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Ecology
Dimorphism and population size of the Mexican redrump tarantula, Brachypelma vagans (Araneae: Theraphosidae), in Southeast Mexico
Dimorfismos y tamaño de poblaciones de la tarántula de cadera roja Brachypelma vagans (Araneae: Theraphosidae), en el sureste de México
Yann Hénauta,
Corresponding author
yhenaut@ecosur.mx

Corresponding author.
, Salima Machkour-M’Rabetb, Holger Weissenbergerc, Roberto Rojoa
a Laboratorio de Conducta Animal, El Colegio de la Frontera Sur., Av. Centenario Km. 5.5, 77014 Chetumal, Quintana Roo, Mexico
b Laboratorio de Ecología Molecular y Conservación, El Colegio de la Frontera Sur., Av. Centenario Km. 5.5, 77014 Chetumal, Quintana Roo, Mexico
c Laboratorio de Análisis de Información Geográfica y Estadística, Av. Centenario Km. 5.5, 77014 Chetumal, Quintana Roo, Mexico
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primarily to protect them from the illegal pet trade in the black market&#44; as these highly sought species are very docile&#44; large&#44; and beautifully colored &#40;<a class="elsevierStyleCrossRef" href="#bib0235">West&#44; 2005</a>&#41;&#46; Species of <span class="elsevierStyleItalic">Brachypelma</span> face other serious problems such as habitat destruction&#44; high juvenile mortality&#44; and late sexual maturity &#40;<a class="elsevierStyleCrossRef" href="#bib0145">Machkour-M&#8217;Rabet et al&#46;&#44; 2007</a>&#41;&#44; leading to population decline or local extinction&#46; Due to the vulnerability of the entire genus&#44; some studies have concentrated on understanding their ecology &#40;<a class="elsevierStyleCrossRefs" href="#bib0040">Criscuolo&#44; Font-Sala&#44; Bouillaud&#44; Poulin&#44; &#38; Trabalon&#44; 2010&#59; Dor&#44; Machkour-M&#8217;Rabet&#44; Legal&#44; Williams&#44; &#38; H&#233;naut&#44; 2008&#59; Machkour-M&#8217;Rabet&#44; H&#233;naut&#44; Rojo&#44; &#38; Calm&#233;&#44; 2005&#59; Machkour-M&#8217;Rabet et al&#46;&#44; 2007&#59; Vilchis-Nestor&#44; Machkour-M&#8217;Rabet&#44; Barriga-Sosa&#44; Winterton&#44; &#38; H&#233;naut&#44; 2013&#59; Y&#225;&#241;ez &#38; Floater&#44; 2000</a>&#41;&#59; behavior &#40;<a class="elsevierStyleCrossRefs" href="#bib0065">Dor&#44; Calm&#233;&#44; &#38; H&#233;naut&#44; 2011&#59; Dor &#38; H&#233;naut&#44; 2011&#44; 2012&#44; 2013&#59; Locht&#44; Y&#225;&#241;ez&#44; &#38; V&#225;zquez&#44; 1999&#59; Reichling&#44; 2000&#59; Y&#225;&#241;ez&#44; Locht&#44; &#38; Mac&#237;as-Ord&#243;&#241;ez&#44; 1999</a>&#41;&#59; genetic structure &#40;<a class="elsevierStyleCrossRefs" href="#bib0125">Longhorn&#44; Nicholas&#44; Chuter&#44; &#38; Volger&#44; 2007&#59; Machkour-M&#8217;Rabet&#44; H&#233;naut&#44; Calm&#233;&#44; &#38; Legal&#44; 2012&#59; Machkour-M&#8217;Rabet et al&#46;&#44; 2009</a>&#41;&#44; and traditional use by local populations &#40;<a class="elsevierStyleCrossRef" href="#bib0150">Machkour-M&#8217;Rabet&#44; H&#233;naut&#44; Winterton&#44; &#38; Rojo&#44; 2011</a>&#41;&#46; The <span class="elsevierStyleItalic">Brachypelma</span> species are restricted to small and specific areas&#44; with one exception&#44; the Mexican redrump tarantula <span class="elsevierStyleItalic">B&#46; vagans</span> Ausserer&#44; 1875&#46; This is the only species of the genus that is widely distributed and has been subject to several studies during recent years &#40;<a class="elsevierStyleCrossRefs" href="#bib0070">Dor et al&#46;&#44; 2008&#44; 2011&#59; Machkour-M&#8217;Rabet et al&#46;&#44; 2005&#44; 2007&#44; 2009&#44; 2011&#44; 2012</a>&#41;&#44; also <span class="elsevierStyleItalic">B&#46; vagans</span> was identified in Florida &#40;<a class="elsevierStyleCrossRef" href="#bib0075">Edwards &#38; Hibbard&#44; 1999</a>&#41; and Cozumel Island &#40;<a class="elsevierStyleCrossRef" href="#bib0130">Machkour-M&#8217;Rabet et al&#46;&#44; 2012</a>&#41; as an invasive exotic tarantula&#46; These studies demonstrate that this tarantula may be present in high densities in rural villages with low levels of anthropogenic disturbance and closed to medium semi-evergreen forests &#40;<a class="elsevierStyleCrossRef" href="#bib0140">Machkour-M&#8217;Rabet et al&#46;&#44; 2005</a>&#41;&#46; In these areas&#44; soil structure appears to be an important factor explaining the presence and high density of <span class="elsevierStyleItalic">B&#46; vagans</span> due to their burrower condition &#40;<a class="elsevierStyleCrossRef" href="#bib0145">Machkour-M&#8217;Rabet et al&#46;&#44; 2007</a>&#41;&#46; In this context&#44; <a class="elsevierStyleCrossRef" href="#bib0080">H&#233;naut and Machkour-M&#8217;Rabet &#40;2005&#41;</a> observed that coexisting females are very aggressive toward congeners and commonly attack other females&#44; which are detected by chemical cues &#40;<a class="elsevierStyleCrossRef" href="#bib0070">Dor et al&#46;&#44; 2008</a>&#41;&#46;</p><p id="par0010" class="elsevierStylePara elsevierViewall">Many species of animals with wide distribution ranges exhibit geographical variations in growth and life history traits &#40;<a class="elsevierStyleCrossRef" href="#bib0210">Stillwell &#38; Fox&#44; 2009</a>&#41;&#46; The body size of many animals varies with latitude and altitude &#40;<a class="elsevierStyleCrossRefs" href="#bib0020">Blanckenhorn &#38; Demont&#44; 2004&#59; Stillwell&#44; Morse&#44; &#38; Fox&#44; 2007</a>&#41;&#44; and the most common environmental variable advocated to explain body size variation is temperature &#40;thermocline&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib0210">Stillwell &#38; Fox&#44; 2009</a>&#41;&#46; The geographic adaptations of animals&#44; which include variations in body size&#44; are generally genetically based &#40;<a class="elsevierStyleCrossRefs" href="#bib0010">Armbruster&#44; Bradshaw&#44; Ruegg&#44; &#38; Holzapfel&#44; 2001&#59; Karl&#44; Janowitz&#44; &#38; Fisher&#44; 2008</a>&#41;&#46; Studies show that a thermocline &#40;or other factors which vary with latitude or altitude&#41; can result in both intra-specific and inter-specific size dimorphism &#40;<a class="elsevierStyleCrossRefs" href="#bib0025">Blanckenhorn&#44; Stillwell&#44; Young&#44; Fox&#44; &#38; Ashton&#44; 2006&#59; Stillwell &#38; Fox&#44; 2007&#59; Teder &#38; Tammaru&#44; 2005</a>&#41;&#46; The body size of animals may also be correlated with population density &#40;<a class="elsevierStyleCrossRef" href="#bib0180">Robinson &#38; Redford&#44; 1986</a>&#41;&#46; Studies on <span class="elsevierStyleItalic">Nephila clavipes</span> Leach&#44; 1815 &#40;Araneae&#44; Nephilidae&#41; &#40;<a class="elsevierStyleCrossRefs" href="#bib0085">Higgins&#44; 1992&#44; 1993&#44; 1995</a>&#41; demonstrated that environmental variations result in differences in the size of individuals between spider populations&#46; <a class="elsevierStyleCrossRef" href="#bib0100">Higgins &#40;2000&#41;</a> showed that seasonal variations or prey availability result in dissimilarities in spider size&#59; females strategically adapt to these fluctuations by reaching maturity earlier when ecological conditions are degraded or reproduce before the end of the favorable season&#46; Furthermore&#44; <a class="elsevierStyleCrossRef" href="#bib0100">Higgins &#40;2000&#41;</a> demonstrated that late maturing <span class="elsevierStyleItalic">N&#46; clavipes</span> females present lower reproductive success in strongly seasonal habitats&#46; In <span class="elsevierStyleItalic">B&#46; vagans</span>&#44; a recent study &#40;<a class="elsevierStyleCrossRef" href="#bib0225">Vilchis-Nestor et al&#46;&#44; 2013</a>&#41; showed that individuals recently introduced to an island &#40;Cozumel Island&#44; Quintana Roo&#44; Mexico&#41;&#44; had larger adults and a lower diversity of body patterns than individuals from mainland populations &#40;e&#46;g&#46;&#44; the Yucat&#225;n Peninsula&#44; Mexico&#41;&#46;</p><p id="par0015" class="elsevierStylePara elsevierViewall">The principal type of dimorphism in spiders is sexual dimorphism&#44; with males usually smaller than females &#40;<a class="elsevierStyleCrossRef" href="#bib0110">Hormiga&#44; Scharff&#44; &#38; Coddington&#44; 2000</a>&#41;&#46; According to <a class="elsevierStyleCrossRef" href="#bib0045">Darwin &#40;1871&#41;</a> and present day authors&#44; the first example of sexual dimorphism in animals was observed in spiders&#44; principally web-building species from various families&#44; but also for non-web-building groups such as the Lycosidae <span class="elsevierStyleItalic">Rabidos rabida</span> &#40;<a class="elsevierStyleCrossRef" href="#bib0230">Walker &#38; Rypstra&#44; 2001</a>&#41;&#46; Although sexual dimorphism is extreme in web-building spiders&#44; non-web-building spiders generally have a lower degree of dimorphism &#40;<a class="elsevierStyleCrossRef" href="#bib0230">Walker &#38; Rypstra&#44; 2001</a>&#41;&#46; Sexual dimorphism is also evident in the relative size of body parts&#44; with males having comparatively longer legs than females &#40;<a class="elsevierStyleCrossRef" href="#bib0170">Prenter&#44; Montgomery&#44; &#38; Elwood&#44; 1995</a>&#41;&#46; In Theraphosidae&#44; males and females appear to be similar sized&#44; but they present sexual dimorphism with respect to metabolic rates &#40;<a class="elsevierStyleCrossRef" href="#bib0190">Shillington&#44; 2005</a>&#41;&#46; Tarantula females are sit-and-wait predators that remain in the same location during large periods of time&#46; In contrast&#44; males disperse by walking and actively search for females over large distances &#40;<a class="elsevierStyleCrossRefs" href="#bib0130">Machkour-M&#8217;Rabet et al&#46;&#44; 2012&#59; Shillington&#44; 2002</a>&#41;&#46;</p><p id="par0020" class="elsevierStylePara elsevierViewall">This study compares populations of <span class="elsevierStyleItalic">B&#46; vagans</span> from different geographical locations in Southeast Mexico&#46; First&#44; the study aims to determine if tarantula population size varies according to different geographical locations&#46; Second&#44; we focus on dimorphism among females from different populations to determine if geographical dimorphism occurs in <span class="elsevierStyleItalic">B&#46; vagans</span>&#46;</p></span><span id="sec0010" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0030">Materials and methods</span><p id="par0025" class="elsevierStylePara elsevierViewall">All data for the morphology of <span class="elsevierStyleItalic">B&#46; vagans</span> were obtained from a total of 6 sites in four different states of southern Mexico &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#44; <a class="elsevierStyleCrossRef" href="#fig0005">Fig&#46; 1</a>&#41;&#46; In Chiapas&#44; we sampled 2 sites&#44; El Casta&#241;o &#40;CH-EC&#59; 15&#176;17&#8242;N&#44; 92&#176;58&#8242;W&#44; 16<span class="elsevierStyleHsp" style=""></span>masl&#41; near the Pacific coast&#44; and in Frontera Corozal &#40;CH-FC&#59; 16&#176;49&#8242;N&#44; 90&#176;53&#8242;W&#44; 117<span class="elsevierStyleHsp" style=""></span>masl&#41; in the Lacandona rainforest&#46; In Campeche&#44; we visited 1 site &#40;11 de Mayo&#59; C-11M&#59; 18&#176;18&#8242;N&#44; 89&#176;27&#8242;W&#44; 281<span class="elsevierStyleHsp" style=""></span>masl&#41; close to the Calakmul Biosphere Reserve&#46; In Quintana Roo&#44; we visited 2 sites&#44; 1 on the mainland &#40;Coba&#59; QR-CB&#59; 20&#176;28&#8242;N&#44; 87&#176;44&#8242;W&#44; 13<span class="elsevierStyleHsp" style=""></span>masl&#41; and 1 on Cozumel Island &#40;Rancho Guadalupe&#59; QR-RG&#59; 20&#176;29&#8242;N&#44; 86&#176;50&#8242;W&#44; 17<span class="elsevierStyleHsp" style=""></span>masl&#41;&#46; We also recorded data from a site in Veracruz &#40;Paso de Milpa&#59; V-PM&#59; 19&#176;26&#8242;N&#44; 96&#176;36&#8242;W&#44; 260<span class="elsevierStyleHsp" style=""></span>masl&#41;&#46; Collections were made at the beginning of the rainy season for Chiapas and Yucat&#225;n Peninsula&#44; which is between May and July&#46; The start of the rains coincides with the reproductive season&#46; All the sites studied were rural communities with similar ecological characteristics&#58; deep clay soil with little vegetation&#44; roots&#44; and limestone rocks &#40;<a class="elsevierStyleCrossRefs" href="#bib0140">Machkour-M&#8217;Rabet et al&#46;&#44; 2005&#44; 2007</a>&#41;&#46; Climatic characteristics were very similar for all study sites &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#41;&#46;</p><elsevierMultimedia ident="tbl0005"></elsevierMultimedia><elsevierMultimedia ident="fig0005"></elsevierMultimedia><p id="par0030" class="elsevierStylePara elsevierViewall">Individual spiders were collected manually throughout the entire area&#46; Despite differences in study area size&#44; most tarantulas were found in their burrows&#44; which were concentrated in particular areas such as gardens or football fields&#44; as observed in previous studies &#40;<a class="elsevierStyleCrossRefs" href="#bib0140">Machkour-M&#8217;Rabet et al&#46;&#44; 2005&#44; 2007</a>&#41;&#46; In Veracruz and Campeche states&#44; we also collected males found walking close to the sampling area &#40;less than 1<span class="elsevierStyleHsp" style=""></span>km&#41;&#46; We also recorded morphological data for these males as they were considered characteristic of the corresponding localities&#46; Males and females were sexed by the tibial apophysis of leg I and the palp embolus in males&#44; and by the females spermathecae&#46; Collected individuals were measured and classified as adults &#40;males and females&#41; or juveniles based on sexual characters&#46; The specimens were collected and measured in the field during the night between 20&#58;30 and 00&#58;30<span class="elsevierStyleHsp" style=""></span>h&#46; A small stick was used to remove the spiders from their burrows and handle them carefully by hand&#46; For each individual&#44; we measured the length and width of the prosoma &#40;Lpro and Wpro&#44; respectively&#41;&#44; the length of patella I &#40;PI&#41; and IV &#40;PIV&#41;&#44; and tibia I &#40;TI&#41; and IV &#40;TIV&#41; as described by <a class="elsevierStyleCrossRef" href="#bib0035">Chamberlin and Ivie &#40;1938&#41;</a>&#44; the same methodology used in previous studies &#40;<a class="elsevierStyleCrossRef" href="#bib0140">Machkour-M&#8217;Rabet et al&#46;&#44; 2005</a>&#41;&#46; Measurements were taken in millimeters using a digital vernier &#40;Truper&#41;&#46; The individuals were weighed &#40;<span class="elsevierStyleItalic">W</span>&#41; to the nearest 0&#46;1<span class="elsevierStyleHsp" style=""></span>g using a spring balance &#40;Pesola&#41;&#44; the measurements of the weight were in grams &#40;g&#41;&#46; Once all the data had been recorded&#44; each spider was released in front of the entrance to its burrow&#46; Specimens from previous research in the same areas are deposited in the Museum of Zoology&#44; El Colegio de la Frontera Sur &#40;ECOSUR&#41;&#44; Chetumal&#44; Quintana Roo&#44; Mexico&#46;</p><p id="par0035" class="elsevierStylePara elsevierViewall">The <span class="elsevierStyleItalic">G</span>-test was used to compare the number of males&#44; females&#44; and juveniles collected among sites&#46; Morphological measurements for males and females recorded at the sites were compared using the nonparametric Mann&#8211;Whitney <span class="elsevierStyleItalic">U</span>-test&#46;</p><p id="par0040" class="elsevierStylePara elsevierViewall">In Veracruz and Campeche state&#44; walking males that were found close to but outside the sampling area were considered for means of analysis&#44; representative of males from within the study area&#46; To compare morphological measurements of females between sites&#44; a nonparametric Kruskal&#8211;Wallis test was carried out&#44; followed by a post hoc comparison of mean rank&#46; All statistical analyses were performed using <span class="elsevierStyleSmallCaps">S</span>tatistica v7&#46;0&#46;</p></span><span id="sec0015" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0035">Results</span><p id="par0045" class="elsevierStylePara elsevierViewall">We collected a total of 126 <span class="elsevierStyleItalic">B&#46; vagans</span> individuals from the 6 sites&#58; 41 juveniles and 85 adults&#46; The number of females &#40;<span class="elsevierStyleItalic">n</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>77&#41; was significantly higher &#40;<span class="elsevierStyleItalic">G</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>64&#44; <span class="elsevierStyleItalic">d&#46;f&#46;</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>1&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;000&#41; than males &#40;<span class="elsevierStyleItalic">n</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>8&#41;&#44; giving an overall sex ratio of approximately 10 females per male&#46; In Campeche and Veracruz we found 6 and 11 males&#44; respectively&#44; outside the sampling area on the village perimeter&#46;</p><p id="par0050" class="elsevierStylePara elsevierViewall">The number of individuals observed presents a high degree of variation according to location&#46; In Veracruz&#44; 10 females were observed at the Paso de Milpa site&#46; In Chiapas&#44; 6 females and 1 male were observed in El Casta&#241;o and 3 females and 1 juvenile in Frontera Corozal&#46; In Quintana Roo&#44; 11 individuals &#40;3 females&#44; 1 male&#44; and 7 juveniles&#41; were observed in Cob&#225; and 26 in Cozumel &#40;9 females and 17 juveniles&#41;&#46; Sixty-seven tarantulas were observed at the 11 de Mayo site in the state of Campeche &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Fig&#46; 1</a>&#41;&#46; The number of collected individuals at each site varied considerably &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Fig&#46; 1</a>&#41;&#44; varying between 4 and 11 individuals at the sites in Chiapas&#44; Veracruz&#44; and mainland Quintana Roo&#46; At Rancho Guadalupe &#40;Cozumel Island&#41; we observed 23 individuals&#44; while the 11 de Mayo site&#44; with 67 individuals&#44; accounted for the largest proportion &#40;53&#37;&#41; of the total number of spiders collected at the 6 sites&#46;</p><p id="par0055" class="elsevierStylePara elsevierViewall">Males and females were compared in populations with a sufficient number of individuals of each sex&#44; fixed at a minimum of 6&#46; Only the sites in Campeche and Veracruz met this minimum number&#46; Male and female spiders presented significant differences in size in Campeche &#40;<a class="elsevierStyleCrossRef" href="#fig0010">Fig&#46; 2</a>&#41;&#46; Males were significantly larger than females with longer legs &#40;tibias from the I and IV legs&#44; and patella from the IV leg&#41; and prosoma&#46; Female weight &#40;mean<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>SE&#58; 7&#46;7<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;6<span class="elsevierStyleHsp" style=""></span>g&#41; was also significantly lower &#40;Mann&#8211;Whitney <span class="elsevierStyleItalic">U</span>-test&#58; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>132&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;008&#41; than male weight &#40;mean<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>SE&#58; 11&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1<span class="elsevierStyleHsp" style=""></span>g&#41;&#46; This was not the case in Veracruz&#44; especially with respect to the females and males found in Paso de Milpa&#46; At this site&#44; males were significantly larger than females in two measurements&#58; TI male&#58; 13&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3<span class="elsevierStyleHsp" style=""></span>mm&#44; TI female&#58; 10&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm &#40;Mann&#8211;Whitney <span class="elsevierStyleItalic">U</span>-test&#58; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>4&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;000&#41;&#44; and TIV male&#58; 13&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#44; TIV female&#58; 11&#46;4<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3<span class="elsevierStyleHsp" style=""></span>mm &#40;Mann&#8211;Whitney <span class="elsevierStyleItalic">U</span>-test&#58; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>17&#46;5&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;008&#41;&#46; Males were significantly lighter than females &#40;males&#58; 8&#46;9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>g vs&#46; females&#58; 21&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1&#46;5<span class="elsevierStyleHsp" style=""></span>g&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;00&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;000&#41;&#46; In this area no differences between sexes were observed for other morphological characters&#58; PI &#40;males&#58; 9&#46;5<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1<span class="elsevierStyleHsp" style=""></span>mm vs&#46; females&#58; 9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>55&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;4&#41;&#44; PIV &#40;males&#58; 8&#46;6<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1<span class="elsevierStyleHsp" style=""></span>mm vs&#46; females&#58; 8&#46;6<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;2<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>50&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;3&#41;&#44; prosoma length &#40;males&#58; 19&#46;7<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm vs&#46; females&#58; 20&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;6<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>52&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;4&#41;&#44; and prosoma width &#40;males&#58; 18<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm vs&#46; females&#58; 19&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;6<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>55&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;5&#41;&#46;</p><elsevierMultimedia ident="fig0010"></elsevierMultimedia><p id="par0060" class="elsevierStylePara elsevierViewall">Comparing males found in different geographical areas&#44; no significant differences were observed between Campeche and Veracruz&#58; PI &#40;Campeche 8&#46;12<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 9&#46;5<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>55&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;4&#41;&#44; TI &#40;Campeche 13&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 13&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>46&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;2&#41;&#44; PIV &#40;Campeche 8&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 8&#46;6<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>50&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;3&#41;&#44; TIV &#40;Campeche 13&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 13&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>44&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;17&#41;&#44; prosoma length &#40;Campeche 18&#46;7<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;7<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 19&#46;7<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>52&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;4&#41;&#44; prosoma width &#40;Campeche 17&#46;5<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;7<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 18<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>55&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;5&#41;&#44; and weight &#40;Campeche 11&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1<span class="elsevierStyleHsp" style=""></span>mm&#59; Veracruz&#58; 8&#46;9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;4<span class="elsevierStyleHsp" style=""></span>mm&#59; <span class="elsevierStyleItalic">U</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>21&#44; <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p><p id="par0065" class="elsevierStylePara elsevierViewall">Due to the low number of female individuals observed at most sites&#44; we could only compare females from 4 of them&#58; Paso de Milpa in Veracruz with 10 females&#44; El Casta&#241;o in Chiapas with 6 females&#44; Rancho Guadalupe in Quintana Roo with 9 females&#44; and 11 de Mayo Campeche with 45 females&#46; Despite the low number of individuals&#44; differences in size between females from the area with a high number of individuals &#40;11 de Mayo&#41; and areas with few individuals &#40;the remaining localities&#41;&#44; demonstrate a significant difference between 11 de Mayo and the other sites&#46; Leg and prosoma size is significantly lower in female tarantulas from Campeche when compared to the other 3 sites&#46; Weight is higher in tarantulas from Chiapas&#44; Cozumel &#40;Rancho Guadalupe&#41;&#44; and Veracruz&#46; Despite their geographical distance&#44; the morphology of tarantulas from populations with low number of individuals appears to be similar &#40;<a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>&#41;&#46;</p><elsevierMultimedia ident="tbl0010"></elsevierMultimedia></span><span id="sec0020" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0040">Discussion</span><p id="par0070" class="elsevierStylePara elsevierViewall">Some studies of tarantulas show different aspects of sexual dimorphism&#46; For example&#44; <a class="elsevierStyleCrossRefs" href="#bib0185">Shillington &#40;2002&#44; 2005&#41;</a> reported the significance of life history sexual dimorphism for <span class="elsevierStyleItalic">Aphonopelma anax</span> Chamberlin&#44; 1940 &#40;Araneae&#58; Theraphosidae&#41;&#46; Mature male tarantulas lead a very active life that revolves around searching for females for reproduction&#46; This implies higher energy demands&#44; leading to a higher resting metabolic rate &#40;RMR&#41; than females&#46; <a class="elsevierStyleCrossRef" href="#bib0165">P&#233;rez-Miles &#40;2002&#41;</a> described sexual dimorphism related to abdominal urticating hairs for different species of tarantula of the Theraphosinae subfamily&#59; however&#44; few formal studies have reported sexual size dimorphism in tarantulas&#46; Shillington observed that males of <span class="elsevierStyleItalic">A&#46; anax</span> have a smaller abdomen and longer legs than females &#40;<a class="elsevierStyleCrossRef" href="#bib0195">Shillington &#38; Peterson&#44; 2002</a>&#41;&#46; <a class="elsevierStyleCrossRef" href="#bib0160">P&#233;rez-Miles &#40;1989&#41;</a> observed variation in somatic characters between males and females&#44; particularly for legs and prosoma&#46; This study presents&#44; for the first time&#44; clear evidence of sexual size dimorphism in tibia length for <span class="elsevierStyleItalic">Brachypelma</span>&#46; Males of <span class="elsevierStyleItalic">B&#46; vagans</span> have a significantly longer tibia than females&#44; probably a result of selective pressure related to the reproductive role of wandering males&#46; For many species of tarantula&#44; sexually mature males abandon their burrow and actively search for females to mate with over a large geographical area &#40;<a class="elsevierStyleCrossRef" href="#bib0185">Shillington&#44; 2002</a>&#41;&#44; which makes it more difficult to find males than females&#46; This capacity of males increases their reproductive success and gene dispersal&#46; A recent genetic study of <span class="elsevierStyleItalic">B&#46; vagans</span> &#40;<a class="elsevierStyleCrossRef" href="#bib0130">Machkour-M&#8217;Rabet et al&#46;&#44; 2012</a>&#41; reported that the males of this species have a very high potential for dispersal&#46; As the males cover long distances while seeking a mate &#40;<a class="elsevierStyleCrossRefs" href="#bib0185">Shillington&#44; 2002&#59; Machkour-M&#8217;Rabet et al&#46;&#44; 2012</a>&#41;&#44; this could have led to the phenotypic adaptation of a longer tibia&#46; Another explanation could be that during reproduction the male hooks the female fangs to elevate the female body and insert the palpal organ into the female genitalia &#40;<a class="elsevierStyleCrossRef" href="#bib0200">Shillington &#38; Verrell&#44; 1997</a>&#41;&#59; consequently&#44; access to female genitalia could be favored by longer legs in males&#44; particularly pair I&#44; to facilitate the reproduction process&#46; Therefore&#44; the evolutionary process may favor longer legs in males&#46; Male size may also be an adaptation for protection against females that may not predate organisms of an equivalent or larger size&#44; given the risk of injuries&#46;</p><p id="par0075" class="elsevierStylePara elsevierViewall">In non-web building spiders&#44; males had longer legs than females &#40;<a class="elsevierStyleCrossRef" href="#bib0230">Walker &#38; Rypstra&#44; 2001</a>&#41;&#44; but not bigger prosoma&#46; At the Campeche site females are numerous and smaller than in other populations&#46; In addition&#44; they are smaller than males found within the same population&#46; Few studies have reported cases of geographical variation in sexual dimorphism in arthropods &#40;<a class="elsevierStyleCrossRefs" href="#bib0015">Bidau &#38; Mart&#237;&#44; 2007&#59; Blanckenhorn &#38; Demont&#44; 2004&#59; Stillwell &#38; Fox&#44; 2009</a>&#41;&#46; For spiders&#44; <a class="elsevierStyleCrossRef" href="#bib0155">Pek&#225;r and Va&#241;hara &#40;2006&#41;</a> described geographical sexual size dimorphism in <span class="elsevierStyleItalic">Zodarion rubidum</span> Simon&#44; 1914 &#40;Araneae&#58; Zodariidae&#41;&#44; an ant-eating spider&#46; Regions with higher temperatures provide optimal conditions for higher prey availability resulting in larger females &#40;<a class="elsevierStyleCrossRef" href="#bib0155">Pek&#225;r &#38; Va&#241;hara&#44; 2006</a>&#41;&#46; In addition to variations in resource availability&#44; the quality of prey can also influence intraspecific variation in body size &#40;<a class="elsevierStyleCrossRef" href="#bib0005">Amarello et al&#46;&#44; 2010</a>&#41;&#46; However&#44; in Campeche where females are smaller&#44; male size does not appear to be affected&#46; Data were collected at sites with human presence that are frequently inhabited by <span class="elsevierStyleItalic">B&#46; vagans</span> &#40;<a class="elsevierStyleCrossRef" href="#bib0140">Machkour-M&#8217;Rabet et al&#46;&#44; 2005</a>&#41;&#59; therefore&#44; there is little pressure from natural predators such as the mammal &#8220;coat&#237;&#8221;&#44; <span class="elsevierStyleItalic">Nasua narica</span> L&#46;&#44; 1766 &#40;Carnivora&#58; Procyonidae&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib0105">Hirsch&#44; 2009</a>&#41;&#44; since wild mammals do not normally venture into human settlements&#46; The Pepsi wasps &#40;<span class="elsevierStyleItalic">Pepsis</span> spp&#46;&#59; Hymenoptera&#44; Pompilidae&#41;&#44; also known as tarantula-hawks&#44; as described by <a class="elsevierStyleCrossRef" href="#bib0030">Cazier and Mortenson &#40;1964&#41;</a>&#44; may also predate on <span class="elsevierStyleItalic">B&#46; vagans</span>&#46; However&#44; during our time in the communities of Campeche &#40;<a class="elsevierStyleCrossRefs" href="#bib0080">H&#233;naut &#38; Machkour-M&#8217;Rabet&#44; 2005&#59; Machkour-M&#8217;Rabet et al&#46;&#44; 2005&#44; 2007&#44; 2011</a>&#41;&#44; this species of wasp was rarely observed and does not appear to be an important predator of tarantula populations&#46; The number of individuals per population may explain the morphological differences between females from different populations and&#44; between males and females in the Campeche population&#46; <a class="elsevierStyleCrossRefs" href="#bib0140">Machkour-M&#8217;Rabet et al&#46; &#40;2005&#44; 2007&#41;</a> describe the relationship between traditional human activity&#44; soil characteristics&#44; and the presence of <span class="elsevierStyleItalic">B&#46; vagans</span> in villages&#46; Using the characteristics from these initial studies&#44; it was easy to find <span class="elsevierStyleItalic">B&#46; vagans</span> individuals at other sites&#44; even though they were geographically distant&#46; However&#44; the number of tarantulas varied considerably with more individuals found at the Campeche site&#46; Although&#44; the causes are still unknown&#44; the high number of tarantulas in 11 de Mayo site may indicate that ecological factors are particularly favorable for these spiders in this particular area&#44; although female tarantulas were smaller than those found in other geographical areas&#46; The latter may be associated with a high degree of intraspecific competition&#44; with large numbers of individuals competing for the same resources&#46; Moreover&#44; cannibalism behavior was commonly observed between females at this site &#40;<a class="elsevierStyleCrossRef" href="#bib0080">H&#233;naut &#38; Machkour-M&#8217;Rabet&#44; 2005</a>&#41;&#46;</p><p id="par0080" class="elsevierStylePara elsevierViewall">The number of spiders collected is a good indicator of the abundance of tarantulas at each locality&#46; The high number of tarantulas observed in Campeche State and the low number of <span class="elsevierStyleItalic">B&#46; vagans</span> observed in others sites of southern Mexico has to be seriously considered when developing conservation strategies for this species&#46; As <span class="elsevierStyleItalic">B&#46; vagans</span> may rapidly go extinct in states where their numbers are so low&#44; there is a clear need for this species to receive adequate protection&#46; Furthermore&#44; large populations of this tarantula in Campeche need to be provided special protection as they represent important centers of dispersal and resilience for this species&#46; Moreover&#44; the population from Campeche&#44; close to the Calakmul Biosphere Reserve&#44; had more individuals than the remaining populations&#46; This may be a result of its proximity to a large protected area&#44; and further research may help us understand the importance of these areas for the conservation of the Mexican redrump Tarantula&#46;</p></span></span>"
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          "titulo" => "Introduction"
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          "titulo" => "Materials and methods"
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        6 => array:2 [
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          "titulo" => "Results"
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            0 => "<span class="elsevierStyleItalic">Brachypelma</span>"
            1 => "Tarantulas"
            2 => "Density"
            3 => "Morphometry"
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        "resumen" => "<span id="abst0005" class="elsevierStyleSection elsevierViewall"><p id="spar0005" class="elsevierStyleSimplePara elsevierViewall">As a general rule&#44; spiders exhibit sexual dimorphism and their populations may differ in size according to season duration and resource availability&#46; However&#44; few studies have focused on dimorphism in tarantulas&#46; Mexican redrump tarantulas&#44; <span class="elsevierStyleItalic">Brachypelma vagans</span>&#44; listed in CITES&#44; have an exceptionally wide distribution&#46; Surprisingly&#44; there are no studies on the possible relationship between the abundance of tarantulas per population and the geographical areas where they are present&#44; or on how the distribution pattern of this spider may affect individual morphological characteristics&#46; Furthermore&#44; there are no studies on sexual dimorphism within the genus <span class="elsevierStyleItalic">Brachypelma</span>&#46; The aim of the study is to determine the existence of sexual and geographical dimorphism in populations of <span class="elsevierStyleItalic">B&#46; vagans</span>&#46; It was observed that the abundance of spiders per population may vary according to the geographical areas where they were recorded&#46; In six localities in southern Mexico&#44; we recorded morphological data on adult tarantulas&#46; Sexual dimorphism was clearly observed at the site that presented numerous spiders characterized by much smaller females&#46; Since the results of this study demonstrate differences in tarantula number of individuals per locality in southern Mexico&#44; they make an important contribution to the conservation of this species&#46;</p></span>"
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        "resumen" => "<span id="abst0010" class="elsevierStyleSection elsevierViewall"><p id="spar0010" class="elsevierStyleSimplePara elsevierViewall">El dimorfismo sexual es muy com&#250;n en las ara&#241;as&#44; pero tambi&#233;n existen diferencias morfol&#243;gicas entre poblaciones en funci&#243;n de las temporadas y la disponibilidad en los recursos&#46; Pocos estudios han analizado el dimorfismo en tar&#225;ntulas&#46; La tar&#225;ntula mexicana de cadera roja <span class="elsevierStyleItalic">Brachypelma vagans</span>&#44; listada en el CITES&#44; presenta una distribuci&#243;n amplia&#46; Sin embargo&#44; se conoce poco sobre sus poblaciones por &#225;reas geogr&#225;ficas y sobre c&#243;mo el patr&#243;n de distribuci&#243;n de estos organismos&#44; puede afectar las caracter&#237;sticas morfol&#243;gicas individuales&#46; Tampoco se ha estudiado el dimorfismo sexual en el g&#233;nero <span class="elsevierStyleItalic">Brachypelma</span>&#46; Nuestro estudio se enfoca en determinar si existen dimorfismos sexual y geogr&#225;fico en poblaciones espec&#237;ficas de <span class="elsevierStyleItalic">B&#46; vagans</span>&#46; Se observ&#243; que la abundancia de tar&#225;ntulas encontradas por poblaci&#243;n suele ser diferente de acuerdo con las &#225;reas geogr&#225;ficas donde fueron observadas&#46; Registramos datos morfol&#243;gicos de tar&#225;ntulas adultas en 6 sitios geogr&#225;ficos distribuidos en el sureste de M&#233;xico&#46; Encontramos dimorfismo sexual &#250;nicamente en un sitio que presenta un n&#250;mero de ara&#241;as muy alto y donde las hembras son m&#225;s peque&#241;as que en otras localidades&#46; Nuestros resultados&#44; considerando la variaci&#243;n en n&#250;mero de individuos por localidades a lo largo del sureste mexicano&#44; tienen consecuencias para la conservaci&#243;n de esta especie&#46;</p></span>"
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          "en" => "<p id="spar0015" class="elsevierStyleSimplePara elsevierViewall">Map of sites where <span class="elsevierStyleItalic">Brachypelma vagans</span> was observed in Southeast Mexico&#46; F&#44; numbers of female individuals&#59; M&#44; number of male individuals&#59; J&#44; number of juvenile individuals&#46; For codes of sites see <a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#46; The number of wandering males found close to but not inside each sampling area is not considered in this figure&#46;</p>"
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          "en" => "<p id="spar0020" class="elsevierStyleSimplePara elsevierViewall">Mean &#40;&#177;standard error&#41; of morphological measurements &#40;mm&#41; among <span class="elsevierStyleItalic">Brachypelma vagans</span> males &#40;gray&#41; and females &#40;white&#41; for the site in Campeche State&#46; For abbreviations of morphological measurements see <a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>&#46; Mann&#8211;Whitney <span class="elsevierStyleItalic">U</span>-test&#58; ns&#44; not significant&#59; &#42;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>0&#46;05&#59; &#42;&#42;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>0&#46;01&#59; &#42;&#42;&#42;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>0&#46;001&#46;</p>"
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                  \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="top" scope="col" style="border-bottom: 2px solid black">Locality &#40;state&#41;&#59; abbreviation&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">CT&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black"><span class="elsevierStyleItalic">T</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black"><span class="elsevierStyleItalic">P</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">Summer rain&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="top">Paso de Milpa &#40;Veracruz&#41;&#59; V-PM&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm subtropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">800&#8211;1&#44;000&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Low winter precipitation &#40;&#60;5&#37;&#41;&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="top">11 de Mayo &#40;Campeche&#41;&#59; C-11M&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm subtropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">1&#44;000&#8211;1&#44;200&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">High winter precipitation &#40;&#62;10&#46;2&#37;&#41;&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="top">Frontera Corozal &#40;Chiapas&#41;&#59; CH-FC&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm tropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">2&#44;000&#8211;2&#44;500&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Medium winter precipitation &#40;5&#37;<span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>WP<span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>10&#46;2&#37;&#41;&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="top">El Casta&#241;o &#40;Chiapas&#41;&#59; CH-EC&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm tropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">2&#44;000&#8211;2&#44;500&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Low winter precipitation &#40;&#60;5&#37;&#41;&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="top">Cob&#225; &#40;Quintana Roo&#41;&#59; QR-CB&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm subtropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">1&#44;200&#8211;1&#44;500&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">High winter precipitation &#40;&#62;10&#46;2&#37;&#41;&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="top">Rancho Guadalupe &#40;Cozumel&#41;&#59; QR-RG&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">Warm tropical&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">22&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">1&#44;500&#8211;1&#44;800&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">High winter precipitation &#40;10&#46;2&#37;&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr></tbody></table>
                  """
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          "en" => "<p id="spar0025" class="elsevierStyleSimplePara elsevierViewall">Climatic and seasonal characteristics of the sampling sites where <span class="elsevierStyleItalic">Brachypelma vagans</span> individuals were collected&#46; CT&#44; climate type&#59; <span class="elsevierStyleItalic">T</span>&#44; annual mean temperature in degrees&#59; <span class="elsevierStyleItalic">P</span>&#44; annual precipitation in mm&#46;</p>"
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        "etiqueta" => "Table 2"
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                  \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="" valign="top" scope="col" style="border-bottom: 2px solid black">&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">C-11M<span class="elsevierStyleItalic">N</span><span class="elsevierStyleInf">F</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>45&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">V-PM<span class="elsevierStyleItalic">N</span><span class="elsevierStyleInf">F</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>10&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">QR-RG<span class="elsevierStyleItalic">N</span><span class="elsevierStyleInf">F</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>9&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">CH-EC<span class="elsevierStyleItalic">N</span><span class="elsevierStyleInf">F</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>6&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</th><th class="td" title="table-head  " align="left" valign="top" scope="col" style="border-bottom: 2px solid black">KW test&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="top">PI&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">7&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3 &#40;a&#41;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">9&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">11&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">8&#46;6<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;2 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">8&#46;4<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&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="top">TIV&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">11&#46;9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;1 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&#42;&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="top">Lpro&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">19&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;8 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">20&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;5 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">15&#46;5<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;5 &#40;a&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">19&#46;8<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;6 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">18&#46;6<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;7 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">18&#46;9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;3 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&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="top"><span class="elsevierStyleItalic">W</span>&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">8&#46;1<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;6 &#40;a&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">21&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1&#46;5 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">NP&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="char" valign="top">13&#46;7<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>0&#46;8 &#40;b&#41;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="table-entry  " align="left" valign="top">&#42;&#42;&#42;&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td></tr></tbody></table>
                  """
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          "en" => "<p id="spar0030" class="elsevierStyleSimplePara elsevierViewall">Mean &#40;&#177;standard error&#41; for all morphological measures at sites with different number of females tarantulas&#46; C-11M&#44; 11 de Mayo site in Campeche&#44; V-PM Paso de Milpa in Veracruz&#59; CH-EC&#44; El Casta&#241;o in Chiapas&#59; QR-RG&#44; Rancho Guadalupe in Cozumel Island Quintana Roo&#46; <span class="elsevierStyleItalic">N</span><span class="elsevierStyleInf">F</span>&#44; number of female individuals&#59; morphological measurements in mm&#59; PI&#44; and PIV&#44; length of patella I and IV respectively&#59; TI and TIV&#44; length of tibia I and IV&#44; respectively&#59; Lpro and Wpro&#44; length and width of the prosoma&#44; respectively&#46; <span class="elsevierStyleItalic">W</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>weight &#40;in g&#41;&#46; Kruskal&#8211;Wallis test&#58; &#42;&#42;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>0&#46;01&#59; &#42;&#42;&#42;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#60;<span class="elsevierStyleHsp" style=""></span>0&#46;001&#59; NP&#44; not possible&#46; Letters following means represent intergroup differences &#40;post hoc test&#41; for each morphometric measurement&#46;</p>"
        ]
      ]
    ]
    "bibliografia" => array:2 [
      "titulo" => "References"
      "seccion" => array:1 [
        0 => array:2 [
          "identificador" => "bibs0005"
          "bibliografiaReferencia" => array:50 [
            0 => array:3 [
              "identificador" => "bib0005"
              "etiqueta" => "Amarello et al&#46;&#44; 2010"
              "referencia" => array:1 [
                0 => array:2 [
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                      "titulo" => "Potential environmental influences on variation in body size and sexual size dimorphism among Arizona populations of the western diamond-backed rattlesnake &#40;<span class="elsevierStyleItalic">Crotalus atrox</span>&#41;"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => true
                          "autores" => array:6 [
                            0 => "M&#46;E&#46; Amarello"
                            1 => "M&#46; Nowak"
                            2 => "E&#46;N&#46; Taylor"
                            3 => "G&#46;W&#46; Schuett"
                            4 => "R&#46;A&#46; Repp"
                            5 => "P&#46;C&#46; Rosen"
                          ]
                        ]
                      ]
                    ]
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                  "host" => array:1 [
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                      "Revista" => array:5 [
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                      ]
                    ]
                  ]
                ]
              ]
            ]
            1 => array:3 [
              "identificador" => "bib0010"
              "etiqueta" => "Armbruster et al&#46;&#44; 2001"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "Geographic variation and the evolution of reproductive allocation in the pitcher-plant mosquito&#44; <span class="elsevierStyleItalic">Wyeomyia smithii</span>"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:4 [
                            0 => "P&#46; Armbruster"
                            1 => "W&#46;E&#46; Bradshaw"
                            2 => "K&#46; Ruegg"
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                          ]
                        ]
                      ]
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                  ]
                  "host" => array:1 [
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                      "Revista" => array:6 [
                        "tituloSerie" => "Evolution"
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                        "volumen" => "55"
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                        "link" => array:1 [
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                            "web" => "Medline"
                          ]
                        ]
                      ]
                    ]
                  ]
                ]
              ]
            ]
            2 => array:3 [
              "identificador" => "bib0015"
              "etiqueta" => "Bidau and Mart&#237;&#44; 2007"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "<span class="elsevierStyleItalic">Dichroplus vittatus</span> &#40;Orthoptera&#58; Acrididae&#41; follows the converse to Bergmann&#39;s rule although male morphological variability increases with latitude"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:2 [
                            0 => "C&#46;J&#46; Bidau"
                            1 => "D&#46;A&#46; Mart&#237;"
                          ]
                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:2 [
                      "doi" => "10.1017/S0007485307004749"
                      "Revista" => array:6 [
                        "tituloSerie" => "Bulletin of Entomological Research"
                        "fecha" => "2007"
                        "volumen" => "97"
                        "paginaInicial" => "69"
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                        "link" => array:1 [
                          0 => array:2 [
                            "url" => "https://www.ncbi.nlm.nih.gov/pubmed/17298684"
                            "web" => "Medline"
                          ]
                        ]
                      ]
                    ]
                  ]
                ]
              ]
            ]
            3 => array:3 [
              "identificador" => "bib0020"
              "etiqueta" => "Blanckenhorn and Demont&#44; 2004"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "Bergmann and converse Bergmann latitudinal clines in arthropods&#58; two ends of a continuum&#63;"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:2 [
                            0 => "W&#46;U&#46; Blanckenhorn"
                            1 => "M&#46; Demont"
                          ]
                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:2 [
                      "doi" => "10.1093/icb/44.6.413"
                      "Revista" => array:6 [
                        "tituloSerie" => "Integrative and Comparative Biology"
                        "fecha" => "2004"
                        "volumen" => "44"
                        "paginaInicial" => "413"
                        "paginaFinal" => "424"
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                          0 => array:2 [
                            "url" => "https://www.ncbi.nlm.nih.gov/pubmed/21676727"
                            "web" => "Medline"
                          ]
                        ]
                      ]
                    ]
                  ]
                ]
              ]
            ]
            4 => array:3 [
              "identificador" => "bib0025"
              "etiqueta" => "Blanckenhorn et al&#46;&#44; 2006"
              "referencia" => array:1 [
                0 => array:2 [
                  "contribucion" => array:1 [
                    0 => array:2 [
                      "titulo" => "When Rensch meets Bergmann&#58; does sexual size dimorphism change systematically with latitude&#63;"
                      "autores" => array:1 [
                        0 => array:2 [
                          "etal" => false
                          "autores" => array:5 [
                            0 => "W&#46;U&#46; Blanckenhorn"
                            1 => "R&#46;C&#46; Stillwell"
                            2 => "K&#46;A&#46; Young"
                            3 => "C&#46;W&#46; Fox"
                            4 => "K&#46;C&#46; Ashton"
                          ]
                        ]
                      ]
                    ]
                  ]
                  "host" => array:1 [
                    0 => array:1 [
                      "Revista" => array:6 [
                        "tituloSerie" => "Evolution"
                        "fecha" => "2006"
                        "volumen" => "60"
                        "paginaInicial" => "2004"
                        "paginaFinal" => "2011"
                        "link" => array:1 [
                          0 => array:2 [
                            "url" => "https://www.ncbi.nlm.nih.gov/pubmed/17133857"
                            "web" => "Medline"
                          ]
                        ]
                      ]
                    ]
                  ]
                ]
              ]
            ]
            5 => array:3 [
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        "texto" => "<p id="par0085" class="elsevierStylePara elsevierViewall">We would like to thank the people from the villages visited during sampling for granting us access to their land and for their hospitality during our stay&#46; Thanks to the reviewers and to the associate editor of the Revista Mexicana de Biodiversidad for the suggestions and comments that improved our work&#46; Thanks to Claudia Vilchis-Nestor for the information on the Cozumel population of <span class="elsevierStyleItalic">Brachypelma vagans</span>&#46; The research was financed by <span class="elsevierStyleGrantSponsor" id="gs0005">Conacyt project</span> 000000000158138&#46; This study was conducted according to relevant national and international guidelines&#46; Permit &#35; FAUT-0241 was granted to Dr&#46; Yann H&#233;naut&#44; issued by the Secretar&#237;a de Medio Ambiente y Recursos Naturales&#44; according to the Norma Oficial Mexicana NOM-126-ECOL-2000&#46;</p>"
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Article information
ISSN: 18703453
Original language: English
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