología, Universidad Nacional Autónoma de México. Av. Ciudad Universitaria 3000, C.P. 04360, Coyoacán, Ciudad de México, Mexico. 3 Centre for Biodiversity & Genomics, Biodiversity Institute of Ontario. University of Guelph, ON N1G 2W1, Ontario, Canada. * Authors for correspondence: [email protected]; [email protected] Manuscript received 26th October 2018 Abstract: Pachyphytum viscidum (Crassulaceae: Saxifragales) is described as a new species to science. The speci- mens analyzed come from the Sierra de Santa Bárbara, Guanajuato, Mexico. Some morphological diagnostic characters for the new species are: sticky stems and leaves; dark green leaves; absence of wax on all their or- gans; and pale pink flowers. The new species is placed in the section Diotostemon of the genus, with pos- sible proximity to P. brevifolium and P. hookeri. DNA sequence analysis from the matK and rbcL chloroplast regions, as well as ITS2 nuclear loci, revealed the presence of molecular diagnostic sites which, in addition to the aforementioned morphological character states, further support the taxonomic description as well as future integrative taxonomy studies in Mexican Crassulaceae. Resumen: Se describe a Pachyphytum viscidum (Crassulaceae: Saxifragales) como una especie nueva para la ciencia. Los especímenes analizados provienen de la Sierra de Santa Bárbara, Guanajuato, México. Algunos caracteres morfológicos diagnósticos para esta especie son: sus tallos y hojas glutinosas; hojas verde oscuro; la ausencia de cera en sus órganos; y el color rosa tenue de sus flores. Se le ubica dentro de la sección Diotoste- mon, en posible cercanía con P. brevifolium y P. hookeri. El análisis de secuencias de ADN de las regiones del cloroplasto matK y rbcL, así como del locus nuclear ITS2, reveló la presencia de sitios diagnósticos molecula- res que, en adición a los morfológicos, también apoyan la descripción taxonómica así como estudios futuros de taxonomía integrativa en Crassulaceae de México. Keywords: Crassulaceae, character-based DNA barcoding, integrative taxonomy, ITS2, matK, Pachyphytum INTRODUCTION Moran´s nor Thiede´s groupings match García- Ruíz´s results. The genus Pachyphytum (Crassulaceae) was pro- A recent molecular phylogenetic analysis of the posed by Link, Klotzsch et Otto (Klotzsch, 1841) genus Echeveria and related genera including Pachy- with P. bracteosum Klotzsch (Moran, 1989) as the phytum, based on a combined DNA matrix of rbcL, type species. Moran (1968) recognized three sec- matK and ITS2 genomic regions, retrieved the lat- tions based on morphological affinities: Pachyphytum, ter genus as a monophyletic group (Cruz-López et Diotostemon and Ixiocaulon. Later, J. Thiede (in Eggli, al., 2019) sister to the rest of Cremnophila, Echeve- 2003) merged the species of sect. Ixiocaulon into sect. ria, Graptopetalum, and some Sedum and Thompson- Pachyphytum, resulting in only two sections: Diotoste- ella species. Another significant result of this work mon and Pachyphytum. Genetic distance-based clas- indicated that Pachyphytum cuicatecanum (J.Reyes, sification of the genus employing AFLP molecular Joel Pérez & Brachet) Kimnach is closely related to markers suggested two groups inside Pachyphytum several Echeveria species (E. penduliflora and oth- (García-Ruíz 2003). In this classification, group A ers) and clearly do not belong to Pachyphytum, de- included species P. bracteosum, P. brevifolium, P. glu- spite sharing many morphological character states. tinicaule, P. hookeri, P. longifolium, and P. oviferum; Nevertheless, low resolution problems do not allow while group B incorporated P. caesium, P. coeruleum, the identification of groups (sections) inside Pachy- P. compactum, P. contrerasii, P. fittkaui, P. garciae, P. phytum. Previous studies on the “Echeveria Group” kimnachii, P. machucae, P. rzedowskii, P. viride, and (Carrillo-Reyes et al., 2009; Vázquez-Cotero et al., P. werdermannii. It should be noted that neither 2017) also recovered Pachyphytum as a monophyletic Author Copy HASELTONIA 26. 2019 15 Table 1. 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. 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 sp. nov. 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 group, resting on partial species sampling. Histori- this national collection program, botanical explora- cally, the placement of taxa in infrageneric catego- tions were necessary, as well as the cultivation and ries (sections) has been complicated, because there maintenance of species of different genera with a is often little congruence between morphological, geo-referenced database. During fieldwork, the genus biogeographic and DNA datasets; this discrepancy Pachyphytum received primary attention. The work between different sources of evidence may have con- with the collection has received a great impulse from trasting explanations (e.g. recent diversification vs. Christian Brachet, member of the Mexican Cactus decoupling between molecular and morphological Society (Sociedad Mexicana de Cactología, A.C.), evolution). who has provided significant help with the logistics The most outstanding character states of Pachy- of the explorations and with finding information phytum are: presence of membranous scale-like struc- about the type localities of the known species. tures known as appendages on the inside of every The present paper derives from the “Crassulaceae petal, one on each side of the epipetalous filaments of Mexico” (“Crasuláceas de México”) long-term re- although, some Echeveria species also possess ap- search project, based on the National Crassulaceae pendages (E. heterosepala, E. longissima var. longis- Collection and hosted by JB-IBUNAM. A primary sima, E. dactylifera, E. purhepecha, E. rulfiana, among aim of this project has been the gradual construction many others); pronounced succulence of the leaves; of a DNA barcoding reference library for Mexican and pendant scorpioid cymose inflorescences (Reyes Crassulaceae, to be linked with other datasets in an et al., 2007). At present, the genus consists of 24 integrative taxonomy (DeSalle et al., 2005; Gold- described species — based on morphological, phe- stein and DeSalle, 2011) framework. Character- nological, ecological and geographical distribution based methods, such as CAOS (Sarkar et al., 2008), aspects — all endemic to Mexico and distributed have been given preference to establish DNA bar- mainly in central and western regions of this country. codes, as they allow the definition of discrete DNA In 2003, a collection of living plants of the fam- diagnostics compatible with apomorphic, morpho- ily Crassulaceae (Saxifragales: Rosidae) was started at logical character states. the Jardín Botánico, Instituto de Biología, Univer- The species described here was first located near sidad Nacional Autónoma de México. In 2005, the La Presita at the base of the Sierra de Santa Bárbara, Mexican Association of Botanical Gardens (Aso- Guanajuato, in May of 2004 by the team of Chris- ciación Mexicana de Jardines Botánicos) declared tian Brachet, Joel Pérez Crisanto, Roxana Mon- this collection as “Colección Nacional” (National dragón, and Jerónimo Reyes Santiago. The discovery Collection), representing 70% of the Crassulaceae was made during a trip to locate populations of Ech- species known in Mexico. For the establishment of everia calderoniae Perez-Calix growing in the same Author Copy 16 REYES SANTIAGO ET AL. — DIAGNOSTICS OF A NEW PACHYPHYTUM Table 2. Primer sequences and PCR thermocycler programs used to amplify DNA fragments. Primer Sequence Program rbcLa-F ATGTCACCACAAACAGAGACTAAAGC 94°C for 4 min; 35 cycles of 94°C for 30 sec, 55°C for 30 sec, rbcLa-R GTAAAATCAAGTCCACCRCG 72°C for 1 min; final extension 72°C for 10 min. MatK-1RKIM-f ACCCAGTCCATCTGGAAATCTTGGTTC 94°C for 1 min; 35 cycles of 94°C for 30 sec, 52°C for 20 sec, MatK-3FKIM-r CGTACAGTACTTTTGTGTTTACGAG 72°C for 50 sec; final extension 72°C for 5 min. ITS2-S2F ATGCGATACTTGGTGTGAAT 94°C for 5 min; 35 cycles of 94°C for 30 sec, 56°C for 30 sec, ITS4 TCCTCCGCTTATTGATATGC 72°C for 45 sec; final extension 72°C for 10 min. Table 3. Molecular diagnostic sites summary for Pachyphytum viscidum and the sampled species of Pachyphytum. Loci rbcL matK ITS2 Taxa/Site 376 50 114 159 198 357 591 624 636 754 820 144 298 388 439 440 P. brachetii C C G T T C C G C A A C C T C C P. bracteosum C C G T A C C G C A A A C C C C P.
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