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Panbiogeography of the Santa María Amajac area, Hidalgo, Mexico
Panbiogeografía del área de Santa María Amajac, Hidalgo, México
Arturo Palma-Ramírez1, Irene Goyenechea2, Jesús M. Castillo-Cerón3
1 Léxico Estratigráfico, Servicio Geológico Mexicano. Av. Mariano Jiménez 465, 78280 San Luis Potosí, San Luis Potosí, Mexico
2 Laboratorio de Sistemática Molecular, Universidad Autónoma del Estado de Hidalgo. Ciudad Universitaria, Carretera Pachuca Tulancingo s/n Km. 4.5, 42184 Mineral de la Reforma, Hidalgo, Mexico
3 Museo de Paleontología, Universidad Autónoma del Estado de Hidalgo. Ciudad Universitaria, Carretera Pachuca Tulancingo s/n Km. 4.5, 42184 Mineral de la Reforma, Hidalgo, Mexico
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    "textoCompleto" => "<span class="elsevierStyleSections"><span id="sec0005" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0025">Introduction</span><p id="par0005" class="elsevierStylePara elsevierViewall">Traditionally&#44; panbiogeographic studies have had a neontological approach&#46; The underestimation or exclusion of the fossil record in this research program has been pointed out by different authors &#40;<a class="elsevierStyleCrossRef" href="#bib0085">Crisci and Morrone&#44; 1992</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0070">Cox and Moore&#44; 1993</a>&#41;&#46; Nevertheless&#44; <a class="elsevierStyleCrossRef" href="#bib0075">Craw et al&#46; &#40;1999&#41;</a> mentioned that fossils are important in panbiogeography because they could extend the distributional area of taxa&#44; and represent the minimal age of a group&#44; which is not necessarily equivalent to the time of the phylogenetic origin of that group&#46; This contrasts with the traditional dispersalist point of view&#44; where biogeographic hypotheses are based upon the place of first occurrence of a group in the fossil record &#40;<a class="elsevierStyleCrossRef" href="#bib0035">Candela and Morrone&#44; 2003</a>&#41;&#46; In Mexico&#44; the fossil record is abundant and there are localities ranging from the Precambrian &#40;541<span class="elsevierStyleHsp" style=""></span>million years ago&#41; to the Holocene &#40;Recent&#41;&#46; However&#44; given the geological history of the country and the nature of the fossil record&#44; some periods and environments are represented more frequently than others&#44; and some biological groups are widely documented&#44; while in others the information is scarce or absent &#40;<a class="elsevierStyleCrossRef" href="#bib0025">Arroyo-Cabrales et al&#46;&#44; 2008</a>&#41;&#46; Lacustrine environments are distinct because their biota could be preserved in the fossil record &#40;<a class="elsevierStyleCrossRef" href="#bib0025">Arroyo-Cabrales et al&#46;&#44; 2008</a>&#41;&#46; In Mexico&#44; there is no marine<a name="p1229"></a> or terrestrial fossil record revealing the total biological richness of a specific place as a reference to understanding current diversity &#40;<a class="elsevierStyleCrossRef" href="#bib0025">Arroyo-Cabrales et al&#46;&#44; 2008</a>&#41;&#46; The Santa Mar&#237;a Amajac paleolake is located in the central part of Hidalgo&#44; and the paleofauna has been the subject of several studies&#46; Its fossils include plants&#44; ostracods&#44; gastropods&#44; fishes&#44; amphibians&#44; and mammals&#44; among other groups &#40;<a class="elsevierStyleCrossRef" href="#bib0010">Aguilar and Ortiz&#44; 2000</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0015">Aguilar et al&#46;&#44; 2002</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0020">Aguilar-Aguilar and Contreras-Medina&#44; 2003</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0170">Mart&#237;nez-Becerra&#44; 2003</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0145">Hern&#225;ndez-L&#243;pez&#44; 2006</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0180">Mart&#237;nez-Mart&#237;nez and Velasco de Le&#243;n&#44; 2006</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0115">Flores-Camargo et al&#46;&#44; 2009</a>&#44; <a class="elsevierStyleCrossRef" href="#bib0215">Palma-Ram&#237;rez&#44; 2012</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0245">Zayas-Ocelotl&#44; 2013</a>&#41;&#46;</p><p id="par0010" class="elsevierStylePara elsevierViewall">The aim of this study was to determine distributional patterns of different taxa making up the paleobiota in the Santa Mar&#237;a Amajac paleolake through a panbiogeographic approach&#46; This work represents the first analysis of its kind in Mexico that uses a significant number of fossil taxa under a panbiogeographical approach&#46; The panbiogeographic method was originally proposed by <a class="elsevierStyleCrossRef" href="#bib0090">Leon Croizat &#40;1958&#41;</a> with the statement &#8220;Earth and Life evolve together&#8221;&#44; meaning that there is a correlated history among 3 factors working as one&#58; space&#44; time&#44; and form &#40;<a class="elsevierStyleCrossRef" href="#bib0185">Morrone&#44; 2001</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; Panbiogeography emphasizes the spatial or geographical dimension of biodiversity&#44; which allows a better understanding of evolutionary patterns and processes &#40;<a class="elsevierStyleCrossRef" href="#bib0075">Craw et al&#46;&#44; 1999</a>&#41;&#46; Also&#44; it highlights the importance of geographical distribution in maps &#40;<a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; This method identifies spatial homologies&#44; which indicate the preexistence of ancestral biotas &#40;<a class="elsevierStyleCrossRef" href="#bib0130">Grehan&#44; 2001</a>&#41;&#44; clarifying the interrelationships between biogeographic areas&#44; and also recognizing transition zones &#40;<a class="elsevierStyleCrossRef" href="#bib0225">Ruggiero and Ezcurra&#44; 2003</a>&#41;&#46;</p><p id="par0015" class="elsevierStylePara elsevierViewall">Panbiogeography allows the possibility of recognizing whether certain areas have a complex origin &#40;<a class="elsevierStyleCrossRef" href="#bib0185">Morrone&#44; 2001</a>&#41;&#44; suggesting localities for future cladistic biogeographic analyses &#40;<a class="elsevierStyleCrossRef" href="#bib0060">Contreras-Medina and Eliosa-Le&#243;n&#44; 2001</a>&#41;&#46; Additionally&#44; this method is especially useful in paleontology&#44; where phylogenetic data of most of the groups is dubious or even incomplete &#40;<a class="elsevierStyleCrossRef" href="#bib0125">Gallo et al&#46;&#44; 2013</a>&#41;&#46;</p></span><span id="sec0010" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0030">Materials and methods</span><p id="par0020" class="elsevierStylePara elsevierViewall">An exhaustive search for fossil distributional records in America from the taxa identified in the area of Santa Mar&#237;a Amajac was conducted&#46; The data analyzed corresponded to the ages determined according to the chronology based on the presence of North American land mammals &#40;NALMA&#41;&#44; proposed by different authors &#40;<a class="elsevierStyleCrossRef" href="#bib0055">Castillo-Cer&#243;n et al&#46;&#44; 1996</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0045">Carranza-Casta&#241;eda&#44; 2006</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0215">Palma-Ram&#237;rez&#44; 2012</a>&#41;&#58; Blancan &#40;Late Pliocene&#41;&#44; Irvingtonian &#40;Late Pliocene- Early Pleistocene&#41;&#44; and Rancholabrean &#40;Late Pleistocene&#41;&#46; Geographic distributions were obtained from the literature &#40;<a class="elsevierStyleCrossRef" href="#bib0150">Kurt&#233;n and Anderson&#44; 1980</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0040">Carranza-Casta&#241;eda&#44; 1991</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0055">Castillo-Cer&#243;n et al&#46;&#44; 1996</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0095">Edmund&#44; 1996</a>&#41;&#44; and supplemented by the available records of online databases of several scientific institutions&#44; as well as the website&#58; The Paleobiology Database &#40;<a href="http://www.paleodb.org/cgi-bin/bridge.pl">http&#58;&#47;&#47;www&#46;paleodb&#46;org&#47;cgi-bin&#47;bridge&#46;pl</a>&#41;&#46; Taxa were used at the taxonomic level of genus&#44; as in many cases a specific identification was not possible&#46; If fossils were considered at the specific level&#44; the number of records gathered would diminish considerably&#46;</p><p id="par0025" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">Track analysis</span>&#46; Panbiogeography involves 3 principal concepts&#58; individual track&#44; generalized or standard track&#44; and node &#40;<a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; An individual track represents the spatial coordinates of a species or group of related species in space &#40;Liria&#44; 1999&#59; <a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; Operationally&#44; individual tracks are line graphs drawn on a map of connected localities or distributional areas&#44; according to geographical proximity &#40;<a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; Spatially congruent individual tracks of unrelated taxa constitute a generalized track&#44; which provides a spatial criterion for biogeographic homology &#40;<a class="elsevierStyleCrossRef" href="#bib0210">Morrone and Crisci&#44; 1995</a>&#41;&#44; and suggest shared histories within biotas &#40;<a class="elsevierStyleCrossRef" href="#bib0240">Torres-Miranda and Luna-Vega&#44; 2006</a>&#41;&#46; When 2 or more generalized tracks converge or overlap in an area&#44; a biogeographic node is identified&#46; A node is considered a complex area&#46; This means that different ancestral biotic and geologic fragments interrelate in spacetime&#44; as a consequence of terrane collision&#44; docking&#44; or suturing &#40;<a class="elsevierStyleCrossRef" href="#bib0210">Morrone and Crisci&#44; 1995</a>&#41;&#46; Nodes are dynamic biogeographic boundaries where remnant fragments of different ancestral biotas came into contact &#40;<a class="elsevierStyleCrossRef" href="#bib0080">Crisci et al&#46;&#44; 2003</a>&#41;&#46; In this work&#44; we constructed individual tracks for those taxa distributed in the Amajac paleolake &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#41;&#58; 12 plants &#40;11 terrestrial and 1 aquatic&#41;&#44; 3 ostracods&#44; 1 gastropod&#44; 2 amphibians&#44; and 9 mammals&#46; Individual tracks were obtained by plotting localities of each taxon on present-day American continent maps&#44; employing ArcView 3&#46;3 software &#40;<a class="elsevierStyleCrossRef" href="#bib0110">ESRI&#44; 2002</a>&#41;&#44; and connecting them by minimum spanning trees using the Trazos2004 extension &#40;<a class="elsevierStyleCrossRef" href="#bib0220">Rojas-Parra&#44; 2004</a>&#41;&#46; Generalized tracks and biogeographic nodes were drawn by hand&#46; Due to the scale used in this research&#44; individual tracks were not oriented&#44; as has been done in other panbiogeographic studies &#40;<a class="elsevierStyleCrossRef" href="#bib0190">Morrone&#44; 2004</a>&#44; <a class="elsevierStyleCrossRef" href="#bib0200">2009&#41;</a>&#46; Generalized tracks were obtained by superimposing the 27 individual tracks for each genus that correspond to the 27 genera included&#46; Generalized tracks were outlined where 2 or more individual tracks overlap &#40;<a class="elsevierStyleCrossRef" href="#bib0090">Croizat&#44; 1958</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0130">Grehan&#44; 2001</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0190">Morrone&#44; 2004</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0240">Torres-Miranda and Luna-Vega&#44; 2006</a>&#41;&#46; Once the generalized tracks were delimited&#44; nodes were identified in places where 2 or more of these tracks converged&#46;<a name="p1230"></a></p><elsevierMultimedia ident="tbl0005"></elsevierMultimedia></span><span id="sec0015" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0035">Results</span><p id="par0030" class="elsevierStylePara elsevierViewall">We constructed 27 individual tracks for the analyzed taxa &#40;<a class="elsevierStyleCrossRefs" href="#fig0005">Figures 1&#8211;27</a>&#41;&#46; Overlapping of individual tracks allowed the identification of 5 generalized tracks and 4 nodes &#40;<a class="elsevierStyleCrossRef" href="#fig0020">Figure 28</a>&#41;&#46; Four of the generalized tracks are located in northern Mexico and northern and central North America&#46; The fifth generalized track extends from southern Mexico to Central America&#44; and southwestern South America&#46; We list the taxa that support the 5 generalized tracks in <a class="elsevierStyleCrossRef" href="#tbl0010">table 2</a>&#46; The Western North American generalized track &#40;track I&#41; extends from the northwestern United States to California&#44; where it overlaps with one of the edges of the Central North American generalized track &#40;track II&#41;&#59; where the Western North American generalized track turns to Colorado&#44; it reaches Mexico through the Sierra Madre Oriental&#44; and reaches the east-central portion of the Mexican Transvolcanic Belt &#40;MTB&#41;&#44; where it overlaps with generalized tracks II&#44; III&#44; and IV&#46; Both edges of the generalized track II overlie the generalized tracks I&#44; III&#44; and IV in the east-central portion of the MTB&#46; One branch of the generalized track II&#44; extends from southern California&#44; and passes through the west coast of Mexico&#44; while the other edge descends from the northeastern United States across the Gulf of Mexico&#46; The Eastern North American generalized track &#40;track III&#41; comes from the central part of the United States&#44; where it overlies one of the branches<a name="p1231"></a> of the generalized track I&#44; and reaches the central part of Mexico overlapping with the generalized tracks I&#44; II&#44; and IV&#46; The Mesoamerican generalized track &#40;track IV&#41; extends from the southeastern United States and descends to the west-central portion of the MTB&#59; it moves to the east-central part of the MTB&#44; where it overlaps with generalized tracks I&#44; II&#44; and III&#46; Track IV runs to Central America&#44; where it overlies the eastern edge of the South American generalized track &#40;track V&#41;&#46; The latter extends to the Andes&#44; where it bifurcates to the Amazon basin on one side&#44; and to the Andes mountain range on the other&#46; We identified 4 nodes&#46; The first is located in California&#44; United States&#46; The second is situated in the west-central United States&#44; in Colorado&#46; The third is located in the Santa Mar&#237;a Amajac area&#44; in Hidalgo&#44; in the east-central MTB&#46; The fourth is situated in Central America&#44; in Guatemala&#46;</p><elsevierMultimedia ident="fig0005"></elsevierMultimedia><elsevierMultimedia ident="fig0010"></elsevierMultimedia><elsevierMultimedia ident="fig0015"></elsevierMultimedia><elsevierMultimedia ident="fig0020"></elsevierMultimedia><elsevierMultimedia ident="tbl0010"></elsevierMultimedia></span><span id="sec0020" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0040">Discussion</span><p id="par0035" class="elsevierStylePara elsevierViewall">Four of the 5 generalized tracks retrieved by the analysis &#40;<a class="elsevierStyleCrossRef" href="#fig0020">Figure 28</a>&#41; display an east-west distribution pattern that agrees with a previous hypothesis of an Atlantic-Pacific division along the Isthmus of Tehuantepec &#40;<a class="elsevierStyleCrossRef" href="#bib0065">Corona et al&#46;&#44; 2007</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0105">Escalante et al&#46;&#44; 2007</a>&#59; and <a class="elsevierStyleCrossRef" href="#bib0155">Le&#243;n-Paniagua et al&#46;&#44; 2007</a>&#41;&#46; The fifth generalized track &#40;South American&#41; is the only generalized track that resembles the Great American Biotic Interchange between North and South America during the late Cenozoic&#44; where 2 of the fossil mammals that support the track are Holarctic taxa&#44; while <span class="elsevierStyleItalic">Pampatherium mexicanum</span> is a Neotropical taxon&#46;</p><p id="par0040" class="elsevierStylePara elsevierViewall">The 5 generalized tracks retrieved by the analysis coincide partially with those proposed by <a class="elsevierStyleCrossRef" href="#bib0060">Contreras-Medina and Eliosa-Le&#243;n &#40;2001&#41;</a> for the distribution of extant gymnosperms&#44; amphibians&#44; reptiles&#44; birds&#44; mammals&#44; and insects&#46; The Western North American generalized track is congruent with the track that <a class="elsevierStyleCrossRef" href="#bib0175">Mart&#237;nez-Mart&#237;nez &#40;2008&#41;</a> recovered with fossil records of 3 genera of aquatic plants <span class="elsevierStyleItalic">&#40;Nymphaea&#44; Scirpus</span>&#44; and <span class="elsevierStyleItalic">Typha</span>&#41;&#46; This track extends from the western United States to the Santa Mar&#237;a Amajac area&#44; in western Mexico&#46; The Mesoamerican generalized track has been retrieved before and partially coincides with the Southern Mesoamerican track proposed by <a class="elsevierStyleCrossRef" href="#bib0165">M&#225;rquez and Morrone &#40;2003&#41;</a> and with the Mesoamerican track proposed by <a class="elsevierStyleCrossRef" href="#bib0235">Toledo et al&#46; &#40;2007&#41;</a>&#46; The South American generalized track agrees in its Central American portion with the track proposed by <a class="elsevierStyleCrossRef" href="#bib0030">Asiain et al&#46; &#40;2010&#41;</a> based on beetles&#44; with the track proposed by <a class="elsevierStyleCrossRef" href="#bib0175">Mart&#237;nez-Mart&#237;nez &#40;2008&#41;</a> for aquatic plants&#44; and with the track that <a class="elsevierStyleCrossRef" href="#bib0100">Escalante et al&#46; &#40;2011&#41;</a> recovered with nematodes and rodents&#46; The<a name="p1232"></a> latter coincides with the South American generalized track as well&#44; but only as far as Colombia&#46;</p><p id="par0045" class="elsevierStylePara elsevierViewall">One of the nodes retrieved by the analysis&#44; overlapping tracks III and IV&#44; is located within the Mexican Transition zone proposed by <a class="elsevierStyleCrossRef" href="#bib0135">Halffter &#40;1978</a>&#44; <a class="elsevierStyleCrossRef" href="#bib0140">1987&#41;</a> and delimited by <a class="elsevierStyleCrossRef" href="#bib0195">Morrone &#40;2006&#41;</a> using the distribution of extant taxa&#46; These authors proposed the Mexican Transition zone as an area where Nearctic and Neotropical elements from the southern United States&#44; Mexico&#44; and Central America overlap&#46; Other authors &#40;<a class="elsevierStyleCrossRef" href="#bib0005">Aguayo and Tr&#225;paga&#44; 1996</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0240">Torres-Miranda and Luna-Vega&#44; 2006</a>&#41; have already recovered nodes in the MTB&#46;</p><p id="par0050" class="elsevierStylePara elsevierViewall">The biogeographic affinity of the Mexican biota has been discussed previously from a paleontological viewpoint by <a class="elsevierStyleCrossRef" href="#bib0050">Carranza-Casta&#241;eda and Ferrusqu&#237;a-Villafranca&#44; &#40;1978&#41;</a> and <a class="elsevierStyleCrossRef" href="#bib0045">Carranza-Casta&#241;eda&#44; &#40;2006&#41;</a>&#44; as well as from a neontological perspective &#40;<a class="elsevierStyleCrossRef" href="#bib0230">Rzedowski&#44; 1978</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0060">Contreras-Medina and Eliosa-Le&#243;n&#44; 2001</a>&#59; <a class="elsevierStyleCrossRef" href="#bib0205">Morrone&#44; 2010</a>&#41;&#46; The generalized tracks identified in our analysis support the idea of a combined origin of the Mexican biota relating Laurasian and Gondwanan components in the central portion of the country&#46; Mexico&#39;s topographic and geographical characteristics have promoted a great variety of ecosystems and one of the highest levels of biodiversity worldwide &#40;<a class="elsevierStyleCrossRef" href="#bib0120">Flores-Villela and G&#233;rez&#44; 1989</a>&#41;&#46; In the central portion of the country the ecological and geological diversity is shown along with evidence of the integration of different organisms&#44; mainly mammals&#44; from the Nearctic region with immigrants from South America&#46; Tracks obtained with the fossil biota of Santa Mar&#237;a Amajac resemble those discovered with extant biotas&#44; suggesting that distribution patterns have prevailed since late Pliocene &#40;Blancan&#41;&#44; because Mexico&#39;s geological configuration experienced minimal changes in the last 2<span class="elsevierStyleHsp" style=""></span>million years&#44; and those changes shown in the central portion of the country are due to climatic variations more than geological modifications &#40;<a class="elsevierStyleCrossRef" href="#bib0150">Kurt&#233;n and Anderson&#44; 1980</a>&#41;&#46;</p></span></span>"
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          "titulo" => "Introduction"
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          "titulo" => "Materials and methods"
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    "fechaRecibido" => "2014-02-07"
    "fechaAceptado" => "2014-06-05"
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        0 => array:4 [
          "clase" => "keyword"
          "titulo" => "Key words"
          "identificador" => "xpalclavsec501913"
          "palabras" => array:6 [
            0 => "paleolake"
            1 => "track analysis"
            2 => "node"
            3 => "birds"
            4 => "mammals"
            5 => "fossils"
          ]
        ]
      ]
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        0 => array:4 [
          "clase" => "keyword"
          "titulo" => "Palabras clave"
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          "palabras" => array:6 [
            0 => "paleolago"
            1 => "an&#225;lisis de trazos"
            2 => "nodo"
            3 => "aves"
            4 => "mam&#237;feros"
            5 => "f&#243;siles"
          ]
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        "titulo" => "Abstract"
        "resumen" => "<span id="abst0005" class="elsevierStyleSection elsevierViewall"><p id="spar0035" class="elsevierStyleSimplePara elsevierViewall">The Santa Mar&#237;a Amajac paleolake is located in the central portion of Hidalgo&#44; Mexico&#46; Twenty-seven fossil taxa of aquatic and terrestrial plants&#44; gastropods&#44; ostracods&#44; amphibians&#44; and mammals identified previously in the area of the paleolake were selected and their distribution in America during the Late Pliocene- Pleistocene was analyzed using the panbiogeographic method&#46; As a result of the overlap of 27 individual tracks&#44; 5 generalized tracks were obtained&#58; <span class="elsevierStyleItalic">I&#41;</span> Western North American&#44; <span class="elsevierStyleItalic">II&#41;</span> Central North American&#44; <span class="elsevierStyleItalic">III&#41;</span> Eastern North American&#44; <span class="elsevierStyleItalic">IV&#41;</span> Mesoamerican&#44; and <span class="elsevierStyleItalic">V&#41;</span> South American&#46; The generalized tracks are consistent with previous proposals for extant gymnosperms&#44; amphibians&#44; sauropsids&#44; birds&#44; mammals&#44; aquatic plants&#44; insects&#44; beetles&#44; and nematodes&#44; suggesting that distribution patterns have prevailed since the late Pliocene &#40;Blancan&#41;&#46; Four biogeographic nodes were identified&#44; 2 situated in North America&#44; in California and Colorado&#44; the third located in the Santa Mar&#237;a Amajac area in central Mexico&#44; and the fourth located in Central America&#46;</p></span>"
      ]
      "es" => array:2 [
        "titulo" => "Resumen"
        "resumen" => "<span id="abst0010" class="elsevierStyleSection elsevierViewall"><p id="spar0040" class="elsevierStyleSimplePara elsevierViewall">El paleolago de Santa Mar&#237;a Amajac se ubica en la parte central del estado de Hidalgo&#44; M&#233;xico&#46; Se seleccionaron 27 taxones f&#243;siles pertenecientes a plantas acu&#225;ticas y terrestres&#44; gaster&#243;podos&#44; ostr&#225;codos&#44; anfibios y mam&#237;feros que hab&#237;an sido previamente identificados en el &#225;rea del paleolago&#44; y se analiz&#243; su distribuci&#243;n en Am&#233;rica durante el Plioceno Tard&#237;o-Pleistoceno a trav&#233;s del m&#233;todo panbiogeogr&#225;fico&#46; Como resultado de la superposici&#243;n de los 27 trazos individuales se obtuvieron 5 trazos generalizados&#58; <span class="elsevierStyleItalic">I&#41;</span> norteamericano occidental&#44; <span class="elsevierStyleItalic">II&#41;</span> norteamericano central&#44; <span class="elsevierStyleItalic">III&#41;</span> norteamerciano oriental&#44; <span class="elsevierStyleItalic">IV&#41;</span> mesoamericano y <span class="elsevierStyleItalic">V&#41;</span> sudamericano&#46; Los trazos generalizados coinciden con los propuestos para gimnospermas&#44; anfibios&#44; saur&#243;psidos&#44; aves&#44; mam&#237;feros&#44; plantas acu&#225;ticas&#44; insectos&#44; cole&#243;pteros y nem&#225;todos recientes&#59; lo que indica que los patrones de distribuci&#243;n han prevalecido desde el Plioceno tard&#237;o &#40;Blancano&#41;&#46; Se identificaron 4 nodos&#59; 2 localizados en Estados Unidos de Norteam&#233;rica&#44; en California y Colorado&#44; otro localizado en el &#225;rea de Santa Mar&#237;a Amajac&#44; en la parte central del pa&#237;s y el &#250;ltimo en Centroam&#233;rica&#46;</p></span>"
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          "en" => "<p id="spar0005" class="elsevierStyleSimplePara elsevierViewall">Individual tracks&#46; 1&#44; <span class="elsevierStyleItalic">Amelanchier</span> sp&#46;&#59; 2&#44; <span class="elsevierStyleItalic">Cercocarpus</span> sp&#46;&#59; 3&#44; <span class="elsevierStyleItalic">Crataegus</span> sp&#46;&#59; 4&#44; <span class="elsevierStyleItalic">Platanus</span> sp&#46;&#59; 5&#44; <span class="elsevierStyleItalic">Prunus</span> sp&#46;&#59; 6&#44; <span class="elsevierStyleItalic">Rhus</span> sp&#46;&#59; 7&#44; <span class="elsevierStyleItalic">Salix</span> sp&#46;&#59; 8&#44; <span class="elsevierStyleItalic">Populus</span> sp&#46;&#59; 9&#44; <span class="elsevierStyleItalic">Candona</span> sp&#46;</p>"
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        "descripcion" => array:1 [
          "en" => "<p id="spar0010" class="elsevierStyleSimplePara elsevierViewall">Individual tracks&#46; 10&#44; <span class="elsevierStyleItalic">Hyla</span> sp&#46;&#59; 11&#44; <span class="elsevierStyleItalic">Tremarctos</span> sp&#46;&#59; 12&#44; <span class="elsevierStyleItalic">Limnocythere</span> sp&#46;&#59; 13&#44; <span class="elsevierStyleItalic">Ambystoma</span> sp&#46;&#59; 14&#44; <span class="elsevierStyleItalic">Juniperus</span> sp&#46;&#59; 15&#44; <span class="elsevierStyleItalic">Typha</span> sp&#46;&#59; 16&#44; <span class="elsevierStyleItalic">Planorbis</span> sp&#46;&#44; 17&#44; <span class="elsevierStyleItalic">Darwinula</span> sp&#46;&#59; 18&#44; <span class="elsevierStyleItalic">Pinus</span> sp&#46;</p>"
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          "en" => "<p id="spar0015" class="elsevierStyleSimplePara elsevierViewall">19&#44; <span class="elsevierStyleItalic">Quercus</span> sp&#46;&#59; 20&#44; <span class="elsevierStyleItalic">Paramylodon</span> sp&#46;&#59; 21&#44; <span class="elsevierStyleItalic">Mammuthus</span> sp&#46;&#59; 22&#44; <span class="elsevierStyleItalic">Pampatherium</span> sp&#46;&#59; 23&#44; <span class="elsevierStyleItalic">Cuvieronius</span> sp&#46;&#59; 24&#44; <span class="elsevierStyleItalic">Sylvilagus</span> sp&#46;&#59; 25&#44; <span class="elsevierStyleItalic">Equus</span> sp&#46;&#59; 26&#44; <span class="elsevierStyleItalic">Platygonus</span> sp&#46;&#59;27&#44; <span class="elsevierStyleItalic">Rhynchotherium</span> sp&#46;</p>"
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          "en" => "<p id="spar0020" class="elsevierStyleSimplePara elsevierViewall">Generalized tracks and nodes obtained&#46; I&#44; Western North American&#59; II&#44; Central North American&#59; III&#44; Eastern North American&#59; IV&#44; Mesoamerican&#59; V&#44; South American&#59; a&#44; b&#44; c&#44; and d&#44; nodes located in California&#44; Colorado&#44; Santa Mar&#237;a Amajac&#44; and Guatemala&#44; respectively&#46;</p>"
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        "etiqueta" => "Table 1"
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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" title="table-head  " align="left" valign="bottom" scope="col" style="border-bottom: 2px solid black"><span class="elsevierStyleItalic">Taxa</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="center" valign="bottom" scope="col" style="border-bottom: 2px solid black"><span class="elsevierStyleItalic">Genera</span>&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" title="table-entry  " align="left" valign="bottom"><span class="elsevierStyleItalic">Terrestrial plants</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Amelanchier</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Cercocarpus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Crataegus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Juniperus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Pinus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Platanus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Populus</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Salix</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">Ostracods&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Darwinula</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Limnocythere</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">Amphibians&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="center" valign="bottom"><span class="elsevierStyleItalic">Ambystoma</span> sp&#46;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr><tr title="table-row"><td class="td" title="table-entry  " align="left" valign="bottom">&nbsp;\t\t\t\t\t\t\n
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              "identificador" => "bib0005"
              "etiqueta" => "Aguayo and Tr&#225;paga&#44; 1996"
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                          "autores" => array:2 [
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                    0 => array:1 [
                      "Libro" => array:5 [
                        "titulo" => "Geodin&#225;mica de M&#233;xico y minerales del mar"
                        "fecha" => "1996"
                        "paginaInicial" => "106"
                        "editorial" => "Fondo de Cultura Econ&#243;mica"
                        "editorialLocalizacion" => "M&#233;xico D&#46;F"
                      ]
                    ]
                  ]
                ]
              ]
            ]
            1 => array:3 [
              "identificador" => "bib0010"
              "etiqueta" => "Aguilar and Ortiz&#44; 2000"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:1 [
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:2 [
                            0 => "F&#46; Aguilar"
                            1 => "M&#46;E&#46; Ortiz"
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                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "Libro" => array:5 [
                        "titulo" => "Estudio paleoecol&#243;gico de la flora plioc&#233;nica de Santa Mar&#237;a Amajac&#44; Hidalgo&#58; Inferencia del paleoclima y de la paleocomunidad"
                        "fecha" => "2000"
                        "paginaInicial" => "79"
                        "editorial" => "Bachelor&#39;s thesis Facultad de Ciencias&#44; Universidad Nacional Aut&#243;noma de M&#233;xico"
                        "editorialLocalizacion" => "M&#233;xico&#44; D&#46; F"
                      ]
                    ]
                  ]
                ]
              ]
            ]
            2 => array:3 [
              "identificador" => "bib0015"
              "etiqueta" => "Aguilar et al&#46;&#44; 2002"
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                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:1 [
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:3 [
                            0 => "F&#46; Aguilar"
                            1 => "A&#46; Silva-Pineda"
                            2 => "M&#46;P&#46; Velasco-de Le&#243;n"
                          ]
                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "Libro" => array:4 [
                        "titulo" => "Registro de <span class="elsevierStyleItalic">Equisetum hyemale</span> en el Plioceno de la regi&#243;n de Santa Mar&#237;a Amajac&#44; Hidalgo&#44; M&#233;xico&#46; VII Congreso Latinoamericano y II Congreso Colombiano de Bot&#225;nica&#46; Cartagena de Indias&#44; Colombia"
                        "fecha" => "2002"
                        "paginaInicial" => "378"
                        "editorial" => "Asociaci&#243;n Latinoamericana de Bot&#225;nica"
                      ]
                    ]
                  ]
                ]
              ]
            ]
            3 => array:3 [
              "identificador" => "bib0020"
              "etiqueta" => "Aguilar-Aguilar and Contreras-Medina&#44; 2003"
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                0 => array:2 [
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                    0 => array:2 [
                      "titulo" => "La distribuci&#243;n de los mam&#237;feros marinos de M&#233;xico&#58; un enfoque panbiogeogr&#225;fico"
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                        "titulo" => "Introducci&#243;n a la biogeograf&#237;a en Latinoam&#233;rica&#58; teor&#237;as&#44; conceptos&#44; m&#233;todos y aplicaciones"
                        "paginaInicial" => "213"
                        "paginaFinal" => "219"
                        "serieFecha" => "2003"
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                ]
              ]
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            4 => array:3 [
              "identificador" => "bib0025"
              "etiqueta" => "Arroyo-Cabrales et al&#46;&#44; 2008"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "La diversidad en el pasado"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:4 [
                            0 => "J&#46; Arroyo-Cabrales"
                            1 => "A&#46;L&#46; Carre&#241;o"
                            2 => "S&#46; Lozano-Garc&#237;a"
                            3 => "M&#46; Montellano-Ballesteros"
                          ]
                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "LibroEditado" => array:4 [
                        "titulo" => "Capital natural de M&#233;xico&#44; vol&#46; I&#58; Conocimiento actual de la biodiversidad"
                        "paginaInicial" => "227"
                        "paginaFinal" => "262"
                        "serieFecha" => "2008"
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                  ]
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            5 => array:3 [
              "identificador" => "bib0030"
              "etiqueta" => "Asiain et al&#46;&#44; 2010"
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                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "Track analysis of the species of <span class="elsevierStyleItalic">Agrodes</span> and <span class="elsevierStyleItalic">Plochionocerus</span> &#40;Coleoptera&#58; Staphylinidae&#41;"
                      "autores" => array:1 [
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                          "autores" => array:3 [
                            0 => "J&#46; Asiain"
                            1 => "J&#46; M&#225;rquez"
                            2 => "J&#46;J&#46; Morrone"
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                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "Revista" => array:5 [
                        "tituloSerie" => "Revista Mexicana de Biodiversidad"
                        "fecha" => "2010"
                        "volumen" => "81"
                        "paginaInicial" => "177"
                        "paginaFinal" => "181"
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            6 => array:3 [
              "identificador" => "bib0035"
              "etiqueta" => "Candela and Morrone&#44; 2003"
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                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "Biogeograf&#237;a de puercoespines neotropicales &#40;Rodentia&#58; Hystricognathi&#41;&#58; Integrando datos f&#243;siles y actuales a trav&#233;s de un enfoque panbiogeogr&#225;fico"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:2 [
                            0 => "A&#46;M&#46; Candela"
                            1 => "J&#46;J&#46; Morrone"
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                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "Revista" => array:5 [
                        "tituloSerie" => "Ameghiniana"
                        "fecha" => "2003"
                        "volumen" => "40"
                        "paginaInicial" => "1"
                        "paginaFinal" => "19"
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                ]
              ]
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            7 => array:3 [
              "identificador" => "bib0040"
              "etiqueta" => "Carranza-Casta&#241;eda&#44; 1991"
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                        "titulo" => "Faunas de vertebrados f&#243;siles del Terciario tard&#237;o del &#225;rea de San Miguel de Allende&#44; Guanajuato"
                        "fecha" => "1991"
                        "paginaInicial" => "20"
                        "editorial" => "M&#233;xico&#44; D&#46; F&#46; III Congreso Nacional de Paleontolog&#237;a&#44; Sociedad Mexicana de Paleontolog&#237;a&#44; memorias"
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                    ]
                  ]
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              ]
            ]
            8 => array:3 [
              "identificador" => "bib0045"
              "etiqueta" => "Carranza-Casta&#241;eda&#44; 2006"
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                    0 => array:2 [
                      "titulo" => "Late Tertiary fossil localities in central M&#233;xico between 19&#176; 23&#176;N"
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                        0 => array:2 [
                          "etal" => false
                          "autores" => array:1 [
                            0 => "O&#46; Carranza-Casta&#241;eda"
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                  ]
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                    0 => array:1 [
                      "LibroEditado" => array:4 [
                        "titulo" => "Advances in late Tertiary Vertebrate Paleontology in Mexico and the Great American biotic interchange"
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            9 => array:3 [
              "identificador" => "bib0050"
              "etiqueta" => "Carranza-Casta&#241;eda and Ferrusqu&#237;a-Villafranca&#44; 1978"
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                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "Nuevas investigaciones sobre la fauna Rancho El Ocote&#44; Plioceno Medio de Guanajuato&#44; M&#233;xico&#59; informe preliminar Universidad Nacional Aut&#243;noma de M&#233;xico"
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                          "autores" => array:2 [
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                    0 => array:1 [
                      "Revista" => array:5 [
                        "tituloSerie" => "Instituto de Geolog&#237;a"
                        "fecha" => "1978"
                        "volumen" => "2"
                        "paginaInicial" => "163"
                        "paginaFinal" => "166"
                      ]
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                  ]
                ]
              ]
            ]
            10 => array:3 [
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