SPECIES DIVERSITY AND PLASTID DNA HAPLOTYPE DISTRIBUTIONS OF PINUS SUBSECTION AUSTRALES (PINACEAE) IN GUERRERO AND OAXACA | TIP. Revista Especializada en Ciencias Químico-Biológicas

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Pair of bars represents assays with different times of exposition to the treatments (24 and 48hours). Results are given as mean ± SD of assays performed in triplicate. Asterisk indicates statistically significant differences (**p value < 0.01) with respect to untreated cells." ] ] ] "autores" => array:1 [ 0 => array:2 [ "autoresLista" => "Dary Luz Mendoza-Meza, Pierine España-Puccini" "autores" => array:2 [ 0 => array:2 [ "nombre" => "Dary Luz" "apellidos" => "Mendoza-Meza" ] 1 => array:2 [ "nombre" => "Pierine" "apellidos" => "España-Puccini" ] ] ] ] ] "idiomaDefecto" => "en" "EPUB" => "https://multimedia.elsevier.es/PublicationsMultimediaV1/item/epub/S1405888X16300018?idApp=UINPBA00004N" "url" => "/1405888X/0000001900000002/v1_201607220355/S1405888X16300018/v1_201607220355/en/main.assets" ] "en" => array:21 [ "idiomaDefecto" => true "cabecera" => "ARTÍCULO ORIGINAL" "titulo" => "SPECIES DIVERSITY AND PLASTID DNA HAPLOTYPE DISTRIBUTIONS OF PINUS SUBSECTION AUSTRALES (PINACEAE) IN GUERRERO AND OAXACA" "tieneTextoCompleto" => true "paginas" => array:1 [ 0 => array:2 [ "paginaInicial" => "92" "paginaFinal" => "101" ] ] "autores" => array:1 [ 0 => array:4 [ "autoresLista" => "Alfredo Ortiz-Martínez, David S. Gernandt" "autores" => array:2 [ 0 => array:3 [ "nombre" => "Alfredo" "apellidos" => "Ortiz-Martínez" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "a" "identificador" => "aff0005" ] ] ] 1 => array:4 [ "nombre" => "David S." "apellidos" => "Gernandt" "email" => array:1 [ 0 => "dgernandt@ib.unam.mx" ] "referencia" => array:2 [ 0 => array:2 [ "etiqueta" => "b" "identificador" => "aff0010" ] 1 => array:2 [ "etiqueta" => "*" "identificador" => "cor0005" ] ] ] ] "afiliaciones" => array:2 [ 0 => array:3 [ "entidad" => "Instituto Tecnológico de Cd. Altamirano, Av. Pungarabato Poniente s/n, Col. Morelos, Cd. Altamirano, C.P. 40660, Guerrero, México" "etiqueta" => "a" "identificador" => "aff0005" ] 1 => array:3 [ "entidad" => "Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México. Ciudad Universitaria, Deleg. Coyoacán, C.P. 04510, Ciudad de México, México" "etiqueta" => "b" "identificador" => "aff0010" ] ] "correspondencia" => array:1 [ 0 => array:3 [ "identificador" => "cor0005" "etiqueta" => "⁎" "correspondencia" => "Corresponding author." ] ] ] ] "titulosAlternativos" => array:1 [ "es" => array:1 [ "titulo" => "Diversidad de especies y distribución de haplotipos de ADN del plastidio de Pinus subsección Australes(Pinaceae) en Guerrero y Oaxaca" ] ] "resumenGrafico" => array:2 [ "original" => 0 "multimedia" => array:7 [ "identificador" => "fig0015" "etiqueta" => "Figure 3" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr3.jpeg" "Alto" => 1043 "Ancho" => 2148 "Tamanyo" => 166660 ] ] "descripcion" => array:1 [ "en" => "Plastid DNA haplotype network for the study species. The area of the circles is proportional to haplotype frequency." ] ] ] "textoCompleto" => "INTRODUCTIONPinus (Pinaceae), arguably the most ecologically and economically important tree genus in the world<a class="elsevierStyleCrossRef" href="#bib0005">1</a>, comprises approximately 110 species distributed naturally in terrestrial environments throughout the Northern Hemisphere<a class="elsevierStyleCrossRefs" href="#bib0010">2–5</a>. Pines are easily recognized by their needle-like leaves arranged in fascicles of 2–8 (although P. monophylla Torr. & Frém. and P. californiarum D.K. Bailey have solitary needles), and woody seed cones with thickened scale apices and a dorsal or terminal umbo (a specialized raised region resulting from more than one season of growth). Mexico is the center of species diversity for pines, with more than 40% of the world's species occurring naturally within its borders<a class="elsevierStyleCrossRefs" href="#bib0010">2–7</a>. The number of species recognized for the country has varied among recent authors; Eckenwalder<a class="elsevierStyleCrossRef" href="#bib0010">2</a> recognized 39 species, Farjon<a class="elsevierStyleCrossRef" href="#bib0015">3</a> recognized 44, and Gernandt and Pérez-de la Rosa<a class="elsevierStyleCrossRef" href="#bib0035">7</a> recognized 49.Pinus is classified in two subgenera, four sections, and 11 subsections<a class="elsevierStyleCrossRef" href="#bib0040">8</a>. Pinus subsection Australes is the most species rich, comprising approximately 29 species distributed throughout North and Central America and the westernmost islands of the Caribbean except Jamaica. It includes the southern yellow pines, the egg-cone pines, and the California closed-cone pines, and is classified together with subsections Contortae, and Ponderosae in Pinus subgenus Pinus section Trifoliae, a group known informally as the North American hard pines<a class="elsevierStyleCrossRef" href="#bib0045">9</a>. All species of Pinus subsection Australes have two vascular bundles in their leaves, and most have persistent fascicle sheaths. All have woody seed cones, and some have strongly developed cone scale apophyses and umbos, often with a mucro. Their seeds have an articulate wing and are primarily wind-dispersed.Numerous taxonomic works have treated Pinus<a class="elsevierStyleCrossRefs" href="#bib0010">2,9–11</a>, including regional floras that include Mexico<a class="elsevierStyleCrossRefs" href="#bib0030">6,12–14</a>. Morphology alone has not been sufficient to resolve the taxonomic questions surrounding the genus. DNA sequences have been very effective at confirming or rejecting the major pine lineages that were originally proposed based primarily on morphology, artificial crosses, and biochemical studies<a class="elsevierStyleCrossRefs" href="#bib0045">9,15</a>. However, natural and artificial hybridization are relatively well documented in pines<a class="elsevierStyleCrossRef" href="#bib0080">16</a>. Both hybridization and retention of ancestral alleles have obscured the phylogenetic relationships among closely related species<a class="elsevierStyleCrossRefs" href="#bib0085">17–24</a>.In most organisms, speciation both generates biological diversity and gives rise to gene genealogies that reflect the history of divergence of the organisms in which the genes are found. These histories can be reconstructed using phylogenetic analysis and used to infer the phylogenetic relationships among species. In pines, hybridization and incomplete lineage sorting cause a remarkable disconnection between individual gene trees and species relationships<a class="elsevierStyleCrossRef" href="#bib0120">24</a>. As a result, many gene trees need to be considered separately (although not in isolation) to form hypotheses regarding species relationships.Plastid (e.g., chloroplast) DNA has been widely studied in pines. In contrast to angiosperms, plastid DNA is paternally inherited in conifers<a class="elsevierStyleCrossRef" href="#bib0125">25</a>. It exhibits low genetic differentiation in conifer populations and species relative to mitochondrial or nuclear DNA; this is because plastid gene flow is mediated by both seed and pollen in conifers<a class="elsevierStyleCrossRef" href="#bib0130">26</a>. One result is that species take a long time to become fixed for a single plastid DNA lineage, and sharing of plastid lineages by closely related species is often observed<a class="elsevierStyleCrossRefs" href="#bib0090">18,27</a>.We chose as our study area the political boundaries marked by Guerrero and Oaxaca. These states are located on the southern Pacific coast of Mexico. Guerrero occupies 63,596km2 and is the fourteenth largest state in the country, and Oaxaca occupies 93,757km2 and is the fifth largest<a class="elsevierStyleCrossRef" href="#bib0140">28</a>. The principal mountain range in these states is the Sierra Madre del Sur, which extends through Jalisco, Colima, southern Michoacán, the State of Mexico, Morelos, Puebla, Guerrero, and Oaxaca, and is bordered to the north by the Transverse Mexican Volcanic Belt<a class="elsevierStyleCrossRef" href="#bib0145">29</a>. Both Guerrero and Oaxaca are highly diverse in species. For example, 9,362 species of plants have been reported for Oaxaca, more than any other state in the country<a class="elsevierStyleCrossRef" href="#bib0150">30</a>. Fourteen pine species have been recognized for Guerrero<a class="elsevierStyleCrossRef" href="#bib0155">31</a> and 17 for Oaxaca<a class="elsevierStyleCrossRef" href="#bib0160">32</a>. Seven of these species belong to Pinus subsection Australes.This is the first comparative study to integrate morphological, anatomical, and molecular characters in pines of Guerrero and Oaxaca, Mexico. Our objectives are to better understand the morphological diversity of Pinus subsection Australes in these two states and to further document which species in this group share plastid DNA haplotypes.MATERIALS AND METHODSDried specimens were obtained from the Inventario Nacional Forestal and the National Herbarium (MEXU). Ninety individuals were examined morphologically (<a class="elsevierStyleCrossRef" href="#fig0005">Figure 1</a>; <a class="elsevierStyleCrossRef" href="#sec0045">Appendix 1</a>). Morphological measurements of needles (secondary leaves), fascicle sheaths, seed cones, and peduncles were taken from dried collections. All measurements were made with a ruler and expressed in cm. For needle and fascicle sheath measurements, 15 arbitrarily chosen fascicles were measured. Seed cone and peduncle measurements were based on a single mature cone per individual (<a class="elsevierStyleCrossRef" href="#fig0010">Figure 2</a>). Transverse sections were made from the medial part of leaves, fixed in formaldehyde-acetic acid-alcohol (FAA), and examined with an Olympus stereomicroscope. The numbers of stomatal lines on the abaxial and adaxial leaf surfaces were counted, and observations were made of the dermal tissues, mesophyll, and endodermis.<elsevierMultimedia ident="fig0005"></elsevierMultimedia><elsevierMultimedia ident="fig0010"></elsevierMultimedia>Statistical analyses were performed in R ver. 3.03<a class="elsevierStyleCrossRef" href="#bib0165">33</a>. Nine variables were analyzed in 83 individuals for which we could obtain a complete set of measurements: sheath length, number of abaxial stomata, number of adaxial stomata, needle length, needle number, cone length, cone width, cone peduncle length, and cone peduncle width. Four individuals with sessile cones were assigned a peduncle length of 0.01 to avoid using zeros. One-way ANOVAs were performed to test for significant differences among species for each of the nine variables. When significant outcomes were obtained, the Tukey method was used as a post hoc test to identify which species had significant differences between their means. Principal components analysis (PCA) was used to explore the relative importance of the leaf and cone variables for explaining overall variance in the data. PCA was run on the correlation matrix resulting from the log-transformed variables, with their means centered and rescaled.Leaf or seed megagametophyte tissue was pulverized with a TissueLyser (Qiagen) and genomic DNA was extracted from leaves with the CTAB method<a class="elsevierStyleCrossRef" href="#bib0170">34</a> or from seed megagametophyte tissue using a Wizard Genomic DNA Purification Kit (Promega). Extractions were eluted in buffer (rehydration solution; Promega) and stored at -20°C. The presence of genomic DNA was confirmed by electrophoresis in agarose gel stained with GelRed (Biotium, Inc.).The first plastid genome, or plastome, reported for a pine was that of P. thunbergii Parl.; it was 119,707 b.p. and composed of 61 protein coding genes, 4, ribosomal RNA genes, 32 tRNA genes, and numerous introns, and intergenic spacers<a class="elsevierStyleCrossRef" href="#bib0175">35</a>. Variable sites are not distributed equally throughout the genome. In addition to differences between protein and RNA coding regions and introns and intergenic spacers noncoding regions, there is variation in substitution rates among the different coding regions. The two most variable exons in pine plastomes are called ycf1 and ycf2; both have an elevated nonsynonymous substitution rate and a considerable number of indels, resulting in changes in the amino acid sequences of the proteins that they encode<a class="elsevierStyleCrossRef" href="#bib0180">36</a>.We evaluated variation in an 840 b.p. fragment of ycf1 in Pinus subsection Australes. This fragment represents 13% of the total gene length with respect to the P. thunbergii reference plastome annotation. DNA amplification was carried out using the polymerase chain reaction (PCR). The final concentrations for the PCR were as follows: 1X Buffer, 1.4mM of MgCl2, 0.2mM of each dNTP, 1μM of forward and reverse primers, and 0.625 U of recombinant Taq polymerase. Reactions were carried out as follows: 94°C, 3min; 35 ×(94°C, 1min; 50°C, 1min; 72°C, 2min); 72°C, 5min; 4°C, 5min. The primers used were Pt96887F (5’- tcatttcgaatctttcggattt-3’) and Pt97833R (5’-taccagaatcggacgtgtca-3’)<a class="elsevierStyleCrossRef" href="#bib0090">18</a>. The PCR products were sent to the University of Washington High Throughput Genomic Center (Seattle, Washington) for purification and sequencing. The same primers used for PCR were used for sequencing.Sequence reads were assembled and edited in Geneious ver. 5<a class="elsevierStyleCrossRef" href="#bib0185">37</a>. Sequences were aligned manually in BioEdit ver. 7.1.9<a class="elsevierStyleCrossRef" href="#bib0190">38</a>. A haplotype network was inferred using statistical parsimony at a 95% confidence level with TCS ver. 1.21<a class="elsevierStyleCrossRef" href="#bib0195">39</a>. Nucleotide diversity (π), the average number of nucleotide differences per site, was also estimated with DnaSP ver. 5<a class="elsevierStyleCrossRef" href="#bib0200">40</a>.RESULTSMorphology and leaf anatomyPinus leiophylla Schiede ex Schltdl. & Cham. is the only species of this group in the study area with deciduous fascicle sheaths, a character that is otherwise almost exclusive to soft pines (Pinus subgenus Strobus). Pinus lawsonii Roezl ex Gordon has nondecurrent bracts on its branches and cone peduncle, compared to decurrent bracts in P. pringlei Shaw. Our measurements of needle lengths coincided with previous works<a class="elsevierStyleCrossRefs" href="#bib0030">6,12,32</a>. The number of leaves per fascicle for P. leiophylla was reported to vary from (2–)3–5 throughout its natural range<a class="elsevierStyleCrossRefs" href="#bib0030">6,12,13</a>; however, we only found needles in fascicles of 5 or very infrequently 4. The number of leaves per fascicle for P. teocote Schiede ex Schltdl. & Cham. was reported by the same authors to vary from 2–5; however, we found only 3 or very occasionally 4 leaves per fascicle.An ANOVA F-test indicated that statistical differences existed among species means for all nine morphological variables evaluated (p < 0.001; results not shown). For example, mean fascicle sheath lengths were significantly different (p < 0.05) for 16 of 28 species pairs, mean numbers of abaxial stomata were significantly different for 17 of 28 species pairs, mean needle length was significantly different for 18 of 28 species pairs, and mean cone length was significantly different for 14 of 28 species pairs.After log transformation, the pairs of variables with the highest correlations were sheath length versus needle length (r=0.649) and the number of abaxial versus dorsal stomatal rows (r=0.844). For the PCAs the cumulative variance for the first, second, and third components were 40.6, 59.7, and 74.5%, respectively. Seven components were needed to explain >95% of the variance. The variables with the highest absolute loading values for the first component were needle length (0.397), number of abaxial stomata (0.390), and number of adaxial stomata (0.386). Needle number was the only variable with a negative value (-0.19). The variables with the highest absolute loading values for the second component were cone peduncle length (0.515), needle number (0.486), and peduncle width (0.334). The variable with the highest negative value for the second component was number of abaxial stomata (-0.312).The results of the statistical analyses coincided with our observation that the most useful characters for identifying species of Pinus subsection Australes in Guerrero and Oaxaca were length of the fascicle sheath and needles, the number of rows of stomatal lines, and seed cone peduncle length; qualitative variables such as persistence of fascicle sheaths and several aspects of the seed cone scale were also useful (<a class="elsevierStyleCrossRef" href="#tbl0005">Table I</a>). Variation in the length or width of the seed cones overlapped greatly among species. Pinus herrerae Martínez had the smallest cones of all species, measuring 4.5–5.5cm in length. The shape of the apophysis and umbo of the cone scale was useful for distinguishing P. oocarpa Schiede, P. pringlei, P. lawsonii, and P. herrerae<a class="elsevierStyleCrossRefs" href="#bib0030">6,13</a>.<elsevierMultimedia ident="tbl0005"></elsevierMultimedia>The anatomical characters of secondary leaves that were compared were the thickness of the hypodermal cell walls, presence or absence of hypodermal cell intrusions into the mesophyll, and position and number of resin canals in the mesophyll (<a class="elsevierStyleCrossRef" href="#tbl0010">Table II</a>). Endodermal characters were not considered here. All characters coincided with or were within the range of variation reported in previous studies<a class="elsevierStyleCrossRefs" href="#bib0030">6,12–14</a>.<elsevierMultimedia ident="tbl0010"></elsevierMultimedia>The thickness of hypodermal cell walls was useful for distinguishing P. lawsonii and P. teocote. Intrusions of hypodermal cells into the mesophyll were observed in some transverse sections of P. lawsonii, but never in P. teocote. The position of resin canals was less variable in these two species than has been reported in studies that take into account their entire geographical distribution<a class="elsevierStyleCrossRefs" href="#bib0030">6,12,13</a>; resin canals were in an internal position (in contact with the endodermis) for both species. Resin canals in a medial position (not in contact with hypodermis or endodermis) were rare. Only P. oocarpa had resin canals in a septal position (in contact with both the endodermis and the hypodermis). Leaf anatomical characters were also useful for confirming morphological identifications of P. patula; none of these individuals agreed in leaf anatomy (e.g., none had five resin canals) with the morphologically similar P. tecunumanii, a species reported for Oaxaca by Farjon & Styles<a class="elsevierStyleCrossRef" href="#bib0030">6</a> but subsequently excluded<a class="elsevierStyleCrossRef" href="#bib0150">30</a>. The number of resin canals in the mesophyll was typically 3 for all species except P. oocarpa (5) and to a lesser extent, P. pringlei (2–5).Identification key for Pinus subsection Australes in Guerrero and Oaxaca<ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005">1.Fascicle sheaths deciduous, needles in fascicles of 5, 7–13cm long. .. P. leiophylla1’. Fascicle sheaths persistent, 11–32cm long, needles in fascicles of 3–5. .. 2<ul class="elsevierStyleList" id="lis0010"><li class="elsevierStyleListItem" id="lsti0010">2.Needles in fascicles of 4–5, 14.5–29cm long, cone peduncle 1.5–3cm long, seed cones ovoid to subglobose. .. P. oocarpa2’. Needles in fascicles of 3–4(–5), cone peduncle < 1.5cm long, seed cones ovoid-attenuate. .. 3<ul class="elsevierStyleList" id="lis0015"><li class="elsevierStyleListItem" id="lsti0015">3.Needles drooping to pendent, in fascicles of 3–5, 12–22cm long, 4 rows of stomata on abaxial face, 4–6 adaxial, cone sessile or with peduncle ≤ 1cm long. .. P. patula3’. Needles spreading or erect, in fascicles of 3–4. .. 4<ul class="elsevierStyleList" id="lis0020"><li class="elsevierStyleListItem" id="lsti0020">4.Bracts non-decurrent, needles 11–27cm long. .. P. lawsonii4’. Bracts decurrent. .. 5<ul class="elsevierStyleList" id="lis0025"><li class="elsevierStyleListItem" id="lsti0025">5.Prickles on cone scale umbo persistent<ul class="elsevierStyleList" id="lis0030"><li class="elsevierStyleListItem" id="lsti0030">6.Needles with 3–5 rows of stomata on abaxial face, 4–6 adaxial, cone peduncle 0.8–1.4cm long × 0.4–0.8cm wide. .. P. patula var. longipedunculata6’. Needles with 8–11 abaxial rows of stomata on abaxial face, 8–12 adaxial, cone peduncle 0.5–2.2cm long × 0.8–1.3cm wide. .. P. pringlei</li></ul>5’. Prickles on cone scale umbo deciduous. .. 7<ul class="elsevierStyleList" id="lis0035"><li class="elsevierStyleListItem" id="lsti0035">7.Needle length 14.5–19cm, 4–7 rows of stomata on abaxial face, 4–8 adaxial. .. P. herrerae7’. Needle length 9–16cm, 8–10(–12) rows of stomata on abaxial face, 10 adaxial. .. P. teocote</li></ul></li></ul></li></ul></li></ul></li></ul></li></ul>Intra- and inter-specific comparison of plastid DNA sequences and haplotypesThe complete plastid DNA sequence matrix of 49 individuals had a total of seven variable sites; six of the site variants were present in more than one individual (parsimony informative). Nucleotide diversity (π) for all species considered together was 0.00191. The haplotype diversity (Hd) was 0.732. The seven variable sites were distributed among nine different haplotypes, five of which were shared by two or more species (<a class="elsevierStyleCrossRef" href="#tbl0015">Table III</a>).<elsevierMultimedia ident="tbl0015"></elsevierMultimedia>The sequences of all the individuals were used to construct a haplotype network (<a class="elsevierStyleCrossRef" href="#fig0015">Figure 3</a>). Haplotype 9 occurred at the highest frequency (n = 23) and was shared by the four species P. lawsonii, P. oocarpa, P. pringlei, and P. teocote. It was near the center of the network, with three connections to other haplotypes. The same four species also shared the second most common haplotype (haplotype 8; n = 9). Only one of these four species, P. oocarpa, shared a haplotype with other species in the area, haplotype 7 (n = 6) with P. leiophylla and P. herrerae. In contrast, the two varieties of P. patula only shared haplotypes with each other and with P. herrerae; Haplotype 6, the fourth most frequent in the network (n = 5), was shared among these three taxa. Haplotype 5 (n = 2) was made up of sequences from the two varieties of P. patula; it had the most connections (four) to other haplotypes in the network. The remaining four haplotypes, found in P. lawsonii, P. teocote, P. patula, and P. patula var. longipedunculata, each only occurred once.<elsevierMultimedia ident="fig0015"></elsevierMultimedia>DISCUSSIONMorphology is central for the study of taxonomy. It permits the documentation of taxonomic limits, geographic distribution, and phenotypic variation. It is often also useful for the detection of hybrids. In pines, a diversity of leaf types, notably bracts along the branches, fascicle sheaths, and secondary needles, are useful for identifying species, as are seed cone size and shape, length of the peduncle, and morphology of the cone scale apex. Needle anatomy has long been used to delimit or identify species of pines<a class="elsevierStyleCrossRefs" href="#bib0030">6,11,12</a>. Careful study of morphology and needle anatomy has the potential of identifying putative natural hybrids. The distribution of the species studied here has been described previously<a class="elsevierStyleCrossRefs" href="#bib0155">31,32</a>, but finer scale information is still lacking. Further field exploration and herbarium work are needed to better establish the geographic distribution of these species in southern Mexico and to capture the full range of morphological and molecular variation of species with ranges that extend beyond Guerrero and Oaxaca.Forest management practices may be altering the geographic distribution of tree taxa in the area; it is important to establish their natural distributions and determine if interspecific gene flow is being promoted by human activities, particularly the planting of economically important species outside of their natural range of distribution.The 840 b.p. fragment of plastid DNA characterized in this study offers corroborating evidence of shared plastid DNA haplotypes in North American pines<a class="elsevierStyleCrossRefs" href="#bib0085">17–23</a>. A previous study that included a 5,425 b.p. plastid DNA alignment for 3–6 individuals per species of Pinus subsection Australes recovered all haplotypes as monophyletic by species for P. herrerae (three individuals), P. leiophylla (four individuals), and P. teocote (three individuals)<a class="elsevierStyleCrossRef" href="#bib0090">18</a>. However, in this study we report a P. herrerae individual (and a P. oocarpa individual) with the typical haplotype of P. leiophylla, and two haplotypes in P. teocote that are widespread in other species of Pinus subsection Australes. Further evidence of shared haplotypes, previously reported at a broader geographical scale<a class="elsevierStyleCrossRef" href="#bib0090">18</a>, was also found within the study region for P. lawsonii, P. oocarpa, P. pringlei, and P. teocote.Sharing of plastid haplotypes may be widespread among closely related species of conifers and some other tree genera. The two processes most likely responsible for this phenomenon in pines are introgressive hybridization and incomplete lineage sorting. These processes are probably most widespread in groups of recently diverged species, such as within Pinus subsection Ponderosae and Pinus subsection Australes<a class="elsevierStyleCrossRef" href="#bib0090">18</a>, or in young putative species pairs, such as the Mexican white pines, P. ayacahuite and P. strobiformis<a class="elsevierStyleCrossRef" href="#bib0110">22</a>. Less plastid haplotype sharing has been reported from other conifer lineages with fewer recently diverged species such as North American pinyon pines<a class="elsevierStyleCrossRefs" href="#bib0205">41,42</a>. Some recently diverged lineages, such as P. caribaea of Pinus subsection Australes<a class="elsevierStyleCrossRef" href="#bib0215">43</a> also show little evidence of shared haplotypes, possibly due to less co-occurrence with close relatives and smaller effective population sizes. Distinguishing between introgressive hybridization and incomplete lineage sorting can be achieved by comparing the genealogical patterns of plastid DNA with those of nuclear and mitochondrial DNA, both of which are likely to result in different patterns than those found with plastid DNA. Further evidence of hybridization can also be documented through morphological and anatomical studies.CONCLUSIONSGuerrero and Oaxaca are rich in species of Pinus subsection Australes. They occur both allopatrically and sympatrically throughout the montane regions of the state. Although they can be distinguished morphologically, they follow a more generalized pattern in North American hard pines of sharing plastid DNA lineages as a result of introgressive hybridization or incomplete lineage sorting. Inferring plastid DNA genealogies is of unquestionable value for studying evolution in the group, but it is important to be aware that adequate infraspecific sampling and inclusion of sufficient variable sites are needed for plastid DNA genealogies to be informative with respect to species relationships.Future studies should incorporate range-wide sampling of species to capture their full range of variation. Sequences from multiple unlinked markers combined with morphometric analysis should permit more accurate detection of interspecific gene flow in the presence of incomplete lineage sorting." "textoCompletoSecciones" => array:1 [ "secciones" => array:13 [ 0 => array:3 [ "identificador" => "xres697146" "titulo" => "ABSTRACT" "secciones" => array:1 [ 0 => array:1 [ "identificador" => "abst0005" ] ] ] 1 => array:2 [ "identificador" => "xpalclavsec706874" "titulo" => "Keywords" ] 2 => array:3 [ "identificador" => "xres697145" "titulo" => "RESUMEN" "secciones" => array:1 [ 0 => array:1 [ "identificador" => "abst0010" ] ] ] 3 => array:2 [ "identificador" => "xpalclavsec706875" "titulo" => "Palabras clave" ] 4 => array:2 [ "identificador" => "sec0005" "titulo" => "INTRODUCTION" ] 5 => array:2 [ "identificador" => "sec0010" "titulo" => "MATERIALS AND METHODS" ] 6 => array:3 [ "identificador" => "sec0015" "titulo" => "RESULTS" "secciones" => array:1 [ 0 => array:2 [ "identificador" => "sec0020" "titulo" => "Morphology and leaf anatomy" ] ] ] 7 => array:2 [ "identificador" => "sec0025" "titulo" => "Identification key for Pinus subsection Australes in Guerrero and Oaxaca" ] 8 => array:2 [ "identificador" => "sec0030" "titulo" => "Intra- and inter-specific comparison of plastid DNA sequences and haplotypes" ] 9 => array:2 [ "identificador" => "sec0035" "titulo" => "DISCUSSION" ] 10 => array:2 [ "identificador" => "sec0040" "titulo" => "CONCLUSIONS" ] 11 => array:2 [ "identificador" => "xack232959" "titulo" => "ACKNOWLEDGEMENTS" ] 12 => array:1 [ "titulo" => "REFERENCES" ] ] ] "pdfFichero" => "main.pdf" "tienePdf" => true "fechaRecibido" => "2015-09-14" "fechaAceptado" => "2016-04-27" "PalabrasClave" => array:2 [ "en" => array:1 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Keywords" "identificador" => "xpalclavsec706874" "palabras" => array:5 [ 0 => "biodiversity" 1 => "gene flow" 2 => "lineage sorting" 3 => "pines" 4 => "species delimitation" ] ] ] "es" => array:1 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Palabras clave" "identificador" => "xpalclavsec706875" "palabras" => array:5 [ 0 => "biodiversidad" 1 => "flujo génico" 2 => "sorteo de linajes" 3 => "pinos" 4 => "delimitación de especies" ] ] ] ] "tieneResumen" => true "resumen" => array:2 [ "en" => array:2 [ "titulo" => "ABSTRACT" "resumen" => "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." ] "es" => array:2 [ "titulo" => "RESUMEN" "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 introgresiva y la retención de polimorfismos ancestrales. Aquí describimos la diversidad de haplotipos de plastidio y la morfología para este grupo de especies en los estados de Guerrero y Oaxaca, México. Siete especies de Pinus subsección Australes son reconocidas en el área de estudio, una de las cuales, P. patula incluye dos variedades. Siete sitios variables y nueve haplotipos fueron encontrados amplificando un fragmento de 840 p. b. de ADN de la región codificante ycf1. Se encontraron haplotipos compartidos para P. patula var. patula, P. patula var. longipedunculata, P. herrerae y P. tecunumanii. Cuatro de los nueve haplotipos encontrados están restringidos a Oaxaca. Aunque las genealogías de genes son valiosas para estudiar la evolución de este grupo, se requieren mayor muestreo de individuos y más sitios variables para la inferencia de relaciones entre las especies." ] ] "apendice" => array:1 [ 0 => array:1 [ "seccion" => array:1 [ 0 => array:4 [ "apendice" => "Pinus herreraeMartínez México. Oaxaca. Loma El Ocote, I. Martínez Martínez 71204. Guerrero. Cerro La Mula, E.O. Valencia Santana 71124. Cerro Faisán, M.A. Juárez 72227. Omiltemi, Chilpancingo, J. A. Pérez de la Rosa 132. Camino Paraíso, L. Lozada 182A. Pinus lawsoniiRoezl ex Gordon México. Oaxaca. Arroyo Cangarec, O. Rodríguez Santamaría 74754. La Ocotera, O. Rodríguez Santamaría 73947. Arroyo Artemio Reyes, G.R. Izaguirre Yáñez 76960. Barranca Pelona, J.C. Calvillo García 71396. El Zacatón, J. Zúñiga Reyes 74549. Agua Cola, J. Zúñiga 74747. Pino Gordo, O. J. Pérez Martínez 72853. Nanche, R, Boquedano Peralta 76955. Guerrero. Exsuyo, J.C. Calvillo García 71587. La Laguna, J.C. Calvillo García 70214. El Chaude, A.J. Fortoul Velasco 70254. Pinus leiophyllaSchiede ex Schltdl. & Cham. México. Oaxaca. La Plaga, R. Baquedano Peralta 72700. Monte de Tesoro, J. Reyes Santiago 883. Miahuatlán, E. Hunn 0096. Santa María del Rosario, A.J. Fortoul Velasco 72252. Ixtlán, A. García-Mendoza 7585. Guerrero. La Guitarra, A.J. Fortoul Velasco 71557. La Culebra, A.J. Fortoul Velasco 71773. Pinus oocarpaSchiede ex Schltdl. México. Oaxaca: El Corolón-El Retén, T. Yescas de los Ángeles 75410. Agua Fría, T. Yescas de los Ángeles 75712. Loma del Chifle, T. Yescas de los Ángeles 76958. La Ermita, J.L. Flores Nicanor 71650. La Cascada, E. Martínez V. 73980, Rancho Viejo, T. Yescas de los Ángeles 72018. Guerrero. La Sidra, J.C. Bautista Martínez 71991. Agua de Obispo, J.C. Calvillo García 72424. La Coscolina, O.J. Pérez Martínez 72838. El Ermitaño, J.C. Calvillo García 70448. Tierra Blanca, J.C. Calvillo García 72023. Pinus patulaSchltdl. & Cham. var.patula México. Oaxaca. Terracería La Venta, J. Reyes 6493. La Cima, M.I. Alvarado Flores 72489. Lomatico, J.C. Bautista Martínez 73310. Pinus patulavar.longipedunculataLoock ex Martínez México. Oaxaca. Aserrado Las Vigas, J. A. Pérez de la Rosa 1849. San Juan Tepeuxila, R. Torres Colín 16168. Santa María Yavesía, I. Trejo 3064. Pinus pringleiShaw México. Oaxaca. Nebrón, J. Zúñiga Reyes 71153. La Caoilla, Ojo de Agua, M. I. Alvarado Flores 69610. El Chaneque, J.L. Flores Nicanor 75148. Silacayuapan, R. Luna Martínez 71600. Tres Cruces, J.L. Flores Nicanor 74544. San Isidro El Chiñón, R. Luna Martínez 71813. Buenavista, O. Rodríguez Santamaría 70069. Guerrero. Cajones, J.C. Calvillo García 65501. Cocuitlazola, J.C. Calvillo García 72446. C. Tlacholoya, J.C. Calvillo García 71371. Ejido El Carrazal, Coyuca de Catalán, E. Rodríguez Ibarra 69525. Huexoapa, F. Rosas Aguilar 72658. Pinus teocoteSchiede ex Schltdl. & Cham.México. Oaxaca. Agua Colorada, I. Martínez Martínez 74372. Cerro Manteca, J.L. Flores Nicanor 75345. Yacuni, M.I. Alvarado Flores 72898. La Ciénaga, J.L. Flores Nicanor 73537, La Joya del Fresno, T. Yescas de los Ángeles 72476. Río Milpas, R. Baquedano Peralta 73534. Barranca del Zopilote, J.C. Calvillo García 71607. San Francisco Higos, R. Lina Martínez 72028. Guerrero. Mesa de la Mujer, A.J. Fortoul Velasco 68303. San Andrés, A.J. Fortoul Velasco72003." "etiqueta" => "APPENDIX 1" "titulo" => "SPECIMENS EXAMINED. ALL ARE DEPOSITED IN MEXU." "identificador" => "sec0045" ] ] ] ] "multimedia" => array:6 [ 0 => array:7 [ "identificador" => "fig0005" "etiqueta" => "Figure 1" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr1.jpeg" "Alto" => 1412 "Ancho" => 2148 "Tamanyo" => 218274 ] ] "descripcion" => array:1 [ "en" => "The collection localities for the individuals included in this study." ] ] 1 => array:7 [ "identificador" => "fig0010" "etiqueta" => "Figure 2" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr2.jpeg" "Alto" => 1208 "Ancho" => 2149 "Tamanyo" => 240316 ] ] "descripcion" => array:1 [ "en" => "Morphological diversity of seed cones for Pinus subsection Australes in Guerrero and Oaxaca. A. Pinus herrerae González & Martínez 770. B. Pinus lawsonii Soto Núñez et al., 5307. C. Pinus leiophylla Reyes 1989. D. Pinus oocarpa Yescas de los Ángeles 75712. E. Pinus patula var. patula Bautista Martínez 73310. F. Pinus patula var. longipedunculata Trejo 3053. G. Pinus pringlei Rico 489. H. Pinus teocote Román 335. Photographs by Carmen Loyola." ] ] 2 => array:7 [ "identificador" => "fig0015" "etiqueta" => "Figure 3" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr3.jpeg" "Alto" => 1043 "Ancho" => 2148 "Tamanyo" => 166660 ] ] "descripcion" => array:1 [ "en" => "Plastid DNA haplotype network for the study species. The area of the circles is proportional to haplotype frequency." ] ] 3 => array:8 [ "identificador" => "tbl0005" "etiqueta" => "Table I" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at1" "detalle" => "Table " "rol" => "short" ] ] "tabla" => array:1 [ "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Taxon \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Fascicle sheath \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Fascicle sheath length (cm) \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Leaves per fascicle \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Leaf length (cm) \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Number of abaxial stomatal rows \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Peduncle length (cm) \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Cone length (cm) \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Umbo shape \t\t\t\t\t\t\n \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. herrerae \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.6–1.7 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3(–4) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">14.5–19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4–7 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.4–1.3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4.5–5.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">slightly raised \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. lawsonii \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1–2.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3(–4) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">11–27 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">7–10 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1–1.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5–8 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">flat to slightly raised \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. leiophylla \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">deciduous \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">7–13 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.8–1.3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4.5–6.2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">flat to slightly raised \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. oocarpa \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1–2.2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">14.5–29 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">(3–)4–6(–14) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1.5–3.0 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4.5–6.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">usually flat \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. patula \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1.1–1.3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3–4(–5) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">12–22 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.5–1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">7–7.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">flat or slightly raised \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. patula var. longipedunculata \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1–1.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3–4(–5) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">14–24.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3–5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.8–1.4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5–8.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">flat or slightly raised \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. pringlei \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1.6–2.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3(–4) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">15–32 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">8–11(–15) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.5–2.2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5–10 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">slightly raised to subpyramidal, \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. teocote \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">persistent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.5–1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3(–4) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">9–16 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">8–10(–12) \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">0.7–1.2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5–6.5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">flat or raised \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab1141315.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Morphological characters of Pinus subsection Australes species in Guerrero and Oaxaca." ] ] 4 => array:8 [ "identificador" => "tbl0010" "etiqueta" => "Table II" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at2" "detalle" => "Table I" "rol" => "short" ] ] "tabla" => array:1 [ "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Taxon \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Hypodermal cell shape \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Hypodermal cell intrusions into mesophyll \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Number of resin canals \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Resin canal position \t\t\t\t\t\t\n \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. herrerae \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">biforme \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">absent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. lawsonii \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">multiforme \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">present \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2–4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. leiophylla \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">absent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2–3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. oocarpa \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">absent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">septal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. patula \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick or multiforme \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">present \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2–3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">medial and internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. patula var. longipedunculata \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick or multiforme \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">present \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2–3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">medial and internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. pringlei \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">biforme or multiforme \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">present \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2–5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">internal \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. tecunumanii \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">absent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">medial \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top">P. teocote \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">thick \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">absent \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">3–4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">internal \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab1141317.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Leaf anatomical characters of Pinus subsection Australes species in Guerrero and Oaxaca. Pinus tecunumanii is also included for comparison (see text)." ] ] 5 => array:8 [ "identificador" => "tbl0015" "etiqueta" => "Table III" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at3" "detalle" => "Table II" "rol" => "short" ] ] "tabla" => array:1 [ "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Haplotype Number \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">208 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">311 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">326 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">470 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">717 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">797 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">819 \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">N \t\t\t\t\t\t\n \t\t\t\t</th><th class="td" title="table-head " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Species \t\t\t\t\t\t\n \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">9 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">23 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. lawsonii, P. oocarpa, P. pringlei, P. teocote \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">8 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">9 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. lawsonii, P. oocarpa, P. pringlei, P. teocote \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">7 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">6 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. herrerae, P. leiophylla, P. oocarpa \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">6 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. herrerae, P. patula, P. patula var. longipedunculata \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">5 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. patula, P. patula var. longipedunculata \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">4 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. patula \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">3 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. patula var. longipedunculata \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">2 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">G \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. lawsonii \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">C \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">T \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="center" valign="top">1 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="table-entry " align="left" valign="top">P. teocote \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab1141316.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Plastid DNA haplotype variation in fragment ycf1 (sequence of primer 5’-3’) and their distribution in the pine species sampled." ] ] ] "bibliografia" => array:2 [ "titulo" => "REFERENCES" "seccion" => array:1 [ 0 => array:2 [ "identificador" => "bibs0005" "bibliografiaReferencia" => array:43 [ 0 => array:3 [ "identificador" => "bib0005" "etiqueta" => "1" "referencia" => array:1 [ 0 => array:2 [ "contribucion" => array:1 [ 0 => array:2 [ "titulo" => "In Ecology and biogeography" "autores" => array:1 [ 0 => array:2 [ "etal" => false "autores" => array:2 [ 0 => "D.M. 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"fecha" => "2010" "volumen" => "107" "paginaInicial" => "229" "paginaFinal" => "241" "link" => array:1 [ 0 => array:2 [ "url" => "https://www.ncbi.nlm.nih.gov/pubmed/21118838" "web" => "Medline" ] ] ] ] ] ] ] ] ] ] ] ] "agradecimientos" => array:1 [ 0 => array:4 [ "identificador" => "xack232959" "titulo" => "ACKNOWLEDGEMENTS" "texto" => "This study formed part of the undergraduate thesis of Alfredo Ortiz-Martínez for the Instituto Tecnológico de Cd. Altamirano. We thank Mauricio Mora for assistance in the herbarium, Carmen Loyola for providing photographs of seed cones, and Jazmín Cortez Sarabia, Erika Oropeza Bruno, Jesús Pascual Orozco, Francisco Zavala Hernández, Lev Jardón Barbolla, and two anonymous reviewers for providing helpful comments on a previous draft of this manuscript. Most of the plant material in this study was provided by the project “Remuestreo del Inventario Nacional Forestal y de Suelos 2009-2013” awarded by the Comisión Nacional Forestal (CONAFOR) to Martin Ricker. We thank the Dirección General de Asuntos de Personal Académico-Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica of the Universidad Nacional Autónoma de México (DGAPA-PAPIIT; IN228209) for financial support for laboratory work, and CONACYT (INFR-2014-1 No. 224743) for supporting the installation of herbarium compactors that have allowed for the storage of the specimens for this study." 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ARTÍCULO ORIGINAL

SPECIES DIVERSITY AND PLASTID DNA HAPLOTYPE DISTRIBUTIONS OF PINUS SUBSECTION AUSTRALES (PINACEAE) IN GUERRERO AND OAXACA

Diversidad de especies y distribución de haplotipos de ADN del plastidio de Pinus subsección Australes(Pinaceae) en Guerrero y Oaxaca

Alfredo Ortiz-Martíneza, David S. Gernandtb,

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dgernandt@ib.unam.mx

Corresponding author.

a Instituto Tecnológico de Cd. Altamirano, Av. Pungarabato Poniente s/n, Col. Morelos, Cd. Altamirano, C.P. 40660, Guerrero, México

b Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México. Ciudad Universitaria, Deleg. Coyoacán, C.P. 04510, Ciudad de México, México

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Bailey have solitary needles&#41;&#44; and woody seed cones with thickened scale apices and a dorsal or terminal umbo &#40;a specialized raised region resulting from more than one season of growth&#41;&#46; Mexico is the center of species diversity for pines&#44; with more than 40&#37; of the world&#39;s species occurring naturally within its borders<a class="elsevierStyleCrossRefs" href="#bib0010"><span class="elsevierStyleSup">2&#8211;7</span></a>&#46; The number of species recognized for the country has varied among recent authors&#59; Eckenwalder<a class="elsevierStyleCrossRef" href="#bib0010"><span class="elsevierStyleSup">2</span></a> recognized 39 species&#44; Farjon<a class="elsevierStyleCrossRef" href="#bib0015"><span class="elsevierStyleSup">3</span></a> recognized 44&#44; and Gernandt and P&#233;rez-de la Rosa<a class="elsevierStyleCrossRef" href="#bib0035"><span class="elsevierStyleSup">7</span></a> recognized 49&#46;</p><p id="par0010" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">Pinus</span> is classified in two subgenera&#44; four sections&#44; and 11 subsections<a class="elsevierStyleCrossRef" href="#bib0040"><span class="elsevierStyleSup">8</span></a>&#46; <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> is the most species rich&#44; comprising approximately 29 species distributed throughout North and Central America and the westernmost islands of the Caribbean except Jamaica&#46; It includes the southern yellow pines&#44; the egg-cone pines&#44; and the California closed-cone pines&#44; and is classified together with subsections <span class="elsevierStyleItalic">Contortae</span>&#44; and <span class="elsevierStyleItalic">Ponderosae</span> in <span class="elsevierStyleItalic">Pinus</span> subgenus <span class="elsevierStyleItalic">Pinus</span> section <span class="elsevierStyleItalic">Trifoliae</span>&#44; a group known informally as the North American hard pines<a class="elsevierStyleCrossRef" href="#bib0045"><span class="elsevierStyleSup">9</span></a>&#46; All species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> have two vascular bundles in their leaves&#44; and most have persistent fascicle sheaths&#46; All have woody seed cones&#44; and some have strongly developed cone scale apophyses and umbos&#44; often with a mucro&#46; Their seeds have an articulate wing and are primarily wind-dispersed&#46;</p><p id="par0015" class="elsevierStylePara elsevierViewall">Numerous taxonomic works have treated <span class="elsevierStyleItalic">Pinus</span><a class="elsevierStyleCrossRefs" href="#bib0010"><span class="elsevierStyleSup">2&#44;9&#8211;11</span></a>&#44; including regional floras that include Mexico<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;12&#8211;14</span></a>&#46; Morphology alone has not been sufficient to resolve the taxonomic questions surrounding the genus&#46; DNA sequences have been very effective at confirming or rejecting the major pine lineages that were originally proposed based primarily on morphology&#44; artificial crosses&#44; and biochemical studies<a class="elsevierStyleCrossRefs" href="#bib0045"><span class="elsevierStyleSup">9&#44;15</span></a>&#46; However&#44; natural and artificial hybridization are relatively well documented in pines<a class="elsevierStyleCrossRef" href="#bib0080"><span class="elsevierStyleSup">16</span></a>&#46; Both hybridization and retention of ancestral alleles have obscured the phylogenetic relationships among closely related species<a class="elsevierStyleCrossRefs" href="#bib0085"><span class="elsevierStyleSup">17&#8211;24</span></a>&#46;</p><p id="par0020" class="elsevierStylePara elsevierViewall">In most organisms&#44; speciation both generates biological diversity and gives rise to gene genealogies that reflect the history of divergence of the organisms in which the genes are found&#46; These histories can be reconstructed using phylogenetic analysis and used to infer the phylogenetic relationships among species&#46; In pines&#44; hybridization and incomplete lineage sorting cause a remarkable disconnection between individual gene trees and species relationships<a class="elsevierStyleCrossRef" href="#bib0120"><span class="elsevierStyleSup">24</span></a>&#46; As a result&#44; many gene trees need to be considered separately &#40;although not in isolation&#41; to form hypotheses regarding species relationships&#46;</p><p id="par0025" class="elsevierStylePara elsevierViewall">Plastid &#40;e&#46;g&#46;&#44; chloroplast&#41; DNA has been widely studied in pines&#46; In contrast to angiosperms&#44; plastid DNA is paternally inherited in conifers<a class="elsevierStyleCrossRef" href="#bib0125"><span class="elsevierStyleSup">25</span></a>&#46; It exhibits low genetic differentiation in conifer populations and species relative to mitochondrial or nuclear DNA&#59; this is because plastid gene flow is mediated by both seed and pollen in conifers<a class="elsevierStyleCrossRef" href="#bib0130"><span class="elsevierStyleSup">26</span></a>&#46; One result is that species take a long time to become fixed for a single plastid DNA lineage&#44; and sharing of plastid lineages by closely related species is often observed<a class="elsevierStyleCrossRefs" href="#bib0090"><span class="elsevierStyleSup">18&#44;27</span></a>&#46;</p><p id="par0030" class="elsevierStylePara elsevierViewall">We chose as our study area the political boundaries marked by Guerrero and Oaxaca&#46; These states are located on the southern Pacific coast of Mexico&#46; Guerrero occupies 63&#44;596<span class="elsevierStyleHsp" style=""></span>km<span class="elsevierStyleSup">2</span> and is the fourteenth largest state in the country&#44; and Oaxaca occupies 93&#44;757<span class="elsevierStyleHsp" style=""></span>km<span class="elsevierStyleSup">2</span> and is the fifth largest<a class="elsevierStyleCrossRef" href="#bib0140"><span class="elsevierStyleSup">28</span></a>&#46; The principal mountain range in these states is the Sierra Madre del Sur&#44; which extends through Jalisco&#44; Colima&#44; southern Michoac&#225;n&#44; the State of Mexico&#44; Morelos&#44; Puebla&#44; Guerrero&#44; and Oaxaca&#44; and is bordered to the north by the Transverse Mexican Volcanic Belt<a class="elsevierStyleCrossRef" href="#bib0145"><span class="elsevierStyleSup">29</span></a>&#46; Both Guerrero and Oaxaca are highly diverse in species&#46; For example&#44; 9&#44;362 species of plants have been reported for Oaxaca&#44; more than any other state in the country<a class="elsevierStyleCrossRef" href="#bib0150"><span class="elsevierStyleSup">30</span></a>&#46; Fourteen pine species have been recognized for Guerrero<a class="elsevierStyleCrossRef" href="#bib0155"><span class="elsevierStyleSup">31</span></a> and 17 for Oaxaca<a class="elsevierStyleCrossRef" href="#bib0160"><span class="elsevierStyleSup">32</span></a>&#46; Seven of these species belong to <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span>&#46;</p><p id="par0035" class="elsevierStylePara elsevierViewall">This is the first comparative study to integrate morphological&#44; anatomical&#44; and molecular characters in pines of Guerrero and Oaxaca&#44; Mexico&#46; Our objectives are to better understand the morphological diversity of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> in these two states and to further document which species in this group share plastid DNA haplotypes&#46;</p></span><span id="sec0010" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0030">MATERIALS AND METHODS</span><p id="par0040" class="elsevierStylePara elsevierViewall">Dried specimens were obtained from the Inventario Nacional Forestal and the National Herbarium &#40;MEXU&#41;&#46; Ninety individuals were examined morphologically &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Figure 1</a>&#59; <a class="elsevierStyleCrossRef" href="#sec0045">Appendix 1</a>&#41;&#46; Morphological measurements of needles &#40;secondary leaves&#41;&#44; fascicle sheaths&#44; seed cones&#44; and peduncles were taken from dried collections&#46; All measurements were made with a ruler and expressed in cm&#46; For needle and fascicle sheath measurements&#44; 15 arbitrarily chosen fascicles were measured&#46; Seed cone and peduncle measurements were based on a single mature cone per individual &#40;<a class="elsevierStyleCrossRef" href="#fig0010">Figure 2</a>&#41;&#46; Transverse sections were made from the medial part of leaves&#44; fixed in formaldehyde-acetic acid-alcohol &#40;FAA&#41;&#44; and examined with an Olympus stereomicroscope&#46; The numbers of stomatal lines on the abaxial and adaxial leaf surfaces were counted&#44; and observations were made of the dermal tissues&#44; mesophyll&#44; and endodermis&#46;</p><elsevierMultimedia ident="fig0005"></elsevierMultimedia><elsevierMultimedia ident="fig0010"></elsevierMultimedia><p id="par0045" class="elsevierStylePara elsevierViewall">Statistical analyses were performed in R ver&#46; 3&#46;03<a class="elsevierStyleCrossRef" href="#bib0165"><span class="elsevierStyleSup">33</span></a>&#46; Nine variables were analyzed in 83 individuals for which we could obtain a complete set of measurements&#58; sheath length&#44; number of abaxial stomata&#44; number of adaxial stomata&#44; needle length&#44; needle number&#44; cone length&#44; cone width&#44; cone peduncle length&#44; and cone peduncle width&#46; Four individuals with sessile cones were assigned a peduncle length of 0&#46;01 to avoid using zeros&#46; One-way ANOVAs were performed to test for significant differences among species for each of the nine variables&#46; When significant outcomes were obtained&#44; the Tukey method was used as a post hoc test to identify which species had significant differences between their means&#46; Principal components analysis &#40;PCA&#41; was used to explore the relative importance of the leaf and cone variables for explaining overall variance in the data&#46; PCA was run on the correlation matrix resulting from the log-transformed variables&#44; with their means centered and rescaled&#46;</p><p id="par0050" class="elsevierStylePara elsevierViewall">Leaf or seed megagametophyte tissue was pulverized with a TissueLyser &#40;Qiagen&#41; and genomic DNA was extracted from leaves with the CTAB method<a class="elsevierStyleCrossRef" href="#bib0170"><span class="elsevierStyleSup">34</span></a> or from seed megagametophyte tissue using a Wizard Genomic DNA Purification Kit &#40;Promega&#41;&#46; Extractions were eluted in buffer &#40;rehydration solution&#59; Promega&#41; and stored at -20<span class="elsevierStyleHsp" style=""></span>&#176;C&#46; The presence of genomic DNA was confirmed by electrophoresis in agarose gel stained with GelRed &#40;Biotium&#44; Inc&#46;&#41;&#46;</p><p id="par0055" class="elsevierStylePara elsevierViewall">The first plastid genome&#44; or plastome&#44; reported for a pine was that of <span class="elsevierStyleItalic">P&#46; thunbergii</span> Parl&#46;&#59; it was 119&#44;707 b&#46;p&#46; and composed of 61 protein coding genes&#44; 4&#44; ribosomal RNA genes&#44; 32 tRNA genes&#44; and numerous introns&#44; and intergenic spacers<a class="elsevierStyleCrossRef" href="#bib0175"><span class="elsevierStyleSup">35</span></a>&#46; Variable sites are not distributed equally throughout the genome&#46; In addition to differences between protein and RNA coding regions and introns and intergenic spacers noncoding regions&#44; there is variation in substitution rates among the different coding regions&#46; The two most variable exons in pine plastomes are called <span class="elsevierStyleItalic">ycf1</span> and <span class="elsevierStyleItalic">ycf2</span>&#59; both have an elevated nonsynonymous substitution rate and a considerable number of indels&#44; resulting in changes in the amino acid sequences of the proteins that they encode<a class="elsevierStyleCrossRef" href="#bib0180"><span class="elsevierStyleSup">36</span></a>&#46;</p><p id="par0060" class="elsevierStylePara elsevierViewall">We evaluated variation in an 840 b&#46;p&#46; fragment of <span class="elsevierStyleItalic">ycf1</span> in <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span>&#46; This fragment represents 13&#37; of the total gene length with respect to the <span class="elsevierStyleItalic">P&#46; thunbergii</span> reference plastome annotation&#46; DNA amplification was carried out using the polymerase chain reaction &#40;PCR&#41;&#46; The final concentrations for the PCR were as follows&#58; 1X Buffer&#44; 1&#46;4<span class="elsevierStyleHsp" style=""></span>mM of MgCl<span class="elsevierStyleInf">2</span>&#44; 0&#46;2<span class="elsevierStyleHsp" style=""></span>mM of each dNTP&#44; 1<span class="elsevierStyleHsp" style=""></span>&#956;M of forward and reverse primers&#44; and 0&#46;625 U of recombinant <span class="elsevierStyleItalic">Taq</span> polymerase&#46; Reactions were carried out as follows&#58; 94<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 3<span class="elsevierStyleHsp" style=""></span>min&#59; 35 &#215;&#40;94<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 1<span class="elsevierStyleHsp" style=""></span>min&#59; 50<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 1<span class="elsevierStyleHsp" style=""></span>min&#59; 72<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 2<span class="elsevierStyleHsp" style=""></span>min&#41;&#59; 72<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 5<span class="elsevierStyleHsp" style=""></span>min&#59; 4<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; 5<span class="elsevierStyleHsp" style=""></span>min&#46; The primers used were Pt96887F &#40;5&#8217;- tcatttcgaatctttcggattt-3&#8217;&#41; and Pt97833R &#40;5&#8217;-taccagaatcggacgtgtca-3&#8217;&#41;<a class="elsevierStyleCrossRef" href="#bib0090"><span class="elsevierStyleSup">18</span></a>&#46; The PCR products were sent to the University of Washington High Throughput Genomic Center &#40;Seattle&#44; Washington&#41; for purification and sequencing&#46; The same primers used for PCR were used for sequencing&#46;</p><p id="par0065" class="elsevierStylePara elsevierViewall">Sequence reads were assembled and edited in Geneious ver&#46; 5<a class="elsevierStyleCrossRef" href="#bib0185"><span class="elsevierStyleSup">37</span></a>&#46; Sequences were aligned manually in BioEdit ver&#46; 7&#46;1&#46;9<a class="elsevierStyleCrossRef" href="#bib0190"><span class="elsevierStyleSup">38</span></a>&#46; A haplotype network was inferred using statistical parsimony at a 95&#37; confidence level with TCS ver&#46; 1&#46;21<a class="elsevierStyleCrossRef" href="#bib0195"><span class="elsevierStyleSup">39</span></a>&#46; Nucleotide diversity &#40;&#960;&#41;&#44; the average number of nucleotide differences per site&#44; was also estimated with DnaSP ver&#46; 5<a class="elsevierStyleCrossRef" href="#bib0200"><span class="elsevierStyleSup">40</span></a>&#46;</p></span><span id="sec0015" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0035">RESULTS</span><span id="sec0020" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0040">Morphology and leaf anatomy</span><p id="par0070" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">Pinus leiophylla</span> Schiede ex Schltdl&#46; &#38; Cham&#46; is the only species of this group in the study area with deciduous fascicle sheaths&#44; a character that is otherwise almost exclusive to soft pines &#40;<span class="elsevierStyleItalic">Pinus</span> subgenus <span class="elsevierStyleItalic">Strobus</span>&#41;&#46; <span class="elsevierStyleItalic">Pinus lawsonii</span> Roezl ex Gordon has nondecurrent bracts on its branches and cone peduncle&#44; compared to decurrent bracts in <span class="elsevierStyleItalic">P&#46; pringlei</span> Shaw&#46; Our measurements of needle lengths coincided with previous works<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;12&#44;32</span></a>&#46; The number of leaves per fascicle for <span class="elsevierStyleItalic">P&#46; leiophylla</span> was reported to vary from &#40;2&#8211;&#41;3&#8211;5 throughout its natural range<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;12&#44;13</span></a>&#59; however&#44; we only found needles in fascicles of 5 or very infrequently 4&#46; The number of leaves per fascicle for <span class="elsevierStyleItalic">P&#46; teocote</span> Schiede ex Schltdl&#46; &#38; Cham&#46; was reported by the same authors to vary from 2&#8211;5&#59; however&#44; we found only 3 or very occasionally 4 leaves per fascicle&#46;</p><p id="par0075" class="elsevierStylePara elsevierViewall">An ANOVA F-test indicated that statistical differences existed among species means for all nine morphological variables evaluated &#40;p &#60; 0&#46;001&#59; results not shown&#41;&#46; For example&#44; mean fascicle sheath lengths were significantly different &#40;p &#60; 0&#46;05&#41; for 16 of 28 species pairs&#44; mean numbers of abaxial stomata were significantly different for 17 of 28 species pairs&#44; mean needle length was significantly different for 18 of 28 species pairs&#44; and mean cone length was significantly different for 14 of 28 species pairs&#46;</p><p id="par0080" class="elsevierStylePara elsevierViewall">After log transformation&#44; the pairs of variables with the highest correlations were sheath length versus needle length &#40;r&#61;0&#46;649&#41; and the number of abaxial versus dorsal stomatal rows &#40;r&#61;0&#46;844&#41;&#46; For the PCAs the cumulative variance for the first&#44; second&#44; and third components were 40&#46;6&#44; 59&#46;7&#44; and 74&#46;5&#37;&#44; respectively&#46; Seven components were needed to explain &#62;95&#37; of the variance&#46; The variables with the highest absolute loading values for the first component were needle length &#40;0&#46;397&#41;&#44; number of abaxial stomata &#40;0&#46;390&#41;&#44; and number of adaxial stomata &#40;0&#46;386&#41;&#46; Needle number was the only variable with a negative value &#40;-0&#46;19&#41;&#46; The variables with the highest absolute loading values for the second component were cone peduncle length &#40;0&#46;515&#41;&#44; needle number &#40;0&#46;486&#41;&#44; and peduncle width &#40;0&#46;334&#41;&#46; The variable with the highest negative value for the second component was number of abaxial stomata &#40;-0&#46;312&#41;&#46;</p><p id="par0085" class="elsevierStylePara elsevierViewall">The results of the statistical analyses coincided with our observation that the most useful characters for identifying species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> in Guerrero and Oaxaca were length of the fascicle sheath and needles&#44; the number of rows of stomatal lines&#44; and seed cone peduncle length&#59; qualitative variables such as persistence of fascicle sheaths and several aspects of the seed cone scale were also useful &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table I</a>&#41;&#46; Variation in the length or width of the seed cones overlapped greatly among species&#46; <span class="elsevierStyleItalic">Pinus herrerae</span> Mart&#237;nez had the smallest cones of all species&#44; measuring 4&#46;5&#8211;5&#46;5<span class="elsevierStyleHsp" style=""></span>cm in length&#46; The shape of the apophysis and umbo of the cone scale was useful for distinguishing <span class="elsevierStyleItalic">P&#46; oocarpa</span> Schiede&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span>&#44; <span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; and <span class="elsevierStyleItalic">P&#46; herrerae</span><a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;13</span></a>&#46;</p><elsevierMultimedia ident="tbl0005"></elsevierMultimedia><p id="par0090" class="elsevierStylePara elsevierViewall">The anatomical characters of secondary leaves that were compared were the thickness of the hypodermal cell walls&#44; presence or absence of hypodermal cell intrusions into the mesophyll&#44; and position and number of resin canals in the mesophyll &#40;<a class="elsevierStyleCrossRef" href="#tbl0010">Table II</a>&#41;&#46; Endodermal characters were not considered here&#46; All characters coincided with or were within the range of variation reported in previous studies<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;12&#8211;14</span></a>&#46;</p><elsevierMultimedia ident="tbl0010"></elsevierMultimedia><p id="par0095" class="elsevierStylePara elsevierViewall">The thickness of hypodermal cell walls was useful for distinguishing <span class="elsevierStyleItalic">P&#46; lawsonii</span> and <span class="elsevierStyleItalic">P&#46; teocote</span>&#46; Intrusions of hypodermal cells into the mesophyll were observed in some transverse sections of <span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; but never in <span class="elsevierStyleItalic">P&#46; teocote</span>&#46; The position of resin canals was less variable in these two species than has been reported in studies that take into account their entire geographical distribution<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;12&#44;13</span></a>&#59; resin canals were in an internal position &#40;in contact with the endodermis&#41; for both species&#46; Resin canals in a medial position &#40;not in contact with hypodermis or endodermis&#41; were rare&#46; Only <span class="elsevierStyleItalic">P&#46; oocarpa</span> had resin canals in a septal position &#40;in contact with both the endodermis and the hypodermis&#41;&#46; Leaf anatomical characters were also useful for confirming morphological identifications of <span class="elsevierStyleItalic">P&#46; patula</span>&#59; none of these individuals agreed in leaf anatomy &#40;e&#46;g&#46;&#44; none had five resin canals&#41; with the morphologically similar <span class="elsevierStyleItalic">P&#46; tecunumanii</span>&#44; a species reported for Oaxaca by Farjon &#38; Styles<a class="elsevierStyleCrossRef" href="#bib0030"><span class="elsevierStyleSup">6</span></a> but subsequently excluded<a class="elsevierStyleCrossRef" href="#bib0150"><span class="elsevierStyleSup">30</span></a>&#46; The number of resin canals in the mesophyll was typically 3 for all species except <span class="elsevierStyleItalic">P&#46; oocarpa</span> &#40;5&#41; and to a lesser extent&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span> &#40;2&#8211;5&#41;&#46;</p></span></span><span id="sec0025" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0045">Identification key for <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> in Guerrero and Oaxaca</span><p id="par0295" class="elsevierStylePara elsevierViewall"><ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005"><span class="elsevierStyleLabel">1&#46;</span><p id="par0300" class="elsevierStylePara elsevierViewall">Fascicle sheaths deciduous&#44; needles in fascicles of 5&#44; 7&#8211;13<span class="elsevierStyleHsp" style=""></span>cm long&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; leiophylla</span></p><p id="par0105" class="elsevierStylePara elsevierViewall">1&#8217;&#46; Fascicle sheaths persistent&#44; 11&#8211;32<span class="elsevierStyleHsp" style=""></span>cm long&#44; needles in fascicles of 3&#8211;5&#46; &#46;&#46; 2<ul class="elsevierStyleList" id="lis0010"><li class="elsevierStyleListItem" id="lsti0010"><span class="elsevierStyleLabel">2&#46;</span><p id="par0110" class="elsevierStylePara elsevierViewall">Needles in fascicles of 4&#8211;5&#44; 14&#46;5&#8211;29<span class="elsevierStyleHsp" style=""></span>cm long&#44; cone peduncle 1&#46;5&#8211;3<span class="elsevierStyleHsp" style=""></span>cm long&#44; seed cones ovoid to subglobose&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; oocarpa</span></p><p id="par0115" class="elsevierStylePara elsevierViewall">2&#8217;&#46; Needles in fascicles of 3&#8211;4&#40;&#8211;5&#41;&#44; cone peduncle &#60; 1&#46;5<span class="elsevierStyleHsp" style=""></span>cm long&#44; seed cones ovoid-attenuate&#46; &#46;&#46; 3<ul class="elsevierStyleList" id="lis0015"><li class="elsevierStyleListItem" id="lsti0015"><span class="elsevierStyleLabel">3&#46;</span><p id="par0120" class="elsevierStylePara elsevierViewall">Needles drooping to pendent&#44; in fascicles of 3&#8211;5&#44; 12&#8211;22<span class="elsevierStyleHsp" style=""></span>cm long&#44; 4 rows of stomata on abaxial face&#44; 4&#8211;6 adaxial&#44; cone sessile or with peduncle &#8804; 1<span class="elsevierStyleHsp" style=""></span>cm long&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; patula</span></p><p id="par0125" class="elsevierStylePara elsevierViewall">3&#8217;&#46; Needles spreading or erect&#44; in fascicles of 3&#8211;4&#46; &#46;&#46; 4<ul class="elsevierStyleList" id="lis0020"><li class="elsevierStyleListItem" id="lsti0020"><span class="elsevierStyleLabel">4&#46;</span><p id="par0130" class="elsevierStylePara elsevierViewall">Bracts non-decurrent&#44; needles 11&#8211;27<span class="elsevierStyleHsp" style=""></span>cm long&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; lawsonii</span></p><p id="par0135" class="elsevierStylePara elsevierViewall">4&#8217;&#46; Bracts decurrent&#46; &#46;&#46; 5<ul class="elsevierStyleList" id="lis0025"><li class="elsevierStyleListItem" id="lsti0025"><span class="elsevierStyleLabel">5&#46;</span><p id="par0140" class="elsevierStylePara elsevierViewall">Prickles on cone scale umbo persistent<ul class="elsevierStyleList" id="lis0030"><li class="elsevierStyleListItem" id="lsti0030"><span class="elsevierStyleLabel">6&#46;</span><p id="par0145" class="elsevierStylePara elsevierViewall">Needles with 3&#8211;5 rows of stomata on abaxial face&#44; 4&#8211;6 adaxial&#44; cone peduncle 0&#46;8&#8211;1&#46;4<span class="elsevierStyleHsp" style=""></span>cm long &#215; 0&#46;4&#8211;0&#46;8<span class="elsevierStyleHsp" style=""></span>cm wide&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span></p><p id="par0150" class="elsevierStylePara elsevierViewall">6&#8217;&#46; Needles with 8&#8211;11 abaxial rows of stomata on abaxial face&#44; 8&#8211;12 adaxial&#44; cone peduncle 0&#46;5&#8211;2&#46;2<span class="elsevierStyleHsp" style=""></span>cm long &#215; 0&#46;8&#8211;1&#46;3<span class="elsevierStyleHsp" style=""></span>cm wide&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; pringlei</span></p></li></ul></p><p id="par0290" class="elsevierStylePara elsevierViewall">5&#8217;&#46; Prickles on cone scale umbo deciduous&#46; &#46;&#46; 7<ul class="elsevierStyleList" id="lis0035"><li class="elsevierStyleListItem" id="lsti0035"><span class="elsevierStyleLabel">7&#46;</span><p id="par0160" class="elsevierStylePara elsevierViewall">Needle length 14&#46;5&#8211;19<span class="elsevierStyleHsp" style=""></span>cm&#44; 4&#8211;7 rows of stomata on abaxial face&#44; 4&#8211;8 adaxial&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; herrerae</span></p><p id="par0165" class="elsevierStylePara elsevierViewall">7&#8217;&#46; Needle length 9&#8211;16<span class="elsevierStyleHsp" style=""></span>cm&#44; 8&#8211;10&#40;&#8211;12&#41; rows of stomata on abaxial face&#44; 10 adaxial&#46; &#46;&#46; <span class="elsevierStyleItalic">P&#46; teocote</span></p></li></ul></p></li></ul></p></li></ul></p></li></ul></p></li></ul></p></li></ul></p></span><span id="sec0030" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0050">Intra- and inter-specific comparison of plastid DNA sequences and haplotypes</span><p id="par0170" class="elsevierStylePara elsevierViewall">The complete plastid DNA sequence matrix of 49 individuals had a total of seven variable sites&#59; six of the site variants were present in more than one individual &#40;parsimony informative&#41;&#46; Nucleotide diversity &#40;&#960;&#41; for all species considered together was 0&#46;00191&#46; The haplotype diversity &#40;Hd&#41; was 0&#46;732&#46; The seven variable sites were distributed among nine different haplotypes&#44; five of which were shared by two or more species &#40;<a class="elsevierStyleCrossRef" href="#tbl0015">Table III</a>&#41;&#46;</p><elsevierMultimedia ident="tbl0015"></elsevierMultimedia><p id="par0175" class="elsevierStylePara elsevierViewall">The sequences of all the individuals were used to construct a haplotype network &#40;<a class="elsevierStyleCrossRef" href="#fig0015">Figure 3</a>&#41;&#46; Haplotype 9 occurred at the highest frequency &#40;n &#61; 23&#41; and was shared by the four species <span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; <span class="elsevierStyleItalic">P&#46; oocarpa</span>&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span>&#44; and <span class="elsevierStyleItalic">P&#46; teocote</span>&#46; It was near the center of the network&#44; with three connections to other haplotypes&#46; The same four species also shared the second most common haplotype &#40;haplotype 8&#59; n &#61; 9&#41;&#46; Only one of these four species&#44; <span class="elsevierStyleItalic">P&#46; oocarpa</span>&#44; shared a haplotype with other species in the area&#44; haplotype 7 &#40;n &#61; 6&#41; with <span class="elsevierStyleItalic">P&#46; leiophylla</span> and <span class="elsevierStyleItalic">P&#46; herrerae</span>&#46; In contrast&#44; the two varieties of <span class="elsevierStyleItalic">P&#46; patula</span> only shared haplotypes with each other and with <span class="elsevierStyleItalic">P&#46; herrerae</span>&#59; Haplotype 6&#44; the fourth most frequent in the network &#40;n &#61; 5&#41;&#44; was shared among these three taxa&#46; Haplotype 5 &#40;n &#61; 2&#41; was made up of sequences from the two varieties of <span class="elsevierStyleItalic">P&#46; patula</span>&#59; it had the most connections &#40;four&#41; to other haplotypes in the network&#46; The remaining four haplotypes&#44; found in <span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; <span class="elsevierStyleItalic">P&#46; teocote</span>&#44; <span class="elsevierStyleItalic">P&#46; patula</span>&#44; and <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&#44; each only occurred once&#46;</p><elsevierMultimedia ident="fig0015"></elsevierMultimedia></span><span id="sec0035" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0055">DISCUSSION</span><p id="par0180" class="elsevierStylePara elsevierViewall">Morphology is central for the study of taxonomy&#46; It permits the documentation of taxonomic limits&#44; geographic distribution&#44; and phenotypic variation&#46; It is often also useful for the detection of hybrids&#46; In pines&#44; a diversity of leaf types&#44; notably bracts along the branches&#44; fascicle sheaths&#44; and secondary needles&#44; are useful for identifying species&#44; as are seed cone size and shape&#44; length of the peduncle&#44; and morphology of the cone scale apex&#46; Needle anatomy has long been used to delimit or identify species of pines<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6&#44;11&#44;12</span></a>&#46; Careful study of morphology and needle anatomy has the potential of identifying putative natural hybrids&#46; The distribution of the species studied here has been described previously<a class="elsevierStyleCrossRefs" href="#bib0155"><span class="elsevierStyleSup">31&#44;32</span></a>&#44; but finer scale information is still lacking&#46; Further field exploration and herbarium work are needed to better establish the geographic distribution of these species in southern Mexico and to capture the full range of morphological and molecular variation of species with ranges that extend beyond Guerrero and Oaxaca&#46;</p><p id="par0185" class="elsevierStylePara elsevierViewall">Forest management practices may be altering the geographic distribution of tree taxa in the area&#59; it is important to establish their natural distributions and determine if interspecific gene flow is being promoted by human activities&#44; particularly the planting of economically important species outside of their natural range of distribution&#46;</p><p id="par0190" class="elsevierStylePara elsevierViewall">The 840 b&#46;p&#46; fragment of plastid DNA characterized in this study offers corroborating evidence of shared plastid DNA haplotypes in North American pines<a class="elsevierStyleCrossRefs" href="#bib0085"><span class="elsevierStyleSup">17&#8211;23</span></a>&#46; A previous study that included a 5&#44;425 b&#46;p&#46; plastid DNA alignment for 3&#8211;6 individuals per species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> recovered all haplotypes as monophyletic by species for <span class="elsevierStyleItalic">P&#46; herrerae</span> &#40;three individuals&#41;&#44; <span class="elsevierStyleItalic">P&#46; leiophylla</span> &#40;four individuals&#41;&#44; and <span class="elsevierStyleItalic">P&#46; teocote</span> &#40;three individuals&#41;<a class="elsevierStyleCrossRef" href="#bib0090"><span class="elsevierStyleSup">18</span></a>&#46; However&#44; in this study we report a <span class="elsevierStyleItalic">P&#46; herrerae</span> individual &#40;and a <span class="elsevierStyleItalic">P&#46; oocarpa</span> individual&#41; with the typical haplotype of <span class="elsevierStyleItalic">P&#46; leiophylla</span>&#44; and two haplotypes in <span class="elsevierStyleItalic">P&#46; teocote</span> that are widespread in other species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span>&#46; Further evidence of shared haplotypes&#44; previously reported at a broader geographical scale<a class="elsevierStyleCrossRef" href="#bib0090"><span class="elsevierStyleSup">18</span></a>&#44; was also found within the study region for <span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; <span class="elsevierStyleItalic">P&#46; oocarpa</span>&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span>&#44; and <span class="elsevierStyleItalic">P&#46; teocote</span>&#46;</p><p id="par0195" class="elsevierStylePara elsevierViewall">Sharing of plastid haplotypes may be widespread among closely related species of conifers and some other tree genera&#46; The two processes most likely responsible for this phenomenon in pines are introgressive hybridization and incomplete lineage sorting&#46; These processes are probably most widespread in groups of recently diverged species&#44; such as within <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Ponderosae</span> and <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span><a class="elsevierStyleCrossRef" href="#bib0090"><span class="elsevierStyleSup">18</span></a>&#44; or in young putative species pairs&#44; such as the Mexican white pines&#44; <span class="elsevierStyleItalic">P&#46; ayacahuite</span> and <span class="elsevierStyleItalic">P&#46; strobiformis</span><a class="elsevierStyleCrossRef" href="#bib0110"><span class="elsevierStyleSup">22</span></a>&#46; Less plastid haplotype sharing has been reported from other conifer lineages with fewer recently diverged species such as North American pinyon pines<a class="elsevierStyleCrossRefs" href="#bib0205"><span class="elsevierStyleSup">41&#44;42</span></a>&#46; Some recently diverged lineages&#44; such as <span class="elsevierStyleItalic">P&#46; caribaea</span> of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span><a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">43</span></a> also show little evidence of shared haplotypes&#44; possibly due to less co-occurrence with close relatives and smaller effective population sizes&#46; Distinguishing between introgressive hybridization and incomplete lineage sorting can be achieved by comparing the genealogical patterns of plastid DNA with those of nuclear and mitochondrial DNA&#44; both of which are likely to result in different patterns than those found with plastid DNA&#46; Further evidence of hybridization can also be documented through morphological and anatomical studies&#46;</p></span><span id="sec0040" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0060">CONCLUSIONS</span><p id="par0200" class="elsevierStylePara elsevierViewall">Guerrero and Oaxaca are rich in species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span>&#46; They occur both allopatrically and sympatrically throughout the montane regions of the state&#46; Although they can be distinguished morphologically&#44; they follow a more generalized pattern in North American hard pines of sharing plastid DNA lineages as a result of introgressive hybridization or incomplete lineage sorting&#46; Inferring plastid DNA genealogies is of unquestionable value for studying evolution in the group&#44; but it is important to be aware that adequate infraspecific sampling and inclusion of sufficient variable sites are needed for plastid DNA genealogies to be informative with respect to species relationships&#46;</p><p id="par0205" class="elsevierStylePara elsevierViewall">Future studies should incorporate range-wide sampling of species to capture their full range of variation&#46; Sequences from multiple unlinked markers combined with morphometric analysis should permit more accurate detection of interspecific gene flow in the presence of incomplete lineage sorting&#46;</p></span></span>"
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          "titulo" => "INTRODUCTION"
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          "titulo" => "MATERIALS AND METHODS"
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          "titulo" => "RESULTS"
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              "titulo" => "Morphology and leaf anatomy"
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          "identificador" => "sec0025"
          "titulo" => "Identification key for Pinus subsection Australes in Guerrero and Oaxaca"
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          "titulo" => "Intra- and inter-specific comparison of plastid DNA sequences and haplotypes"
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          "titulo" => "DISCUSSION"
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          "titulo" => "ACKNOWLEDGEMENTS"
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    "fechaRecibido" => "2015-09-14"
    "fechaAceptado" => "2016-04-27"
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          "palabras" => array:5 [
            0 => "biodiversity"
            1 => "gene flow"
            2 => "lineage sorting"
            3 => "pines"
            4 => "species delimitation"
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          "titulo" => "Palabras clave"
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          "palabras" => array:5 [
            0 => "biodiversidad"
            1 => "flujo g&#233;nico"
            2 => "sorteo de linajes"
            3 => "pinos"
            4 => "delimitaci&#243;n de especies"
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        "titulo" => "ABSTRACT"
        "resumen" => "<span id="abst0005" class="elsevierStyleSection elsevierViewall"><p id="spar0005" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> 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&#46; Previous studies have shown that some species of this subsection share plastid DNA haplotypes&#44; a pattern that is attributed to introgressive hybridization or the retention of ancestral polymorphisms&#46; Here we describe the morphological and plastid haplotype diversity for this group of species in the states of Guerrero and Oaxaca&#44; Mexico&#46; Seven species of <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> are recognized in the study area&#44; one of which&#44; <span class="elsevierStyleItalic">P&#46; patula</span>&#44; includes two varieties&#46; Seven variable sites and nine haplotypes were found in an 840 b&#46;p&#46; fragment of the DNA coding region <span class="elsevierStyleItalic">ycf1</span>&#46; Shared haplotypes were found for <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">patula</span>&#44; <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&#44; <span class="elsevierStyleItalic">P&#46; herrerae</span>&#44; and <span class="elsevierStyleItalic">P&#46; tecunumanii</span>&#46; Four of the nine haplotypes found were restricted to Oaxaca&#46; Although plastid DNA genealogies are valuable for studying evolution in this group&#44; greater sampling of individuals and the inclusion of more variable sites are needed to more accurately infer species relationships&#46;</p></span>"
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      "es" => array:2 [
        "titulo" => "RESUMEN"
        "resumen" => "<span id="abst0010" class="elsevierStyleSection elsevierViewall"><p id="spar0010" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleItalic">Pinus</span> subsecci&#243;n <span class="elsevierStyleItalic">Australes</span> es un grupo de pinos duros de Am&#233;rica del Norte que comprende aproximadamente 29 especies de &#225;rboles importantes econ&#243;micamente y ecol&#243;gicamente distribuidos a lo largo de toda Am&#233;rica del Norte y Central y las Islas Caribe&#241;as&#46; Estudios previos han mostrado que las especies de esta subsecci&#243;n a menudo comparten haplotipos de ADN de plastidio&#44; un patr&#243;n que es atribuido a la hibridaci&#243;n introgresiva y la retenci&#243;n de polimorfismos ancestrales&#46; Aqu&#237; describimos la diversidad de haplotipos de plastidio y la morfolog&#237;a para este grupo de especies en los estados de Guerrero y Oaxaca&#44; M&#233;xico&#46; Siete especies de <span class="elsevierStyleItalic">Pinus</span> subsecci&#243;n <span class="elsevierStyleItalic">Australes</span> son reconocidas en el &#225;rea de estudio&#44; una de las cuales&#44; <span class="elsevierStyleItalic">P&#46; patula</span> incluye dos variedades&#46; Siete sitios variables y nueve haplotipos fueron encontrados amplificando un fragmento de 840 p&#46; b&#46; de ADN de la regi&#243;n codificante <span class="elsevierStyleItalic">ycf1</span>&#46; Se encontraron haplotipos compartidos para <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">patula</span>&#44; <span class="elsevierStyleItalic">P</span>&#46; <span class="elsevierStyleItalic">patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&#44; <span class="elsevierStyleItalic">P&#46; herrerae</span> y <span class="elsevierStyleItalic">P&#46; tecunumanii</span>&#46; Cuatro de los nueve haplotipos encontrados est&#225;n restringidos a Oaxaca&#46; Aunque las genealog&#237;as de genes son valiosas para estudiar la evoluci&#243;n de este grupo&#44; se requieren mayor muestreo de individuos y m&#225;s sitios variables para la inferencia de relaciones entre las especies&#46;</p></span>"
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    "apendice" => array:1 [
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          0 => array:4 [
            "apendice" => "<p id="par0215" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus herrerae</span></span><span class="elsevierStyleBold">Mart&#237;nez</span></p> <p id="par0220" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> Loma El Ocote&#44; I&#46; Mart&#237;nez Mart&#237;nez 71204&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; Cerro La Mula&#44; E&#46;O&#46; Valencia Santana 71124&#46; Cerro Fais&#225;n&#44; M&#46;A&#46; Ju&#225;rez 72227&#46; Omiltemi&#44; Chilpancingo&#44; J&#46; A&#46; P&#233;rez de la Rosa 132&#46; Camino Para&#237;so&#44; L&#46; Lozada 182A&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0225" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus lawsonii</span></span><span class="elsevierStyleBold">Roezl ex Gordon</span></p> <p id="par0230" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> Arroyo Cangarec&#44; O&#46; Rodr&#237;guez Santamar&#237;a 74754&#46; La Ocotera&#44; O&#46; Rodr&#237;guez Santamar&#237;a 73947&#46; Arroyo Artemio Reyes&#44; G&#46;R&#46; Izaguirre Y&#225;&#241;ez 76960&#46; Barranca Pelona&#44; J&#46;C&#46; Calvillo Garc&#237;a 71396&#46; El Zacat&#243;n&#44; J&#46; Z&#250;&#241;iga Reyes 74549&#46; Agua Cola&#44; J&#46; Z&#250;&#241;iga 74747&#46; Pino Gordo&#44; O&#46; J&#46; P&#233;rez Mart&#237;nez 72853&#46; Nanche&#44; R&#44; Boquedano Peralta 76955&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; Exsuyo&#44; J&#46;C&#46; Calvillo Garc&#237;a 71587&#46; La Laguna&#44; J&#46;C&#46; Calvillo Garc&#237;a 70214&#46; El Chaude&#44; A&#46;J&#46; Fortoul Velasco 70254&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0235" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus leiophylla</span></span><span class="elsevierStyleBold">Schiede ex Schltdl&#46; &#38; Cham&#46;</span></p> <p id="par0240" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> La Plaga&#44; R&#46; Baquedano Peralta 72700&#46; Monte de Tesoro&#44; J&#46; Reyes Santiago 883&#46; Miahuatl&#225;n&#44; E&#46; Hunn 0096&#46; Santa Mar&#237;a del Rosario&#44; A&#46;J&#46; Fortoul Velasco 72252&#46; Ixtl&#225;n&#44; A&#46; Garc&#237;a-Mendoza 7585&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; La Guitarra&#44; A&#46;J&#46; Fortoul Velasco 71557&#46; La Culebra&#44; A&#46;J&#46; Fortoul Velasco 71773&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0245" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus oocarpa</span></span><span class="elsevierStyleBold">Schiede ex Schltdl&#46;</span></p> <p id="par0250" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca</span>&#58; El Corol&#243;n-El Ret&#233;n&#44; T&#46; Yescas de los &#193;ngeles 75410&#46; Agua Fr&#237;a&#44; T&#46; Yescas de los &#193;ngeles 75712&#46; Loma del Chifle&#44; T&#46; Yescas de los &#193;ngeles 76958&#46; La Ermita&#44; J&#46;L&#46; Flores Nicanor 71650&#46; La Cascada&#44; E&#46; Mart&#237;nez V&#46; 73980&#44; Rancho Viejo&#44; T&#46; Yescas de los &#193;ngeles 72018&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; La Sidra&#44; J&#46;C&#46; Bautista Mart&#237;nez 71991&#46; Agua de Obispo&#44; J&#46;C&#46; Calvillo Garc&#237;a 72424&#46; La Coscolina&#44; O&#46;J&#46; P&#233;rez Mart&#237;nez 72838&#46; El Ermita&#241;o&#44; J&#46;C&#46; Calvillo Garc&#237;a 70448&#46; Tierra Blanca&#44; J&#46;C&#46; Calvillo Garc&#237;a 72023&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0255" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus patula</span></span><span class="elsevierStyleBold">Schltdl&#46; &#38; Cham&#46; var&#46;</span><span class="elsevierStyleItalic"><span class="elsevierStyleBold">patula</span></span></p> <p id="par0260" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> Terracer&#237;a La Venta&#44; J&#46; Reyes 6493&#46; La Cima&#44; M&#46;I&#46; Alvarado Flores 72489&#46; Lomatico&#44; J&#46;C&#46; Bautista Mart&#237;nez 73310&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0265" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus patula</span></span><span class="elsevierStyleBold">var&#46;</span><span class="elsevierStyleItalic"><span class="elsevierStyleBold">longipedunculata</span></span><span class="elsevierStyleBold">Loock ex Mart&#237;nez</span></p> <p id="par0270" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> Aserrado Las Vigas&#44; J&#46; A&#46; P&#233;rez de la Rosa 1849&#46; San Juan Tepeuxila&#44; R&#46; Torres Col&#237;n 16168&#46; Santa Mar&#237;a Yaves&#237;a&#44; I&#46; Trejo 3064&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0275" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus pringlei</span></span><span class="elsevierStyleBold">Shaw</span></p> <p id="par0280" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleBold">M&#233;xico&#46; Oaxaca&#46;</span> Nebr&#243;n&#44; J&#46; Z&#250;&#241;iga Reyes 71153&#46; La Caoilla&#44; Ojo de Agua&#44; M&#46; I&#46; Alvarado Flores 69610&#46; El Chaneque&#44; J&#46;L&#46; Flores Nicanor 75148&#46; Silacayuapan&#44; R&#46; Luna Mart&#237;nez 71600&#46; Tres Cruces&#44; J&#46;L&#46; Flores Nicanor 74544&#46; San Isidro El Chi&#241;&#243;n&#44; R&#46; Luna Mart&#237;nez 71813&#46; Buenavista&#44; O&#46; Rodr&#237;guez Santamar&#237;a 70069&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; Cajones&#44; J&#46;C&#46; Calvillo Garc&#237;a 65501&#46; Cocuitlazola&#44; J&#46;C&#46; Calvillo Garc&#237;a 72446&#46; C&#46; Tlacholoya&#44; J&#46;C&#46; Calvillo Garc&#237;a 71371&#46; Ejido El Carrazal&#44; Coyuca de Catal&#225;n&#44; E&#46; Rodr&#237;guez Ibarra 69525&#46; Huexoapa&#44; F&#46; Rosas Aguilar 72658&#46;<span class="elsevierStyleVsp" style="height:0.5px"></span></p> <p id="par0285" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic"><span class="elsevierStyleBold">Pinus teocote</span></span><span class="elsevierStyleBold">Schiede ex Schltdl&#46; &#38; Cham&#46;M&#233;xico&#46; Oaxaca&#46;</span> Agua Colorada&#44; I&#46; Mart&#237;nez Mart&#237;nez 74372&#46; Cerro Manteca&#44; J&#46;L&#46; Flores Nicanor 75345&#46; Yacuni&#44; M&#46;I&#46; Alvarado Flores 72898&#46; La Ci&#233;naga&#44; J&#46;L&#46; Flores Nicanor 73537&#44; La Joya del Fresno&#44; T&#46; Yescas de los &#193;ngeles 72476&#46; R&#237;o Milpas&#44; R&#46; Baquedano Peralta 73534&#46; Barranca del Zopilote&#44; J&#46;C&#46; Calvillo Garc&#237;a 71607&#46; San Francisco Higos&#44; R&#46; Lina Mart&#237;nez 72028&#46; <span class="elsevierStyleBold">Guerrero</span>&#46; Mesa de la Mujer&#44; A&#46;J&#46; Fortoul Velasco 68303&#46; San Andr&#233;s&#44; A&#46;J&#46; Fortoul Velasco72003&#46;</p>"
            "etiqueta" => "APPENDIX 1"
            "titulo" => "SPECIMENS EXAMINED&#46; ALL ARE DEPOSITED IN MEXU&#46;"
            "identificador" => "sec0045"
          ]
        ]
      ]
    ]
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        "etiqueta" => "Figure 1"
        "tipo" => "MULTIMEDIAFIGURA"
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        "mostrarDisplay" => false
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          "en" => "<p id="spar0015" class="elsevierStyleSimplePara elsevierViewall">The collection localities for the individuals included in this study&#46;</p>"
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      1 => array:7 [
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        "etiqueta" => "Figure 2"
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          "en" => "<p id="spar0020" class="elsevierStyleSimplePara elsevierViewall">Morphological diversity of seed cones for <span class="elsevierStyleItalic">Pinus</span> subsection <span class="elsevierStyleItalic">Australes</span> in Guerrero and Oaxaca&#46; A&#46; <span class="elsevierStyleItalic">Pinus herrerae</span> Gonz&#225;lez &#38; Mart&#237;nez 770&#46; B&#46; <span class="elsevierStyleItalic">Pinus lawsonii</span> Soto N&#250;&#241;ez <span class="elsevierStyleItalic">et al&#46;</span>&#44; 5307&#46; C&#46; <span class="elsevierStyleItalic">Pinus leiophylla</span> Reyes 1989&#46; D&#46; <span class="elsevierStyleItalic">Pinus oocarpa</span> Yescas de los &#193;ngeles 75712&#46; E&#46; <span class="elsevierStyleItalic">Pinus patula</span> var&#46; patula Bautista Mart&#237;nez 73310&#46; F&#46; <span class="elsevierStyleItalic">Pinus patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span> Trejo 3053&#46; G&#46; <span class="elsevierStyleItalic">Pinus pringlei</span> Rico 489&#46; H&#46; <span class="elsevierStyleItalic">Pinus teocote</span> Rom&#225;n 335&#46; Photographs by Carmen Loyola&#46;</p>"
        ]
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        "etiqueta" => "Figure 3"
        "tipo" => "MULTIMEDIAFIGURA"
        "mostrarFloat" => true
        "mostrarDisplay" => false
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        "descripcion" => array:1 [
          "en" => "<p id="spar0025" class="elsevierStyleSimplePara elsevierViewall">Plastid DNA haplotype network for the study species&#46; The area of the circles is proportional to haplotype frequency&#46;</p>"
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      ]
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        "etiqueta" => "Table I"
        "tipo" => "MULTIMEDIATABLA"
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        "detalles" => array:1 [
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            "detalle" => "Table "
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                  <table border="0" frame="\n
                  \t\t\t\t\tvoid\n
                  \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Taxon&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Fascicle sheath&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Fascicle sheath length &#40;cm&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Leaves per fascicle&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Leaf length &#40;cm&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Number of abaxial stomatal rows&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Peduncle length &#40;cm&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Cone length &#40;cm&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Umbo shape&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; herrerae</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;6&#8211;1&#46;7&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#40;&#8211;4&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">14&#46;5&#8211;19&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&#8211;7&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;4&#8211;1&#46;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&#46;5&#8211;5&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">slightly raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; lawsonii</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#8211;2&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#40;&#8211;4&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">11&#8211;27&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">7&#8211;10&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#8211;1&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&#8211;8&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">flat to slightly raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; leiophylla</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">deciduous&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">7&#8211;13&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;8&#8211;1&#46;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&#46;5&#8211;6&#46;2&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">flat to slightly raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; oocarpa</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#8211;2&#46;2&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">14&#46;5&#8211;29&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">&#40;3&#8211;&#41;4&#8211;6&#40;&#8211;14&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#46;5&#8211;3&#46;0&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&#46;5&#8211;6&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">usually flat&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; patula</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#46;1&#8211;1&#46;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#8211;4&#40;&#8211;5&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">12&#8211;22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;5&#8211;1&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">7&#8211;7&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">flat or slightly raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#8211;1&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#8211;4&#40;&#8211;5&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">14&#8211;24&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#8211;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;8&#8211;1&#46;4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&#8211;8&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">flat or slightly raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; pringlei</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&#46;6&#8211;2&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#40;&#8211;4&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">15&#8211;32&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">8&#8211;11&#40;&#8211;15&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;5&#8211;2&#46;2&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&#8211;10&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">slightly raised to subpyramidal&#44;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; teocote</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">persistent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;5&#8211;1&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#40;&#8211;4&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">9&#8211;16&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">8&#8211;10&#40;&#8211;12&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">0&#46;7&#8211;1&#46;2&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&#8211;6&#46;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">flat or raised&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr></tbody></table>
                  """
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        "descripcion" => array:1 [
          "en" => "<p id="spar0030" class="elsevierStyleSimplePara elsevierViewall">Morphological characters of <span class="elsevierStyleItalic">Pinus subsection Australes</span> species in Guerrero and Oaxaca&#46;</p>"
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                  <table border="0" frame="\n
                  \t\t\t\t\tvoid\n
                  \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Taxon&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Hypodermal cell shape&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Hypodermal cell intrusions into mesophyll&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Number of resin canals&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Resin canal position&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; herrerae</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">biforme&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">absent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; lawsonii</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">multiforme&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">present&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&#8211;4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; leiophylla</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">absent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&#8211;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; oocarpa</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">absent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">septal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; patula</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick or multiforme&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">present&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&#8211;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">medial and internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick or multiforme&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">present&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&#8211;3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">medial and internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; pringlei</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">biforme or multiforme&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">present&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&#8211;5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; tecunumanii</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">absent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">medial&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; teocote</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">thick&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">absent&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">3&#8211;4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">internal&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr></tbody></table>
                  """
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          "en" => "<p id="spar0035" class="elsevierStyleSimplePara elsevierViewall">Leaf anatomical characters of <span class="elsevierStyleItalic">Pinus subsection Australes</span> species in Guerrero and Oaxaca&#46; <span class="elsevierStyleItalic">Pinus tecunumanii</span> is also included for comparison &#40;see text&#41;&#46;</p>"
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                0 => """
                  <table border="0" frame="\n
                  \t\t\t\t\tvoid\n
                  \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td-with-role" title="table-head ; entry_with_role_rowhead " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Haplotype Number&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">208&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">311&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">326&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">470&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">717&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">797&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">819&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">N&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="top" scope="col" style="border-bottom: 2px solid black">Species&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">9&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">23&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; <span class="elsevierStyleItalic">P&#46; oocarpa</span>&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span>&#44; <span class="elsevierStyleItalic">P&#46; teocote</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">8&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">9&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; lawsonii</span>&#44; <span class="elsevierStyleItalic">P&#46; oocarpa</span>&#44; <span class="elsevierStyleItalic">P&#46; pringlei</span>&#44; <span class="elsevierStyleItalic">P&#46; teocote</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">7&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">6&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; herrerae</span>&#44; <span class="elsevierStyleItalic">P&#46; leiophylla&#44; P&#46; oocarpa</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">6&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; herrerae</span>&#44; <span class="elsevierStyleItalic">P&#46; patula&#44; P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">5&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">2&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">P&#46; patula&#44; <span class="elsevierStyleItalic">P&#46; patula</span> var&#46; <span class="elsevierStyleItalic">longipedunculata</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">4&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">G&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">1&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top"><span class="elsevierStyleItalic">P&#46; patula</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="table-entry ; entry_with_role_rowhead " align="center" valign="top">3&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">T&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="top">C&nbsp;\t\t\t\t\t\t\n
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Article information

ISSN: 1405888X
Original language: English

DOI:10.1016/j.recqb.2016.06.002

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Year/Month	Html	Pdf	Total

2024 November	5	0	5
2024 October	28	2	30
2024 September	22	0	22
2024 August	22	1	23
2024 July	20	5	25
2024 June	13	5	18
2024 May	17	3	20
2024 April	11	2	13
2024 March	25	7	32
2024 February	15	2	17
2024 January	21	5	26
2023 December	24	7	31
2023 November	17	12	29
2023 October	20	9	29
2023 September	22	3	25
2023 August	10	4	14
2023 July	12	3	15
2023 June	30	7	37
2023 May	53	6	59
2023 April	28	5	33
2023 March	24	4	28
2023 February	35	3	38
2023 January	39	4	43
2022 December	39	3	42
2022 November	46	16	62
2022 October	30	8	38
2022 September	21	2	23
2022 August	15	10	25
2022 July	18	12	30
2022 June	28	3	31
2022 May	14	8	22
2022 April	20	6	26
2022 March	38	12	50
2022 February	29	3	32
2022 January	25	14	39
2021 December	25	9	34
2021 November	27	6	33
2021 October	40	12	52
2021 September	16	7	23
2021 August	26	5	31
2021 July	23	9	32
2021 June	20	13	33
2021 May	15	7	22
2021 April	81	17	98
2021 March	54	8	62
2021 February	39	7	46
2021 January	27	19	46
2020 December	25	5	30
2020 November	16	3	19
2020 October	13	4	17
2020 September	21	6	27
2020 August	14	11	25
2020 July	15	12	27
2020 June	22	7	29
2020 May	34	10	44
2020 April	27	2	29
2020 March	30	5	35
2020 February	27	3	30
2020 January	22	8	30
2019 December	31	5	36
2019 November	24	12	36
2019 October	30	1	31
2019 September	34	8	42
2019 August	30	6	36
2019 July	30	2	32
2019 June	68	15	83
2019 May	125	23	148
2019 April	67	10	77
2019 March	17	6	23
2019 February	14	3	17
2019 January	17	3	20
2018 December	12	3	15
2018 November	15	5	20
2018 October	24	8	32
2018 September	29	5	34
2018 August	18	6	24
2018 July	10	0	10
2018 June	9	6	15
2018 May	18	1	19
2018 April	18	1	19
2018 March	9	0	9
2018 February	12	0	12
2018 January	13	0	13
2017 December	5	0	5
2017 November	11	1	12
2017 October	12	2	14
2017 September	15	1	16
2017 August	8	2	10
2017 July	21	1	22
2017 June	18	1	19
2017 May	20	11	31
2017 April	17	1	18
2017 March	14	28	42
2017 February	43	2	45
2017 January	17	1	18
2016 December	32	6	38
2016 November	39	11	50
2016 October	44	6	50
2016 September	49	3	52
2016 August	28	3	31
2016 July	6	2	8

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