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Species Diversity and Plastid Dna Haplotype Distributions of Pinus Subsection Australes (Pinaceae) in Guerrero and Oaxaca

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Species Diversity and Plastid Dna Haplotype Distributions of Pinus Subsection Australes (Pinaceae) in Guerrero and Oaxaca ARTÍCULO ORIGINAL D.R. © TIP Revista Especializada en Ciencias Químico-Biológicas, 19(2):92-101, 2016 SPECIES DIVERSITY AND PLASTID DNA HAPLOTYPE DISTRIBUTIONS OF PINUS SUBSECTION AUSTRALES (PINACEAE) IN GUERRERO AND OAXACA Alfredo Ortiz-Martíneza and David S. Gernandtb aInstituto Tecnológico de Cd. Altamirano, Av. Pungarabato Poniente s/n, Col. Morelos, Cd. Altamirano, C.P. 40660, Guerrero, México. bDepartamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México. Ciudad Universitaria, Deleg. Coyoacán, C.P. 04510, Ciudad de México, México. E-mail: [email protected] ABSTRACT Pinus subsection Australes is a group of North American hard pines comprising approximately 29 ecologically and economically important tree species distributed throughout North and Central America and the Caribbean Islands. Previous studies have shown that some species of this subsection share plastid DNA haplotypes, a pattern that is attributed to introgressive hybridization or the retention of ancestral polymorphisms. Here we describe the morphological and plastid haplotype diversity for this group of species in the states of Guerrero and Oaxaca, Mexico. Seven species of Pinus subsection Australes are recognized in the study area, one of which, P. patula, includes two varieties. Seven variable sites and nine haplotypes were found in an 840 b.p. fragment of the DNA coding region ycf1. Shared haplotypes were found for P. patula var. patula, P. patula var. longipedunculata, P. herrerae, and P. tecunumanii. Four of the nine haplotypes found were restricted to Oaxaca. Although plastid DNA genealogies are valuable for studying evolution in this group, greater sampling of individuals and the inclusion of more variable sites are needed to more accurately infer species relationships. Key Words:ELRGLYHUVLW\JHQHÀRZOLQHDJHVRUWLQJSLQHVVSHFLHVGHOLPLWDWLRQ Diversidad de especies y distribución de haplotipos de ADN del plastidio de Pinus subsección Australes(Pinaceae) en Guerrero y Oaxaca RESUMEN Pinus subsección Australes es un grupo de pinos duros de América del Norte que comprende aproximadamente 29 especies de árboles importantes económicamente y ecológicamente distribuidos a lo largo de toda América del Norte y Central y las Islas Caribeñas. Estudios previos han mostrado que las especies de esta subsección a menudo comparten haplotipos de ADN de plastidio, un patrón que es atribuido a la hibridación LQWURJUHVLYD\ODUHWHQFLyQGHSROLPRUÀVPRVDQFHVWUDOHV$TXtGHVFULELPRVODGLYHUVLGDGGHKDSORWLSRVGH SODVWLGLR\ODPRUIRORJtDSDUDHVWHJUXSRGHHVSHFLHVHQORVHVWDGRVGH*XHUUHUR\2D[DFD0p[LFR6LHWH especies de Pinus subsección Australes son reconocidas en el área de estudio, una de las cuales, P. patula LQFOX\HGRVYDULHGDGHV6LHWHVLWLRVYDULDEOHV\QXHYHKDSORWLSRVIXHURQHQFRQWUDGRVDPSOLÀFDQGRXQIUDJPHQWR GHSEGH$'1GHODUHJLyQFRGLÀFDQWHycf1. Se encontraron haplotipos compartidos para P. patula var. patula, P. patula var. longipedunculata, P. herrerae y P. tecunumanii. Cuatro de los nueve haplotipos HQFRQWUDGRVHVWiQUHVWULQJLGRVD2D[DFD$XQTXHODVJHQHDORJtDVGHJHQHVVRQYDOLRVDVSDUDHVWXGLDUOD evolución de este grupo, se requieren mayor muestreo de individuos y más sitios variables para la inferencia de relaciones entre las especies. Palabras Clave:ELRGLYHUVLGDGÀXMRJpQLFRVRUWHRGHOLQDMHVSLQRVGHOLPLWDFLyQGHHVSHFLHV Nota: Artículo recibido el 14 de septiembre de 2015 y aceptado el 27 de abril de 2016. agosto, 2016 Ortiz-Martínez, A. & Gernandt, D.S.: Pines of Guerrero and Oaxaca 93 INTRODUCTION 3ODVWLG HJFKORURSODVW '1$KDVEHHQZLGHO\VWXGLHGLQSLQHV inus 3LQDFHDH DUJXDEO\WKHPRVWHFRORJLFDOO\DQG ,QFRQWUDVWWRDQJLRVSHUPVSODVWLG'1$LVSDWHUQDOO\LQKHULWHG HFRQRPLFDOO\ LPSRUWDQW WUHH JHQXV LQ WKH ZRUOG1, LQFRQLIHUV25,WH[KLELWVORZJHQHWLFGL൵HUHQWLDWLRQLQFRQLIHU P FRPSULVHV DSSUR[LPDWHO\ VSHFLHV GLVWULEXWHG SRSXODWLRQVDQGVSHFLHVUHODWLYHWRPLWRFKRQGULDORUQXFOHDU QDWXUDOO\LQWHUUHVWULDOHQYLURQPHQWVWKURXJKRXWWKH '1$WKLVLVEHFDXVHSODVWLGJHQHÀRZLVPHGLDWHGE\ERWK 1RUWKHUQ+HPLVSKHUH2-53LQHVDUHHDVLO\UHFRJQL]HGE\WKHLU VHHGDQGSROOHQLQFRQLIHUV262QHUHVXOWLVWKDWVSHFLHVWDNHD QHHGOHOLNHOHDYHVDUUDQJHGLQIDVFLFOHVRI± DOWKRXJKP. ORQJWLPHWREHFRPH¿[HGIRUDVLQJOHSODVWLG'1$OLQHDJH monophylla7RUU )UpPDQGP. californiarum '.%DLOH\ DQGVKDULQJRISODVWLGOLQHDJHVE\FORVHO\UHODWHGVSHFLHVLV KDYHVROLWDU\QHHGOHV DQGZRRG\VHHGFRQHVZLWKWKLFNHQHG RIWHQREVHUYHG VFDOH DSLFHV DQG D GRUVDO RU WHUPLQDO XPER D VSHFLDOL]HG UDLVHGUHJLRQUHVXOWLQJIURPPRUHWKDQRQHVHDVRQRIJURZWK :HFKRVHDVRXUVWXG\DUHDWKHSROLWLFDOERXQGDULHVPDUNHGE\ 0H[LFRLVWKHFHQWHURIVSHFLHVGLYHUVLW\IRUSLQHVZLWKPRUH *XHUUHURDQG2D[DFD7KHVHVWDWHVDUHORFDWHGRQWKHVRXWKHUQ WKDQRIWKHZRUOG¶VVSHFLHVRFFXUULQJQDWXUDOO\ZLWKLQLWV 3DFL¿FFRDVWRI0H[LFR*XHUUHURRFFXSLHVNP2DQGLV borders2-77KHQXPEHURIVSHFLHVUHFRJQL]HGIRUWKHFRXQWU\ WKHIRXUWHHQWKODUJHVWVWDWHLQWKHFRXQWU\DQG2D[DFDRFFXSLHV KDVYDULHGDPRQJUHFHQWDXWKRUV(FNHQZDOGHU2UHFRJQL]HG NP2DQGLVWKH¿IWKODUJHVW7KHSULQFLSDOPRXQWDLQ VSHFLHV)DUMRQUHFRJQL]HGDQG*HUQDQGWDQG3pUH]GHOD UDQJHLQWKHVHVWDWHVLVWKH6LHUUD0DGUHGHO6XUZKLFKH[WHQGV 5RVD7UHFRJQL]HG WKURXJK -DOLVFR &ROLPD VRXWKHUQ 0LFKRDFiQ WKH 6WDWH RI 0H[LFR0RUHORV3XHEOD*XHUUHURDQG2D[DFDDQGLVERUGHUHG Pinus LV FODVVL¿HG LQ WZR VXEJHQHUD IRXU VHFWLRQV DQG WRWKHQRUWKE\WKH7UDQVYHUVH0H[LFDQ9ROFDQLF%HOW%RWK VXEVHFWLRQVPinusVXEVHFWLRQ AustralesLVWKHPRVWVSHFLHVULFK *XHUUHURDQG2D[DFDDUHKLJKO\GLYHUVHLQVSHFLHV)RUH[DPSOH FRPSULVLQJDSSUR[LPDWHO\VSHFLHVGLVWULEXWHGWKURXJKRXW VSHFLHVRISODQWVKDYHEHHQUHSRUWHGIRU2D[DFDPRUH 1RUWK DQG &HQWUDO$PHULFD DQG WKH ZHVWHUQPRVW LVODQGV RI WKDQDQ\RWKHUVWDWHLQWKHFRXQWU\)RXUWHHQSLQHVSHFLHVKDYH WKH&DULEEHDQH[FHSW-DPDLFD,WLQFOXGHVWKHVRXWKHUQ\HOORZ EHHQUHFRJQL]HGIRU*XHUUHURDQGIRU2D[DFD6HYHQRI SLQHV WKH HJJFRQH SLQHV DQG WKH &DOLIRUQLD FORVHGFRQH WKHVHVSHFLHVEHORQJWRPinusVXEVHFWLRQAustrales SLQHVDQGLVFODVVL¿HGWRJHWKHUZLWKVXEVHFWLRQVContortae, DQGPonderosae in PinusVXEJHQXVPinusVHFWLRQTrifoliaeD 7KLVLVWKH¿UVWFRPSDUDWLYHVWXG\WRLQWHJUDWHPRUSKRORJLFDO JURXSNQRZQLQIRUPDOO\DVWKH1RUWK$PHULFDQKDUGSLQHV DQDWRPLFDODQGPROHFXODUFKDUDFWHUVLQSLQHVRI*XHUUHURDQG $OOVSHFLHVRIPinusVXEVHFWLRQAustralesKDYHWZRYDVFXODU 2D[DFD0H[LFR2XUREMHFWLYHVDUHWREHWWHUXQGHUVWDQGWKH EXQGOHVLQWKHLUOHDYHVDQGPRVWKDYHSHUVLVWHQWIDVFLFOHVKHDWKV PRUSKRORJLFDOGLYHUVLW\RIPinusVXEVHFWLRQAustralesLQWKHVH $OOKDYHZRRG\VHHGFRQHVDQGVRPHKDYHVWURQJO\GHYHORSHG WZRVWDWHVDQGWRIXUWKHUGRFXPHQWZKLFKVSHFLHVLQWKLVJURXS FRQHVFDOHDSRSK\VHVDQGXPERVRIWHQZLWKDPXFUR7KHLU VKDUHSODVWLG'1$KDSORW\SHV VHHGVKDYHDQDUWLFXODWHZLQJDQGDUHSULPDULO\ZLQGGLVSHUVHG MATERIALS AND METHODS 1XPHURXVWD[RQRPLFZRUNVKDYHWUHDWHG PinusLQFOXGLQJ 'ULHGVSHFLPHQVZHUHREWDLQHGIURPWKH,QYHQWDULR1DFLRQDO UHJLRQDOÀRUDVWKDWLQFOXGH0H[LFR0RUSKRORJ\DORQHKDV )RUHVWDO DQG WKH 1DWLRQDO +HUEDULXP 0(;8 1LQHW\ QRWEHHQVX൶FLHQWWRUHVROYHWKHWD[RQRPLFTXHVWLRQVVXUURXQGLQJ LQGLYLGXDOV ZHUH H[DPLQHG PRUSKRORJLFDOO\ )LJXUH WKHJHQXV'1$VHTXHQFHVKDYHEHHQYHU\H൵HFWLYHDWFRQ¿UPLQJ $SSHQGL[ 0RUSKRORJLFDO PHDVXUHPHQWV RI QHHGOHV RUUHMHFWLQJWKHPDMRUSLQHOLQHDJHVWKDWZHUHRULJLQDOO\SURSRVHG VHFRQGDU\OHDYHV IDVFLFOHVKHDWKVVHHGFRQHVDQGSHGXQFOHV EDVHGSULPDULO\RQPRUSKRORJ\DUWL¿FLDOFURVVHVDQGELRFKHPLFDO ZHUHWDNHQIURPGULHGFROOHFWLRQV$OOPHDVXUHPHQWVZHUH VWXGLHV +RZHYHU QDWXUDO DQG DUWL¿FLDO K\EULGL]DWLRQ DUH PDGHZLWKDUXOHUDQGH[SUHVVHGLQFP)RUQHHGOHDQGIDVFLFOH UHODWLYHO\ZHOOGRFXPHQWHGLQSLQHV16%RWKK\EULGL]DWLRQDQG VKHDWK PHDVXUHPHQWV DUELWUDULO\ FKRVHQ IDVFLFOHV ZHUH UHWHQWLRQ RI DQFHVWUDO DOOHOHV KDYH REVFXUHG WKH SK\ORJHQHWLF PHDVXUHG6HHGFRQHDQGSHGXQFOHPHDVXUHPHQWVZHUHEDVHG UHODWLRQVKLSVDPRQJFORVHO\UHODWHGVSHFLHV RQDVLQJOHPDWXUHFRQHSHULQGLYLGXDO )LJXUH 7UDQVYHUVH VHFWLRQVZHUHPDGHIURPWKHPHGLDOSDUWRIOHDYHV¿[HGLQ ,QPRVWRUJDQLVPVVSHFLDWLRQERWKJHQHUDWHVELRORJLFDOGLYHUVLW\ IRUPDOGHK\GHDFHWLFDFLGDOFRKRO )$$ DQGH[DPLQHGZLWK DQGJLYHVULVHWRJHQHJHQHDORJLHVWKDWUHÀ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ev.Esp.Cienc.Quím.Biol. Vol. 19, No. 2 Figure 1. The collection localities for the individuals included in this study. 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