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Programa de Estudios de Posgrado HISTORIA DE VIDA DE Homalonychus selenopoides (ARANEAE: HOMALONYCHIDAE) EN EL DESIERTO DE SONORA TESIS Que para obtener el grado de Doctor en Ciencias Uso, Manejo y Preservación de los Recursos Naturales ( Orientación en Ecología ) P r e s e n t a JOSÉ ANDRÉS ALVARADO CASTRO La Paz, Baja California Sur, Agosto de 2011 iii COMITÉ TUTORIAL Dra. María Luisa Jiménez Jiménez Directora de Tesis Centro de Investigaciones Biológicas del Noroeste, S.C. (CIBNOR) Dr. Yann Lucien Hénaut Seguin Co-Tutor El Colegio de la Frontera Sur (ECOSUR) Dra. Aurora Margarita Breceda Solís Cámara Co-Tutora Centro de Investigaciones Biológicas del Noroeste, S.C. (CIBNOR) Dra. María del Cármen Blázquez Moreno Co-Tutora Centro de Investigaciones Biológicas del Noroeste, S.C. (CIBNOR) Dr. Alejandro Manuel Maeda Martínez Co-Tutor Centro de Investigaciones Biológicas del Noroeste, S.C. (CIBNOR) iv COMITÉ REVISOR DE TESIS Dra. María Luisa Jiménez Jiménez (CIBNOR) Dr. Yann Lucien Hénaut Seguin (ECOSUR) Dra. Aurora Margarita Breceda Solís Cámara (CIBNOR) Dra. María del Cármen Blázquez Moreno (CIBNOR) Dr. Alejandro Manuel Maeda Martínez (CIBNOR) JURADO DE LA DEFENSA DE TESIS Dra. María Luisa Jiménez Jiménez (CIBNOR) Dr. Alejandro Manuel Maeda Martínez (CIBNOR) Dr. Ricardo Rodríguez Estrella (CIBNOR) Dra. Sara Cecilia Díaz Castro (CIBNOR) Dra. Yolanda Maya Delgado (CIBNOR) SUPLENTE Dra. Patricia Galina Tessaro (CIBNOR). v RESUMEN Homalonychus selenopoides Marx, 1891 es una especie de araña endémica de las planicies costeras del Desierto Sonorense en el estado de Sonora, México y del suroeste de los Estados Unidos de Norteamérica. Aunque la especie fue descrita taxonómicamente hace más de un siglo, nada se sabe acerca de su biología y ecología. El propósito general del presente trabajo fue caracterizar la historia de vida de esta especie; particularmente, su fenología y microhábitat en el campo, y su comportamiento reproductivo, defensivo, y ciclo de vida en el laboratorio. Estos rasgos fueron comparados con los reportados para H. theologus para inferir sobre su valor adaptativo. Durante el 2008 se recolectaron 473 especímenes y 67 ovisacos en el sur del Desierto Sonorense, en México. En el campo, el ciclo de vida es anual y la fenología es marcadamente estacional. Los ovisacos, al igual que los reclutas, ocurren en el campo sólo de abril a julio, juveniles todo el año, machos de noviembre a abril y hembras de noviembre a mayo. Se encontró una correlación lineal negativa entre las temperaturas promedio y la abundancia de adultos (R = -0.9353; P < 0.0001), como un posible indicador de la influencia de la temperatura en el entorno biótico de la región. Se criaron dos lotes de arañas para el estudio de ciclo de vida. Después de la oviposición, las arañuelas de instar II tardan ~50 días en emerger del ovisaco. Los machos maduran después de 8-10 mudas y las hembras después de 9-11 mudas. Los machos maduraron más rápido (577.3 ± 35.9 días) que las hembras (622 ± 11.6 días) (t = 4.499; P < 0.0001). El tiempo de desarrollo de los instares ninfales no se incrementó progresivamente sino que fue irregular, y diferente entre ellos (Lote 1: H = 196.525; P = 0.0000; Lote 2: H = 132.544; P = 0.0000). Las temperaturas promedio del laboratorio y el porcentaje de mudas por mes estuvieron lineal y positivamente correlacionados (Lote 1: R = 0.6633; P < 0.005; Lote 2: R = 0.8466; P < 0.0001). Un alto porcentaje de arañas murieron antes de alcanzar el estado adulto (69.6% en el Lote 1 y 85.9% en el Lote 2); esta mortalidad estuvo asociada principalmente a la muda (69.6% en el Lote 1 y 65.6% en el Lote 2). Una proporción importante de estas muertes se asoció a ataques de larvas de Tenebrio contra las arañas antes o durante la muda. Las curvas de sobrevivencia resultaron intermedias entre el Tipo I y el Tipo II de Pearl (1928). La longevidad de machos y hembras recolectados en campo como instares tardíos fue de 82.9 ± 23.8 días y de 481.4 ± 183.5 días, respectivamente. El comportamiento reproductivo fue analizado en el laboratorio, para lo cual se grabó con una cámara infrarroja, principalmente en la noche. La inducción de esperma es de tipo indirecto; los machos tejieron una tela espermática de aproximadamente 2 cm2 cerca del suelo. Los machos y las hembras elaboran hilos de seda y arena, como rastros. El comportamiento de cortejo fue intermedio entre los niveles I y II, y la posición de cópula fue de Tipo 3 modificado, donde el macho ata las patas de la hembra con seda antes de copular. El canibalismo sexual puede ocurrir durante o después de la cópula. Las hembras inician la construcción de su ovisaco ~11 días después del copular y lo concluyen en ~15 h, incluyendo la oviposición. La lámina externa del ovisaco contiene arena, y el saco estuvo rodeado por una guarnición de cordones de seda y arena, posiblemente para proteger a los huevos de la desecación y como una barrera contra vi parásitos y depredadores. Los juveniles, los subadultos y los adultos exhiben todos los actos del comportamiento de empolvado y enterramiento. Aunque los machos exhiben estos comportamientos, las partículas de suelo no se adhieren a las sedas de su cutícula. El trampeo pitfall fue ineficiente para la recolecta general de individuos de esta especie, pero resultó valioso para indicar su época reproductiva. Esta es una de las pocas especies de arañas que son activas en invierno y que los adultos ocurren y tienen su actividad sexual en este periodo. Su comportamiento reproductivo, defensivo, su ciclo de vida y su fenología le confieren aptitud y especialidad para existir en el Desierto Sonorense. Palabras clave: Comportamiento reproductivo, ciclo de vida, fenología. Resumen aprobado por ___________________________________ Dra. María Luisa Jiménez Jiménez Directora de tesis vii ABSTRACT Homalonychus selenopoides Marx, 1891 is an endemic spider species to the coast plains of the Sonoran Desert. It is distributed from Sonora, México to the southeastern part of the United States. Although this species was described for more than a century, nothing is known about its biology and ecology. The overall purpose of this study was to characterize the life history of this species, particularly its phenology and microhabitat in the field, and its reproductive and defensive behavior and life cycle in the laboratory. These features were compared with those reported for H. theologus to infer about its adaptive value. Throughout 2008 we collected 473 spiders and 67 egg sacs at the southern part of the Sonoran Desert in México. In the field the species’ life cycle is annual and its phenology is markedly seasonal. Egg sacs and recruits occur in the field only from April-July, juveniles throughout the year, males from November to April and females from November to May. Average temperature and adult frequency were negatively correlated (R = -0.9353; P < 0.0001) as a possible indicator of the temperature influence on the surrounding biotic environment. Two trials of spiders were reared to the life cycle survey. After oviposition, the spiderlings of the second instar took ~50 days to emerge from the egg sac. Males matured after 8-10 molts and females after 9-11 molts. Males matured faster (577.3 ± 35.9 days) than females (622 ± 11.6 days) (t = 4.499; P < 0.0001). Development time of the nymph instars did not increase gradually, but it was irregular and different between them (Trial 1: H = 196.525; P = 0.0000; Trial 2: H = 132.544; P = 0.0000). Lab mean temperature and monthly molt percentage were lineal and positively correlated (Trial 1: R = 0.6633; P < 0.005; Trial 2: R = 0.8466; P < 0.0001). A high percentage of spiders died before maturity (69.6% in Trial 1 and 85.9% in Trial 2), mainly during molting, including attacks by Tenebrio larvae before or during molting. Survivorship curves were intermediate between Type I and Type II of Pearl (1928). Longevity of male and females collected in the field as late instars was 82.9 ± 23.8 days and 481.4 ± 183.5 days, respectively. The species’ reproductive behavior was recorded mainly at night with an infrared camera and analyzed in the laboratory. Sperm induction is indirect; males wove a spermatic web about 2 cm2 in size near the ground. Males and females make silk threads and sand. Courtship behavior was considered between levels I and II; mating position was Type 3 modified, where the male ties the female’s legs with silk before mating. Sexual cannibalism can take place during or after mating. Females began to spin their egg sac ~11 days after mating and completed it in ~15 h, including oviposition. The outer layer of the egg sac contained sand; the egg sac was surrounded by a garniture of cords of silk and sand, most likely to protect the eggs from desiccation and as a barrier against predators and parasites. Juveniles, sub-adults, and adults show self-burying behavior and self-covering with sand. Although males show these behaviors, their setae do not retain soil particles. Pitfall sampling was inefficient to general collection of these spiders, but it was invaluable to know their reproductive season. This is one of the few spider species that are active in winter, and adults are viii also sexually active during this season. Their reproductive and defensive behavior, life cycle and phenology are adaptations to survive and to be a successful spider species in the Sonoran Desert. Key words: Reproductive behavior, life cycle, phenology. ix DEDICATORIA Con mucho amor a mi Familia. A mi esposa Sandra, quien ha sabido llevar varios años de abandono.
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    ΠΕΡΙΕΧΟΜΕΝΑ ΠΕΡΙΛΗΨΗ 3 EXTENDED SUMMARY 5 1. ΕΙΣΑΓΩΓΗ 7 1.1 Μιτοχόνδριο 7 1.1.1 Λίγα Λόγια για το DNA 7 1.1.2. Κληρονόμηση Μιτοχονδριακού DNA 9 1.1.3. Ζωικό μιτοχονδριακό DNA 9 1.1.4. Μειονεκτήματα μιτοχονδριακών γενετικών δεικτών 15 1.1.5. Φυτικό μιτοχονδριακό DNA 16 1.2 Οργανισμοί μελέτης 18 1.2.1 Γνωριμία με τους οργανισμούς 18 1.2.2. Αρθρόποδα 18 1.2.3 Χηληκεραιωτά 20 1.2.4. Αραχνίδια 20 1.2.5. Αράχνες 20 1.2.6. Λίγα λόγια για τις Οικογένειες Αραχνών της μελέτης 25 1.3 DNA Barcoding στις αράχνες 40 1.3.1. Γενικά 40 1.3.2 Παρουσίαση της ιστοσελίδας www.boldsystems.org 44 1.3.2.1 Database 46 1.3.2.2. Taxonomy 47 1.3.2.3. Identification 48 1.3.3. Παρουσίαση της ιστοσελίδας www.araneae.unibe.ch 52 1.3.4. Επιλογή των Δειγμάτων 53 1.3.5 Ελληνικές και Ξένες Μελέτες και Προγράμματα για το DNA Barcoding στις Αράχνες 53 1.4 Σκοπός της Μελέτης 56 2. ΥΛΙΚΑ και ΜΕΘΟΔΟΙ 57 2.1. Επιλογή και Συλλογή Δειγμάτων 57 2.2. Είδη, Οικογένειες και Παραομάδες 58 2.3. Πειραματική Διαδικασία 62 2.3.1. Εξαγωγή ολικού γενωμικού DNA 62 2.3.2. Πολλαπλασιασμός των γονιδίων-στόχων μέσω της PCR 64 1 2.3.3. Καθαρισμός του προιόντος της PCR 65 2.4. Επεξεργασία των αλληλουχιών 66 2.5 Εκτίμηση γενετικών αποστάσεων 66 2.6 Φυλογενετικές αναλύσεις 67 3. ΑΠΟΤΕΛΕΣΜΑΤΑ 68 3.1. Γενικά στοιχεία για τις αλληλουχίες του COI 68 3.2.
  • Evidence That Olfaction-Based Affinity for Particular Plant Species Is a Special Characteristic of Evarcha Culicivora, a Mosquit

    Evidence That Olfaction-Based Affinity for Particular Plant Species Is a Special Characteristic of Evarcha Culicivora, a Mosquit

    Evidence that olfaction-based affinity for particular plant species is a special characteristic of Evarcha culicivora, a mosquito-specialist jumping spider Running title: Plant affinity in a jumping spider Ximena J. NELSON1*, Robert R. JACKSON1,2 1School of Biological Sciences, University of Canterbury, Private Bag 4800, Christchurch, New Zealand. 2International Centre of Insect Physiology and Ecology, Thomas Odhiambo Campus, P.0. Box 30 Mbita Point, Kenya. *Email: [email protected] Phone: 64-3-3642987 extn. 4050 Fax: 64-3-3642590 Key words: Plant-arthropod interactions, Evarcha culicivora, Lantana camara, Salticidae, olfaction ABSTRACT. Evarcha culicivora, an East African jumping spider (family Salticidae), was shown in an earlier study to have an affinity for the odor from two particular plant species, namely Lantana camara and Ricinus communis. The olfactometer used in the earlier study was designed for choice testing. Here we focus on L. camara and, by using a second olfactometer method (retention testing), add to the evidence that the odor of this plant is salient to E. culicivora. Another 17 East African salticid species, all from different genera, were investigated using the same two olfactometer designs as used when investigating E. culicivora. The number of individuals of each of these 17 species that chose L. camara odor was not significantly different from the number that chose a no-odor control and, for each species, the latency to leave a holding chamber (retention time) in the presence of L. camara odor was not significantly different from retention time in the presence of a no-odor control. Based on these findings, we conclude that, rather than being a widespread salticid characteristic, an affinity for the odor of L.