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(Amphibia, Anura): Morfología Y Desarrollo Del Sistema Esquelético

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(Amphibia, Anura): Morfología Y Desarrollo Del Sistema Esquelético UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Biología Animal I (Zoología) TESIS DOCTORAL Evolución de Pelobatidos y Peloditidos (Amphibia, Anura): morfología y desarrollo del sistema esquelético MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR María del Rosario Rodríguez Talavera Director Francisco de Borja Sanchiz y Gil de Avalle Madrid Ed. electrónica 2019 © María del Rosario Rodríguez Talavera, 1989 /L °'`^`^^ Departamento de Biología Animal I Facultad de Ciencias Biológicas Universidad Complutense de Madrid ireeemema HVOLIICIOB DH PHLOBATIDOS Y PHLODITIDOS (A1^[PHIBIA, AHURA): XORFOLOGIA Y DHSARROLLO DHL SISTHxA HSQIIñLHTICO Memoria presentada para optar al grado de doctor en Ciencias Biológicas por l[aría del xosarío Rodríguez Talavera v^ B^ del director de la tesis ^^%^ ^a ^-...^r ^ Dr. D. Francisco de Borja Sanchíz y Gil de Avalle Invest:igador del CSIC en el Museo Nacional de Ciencias Iiaturales Madrid, octubre de 1989 AGRADfiCII[ IfiHTOS En primer lugar deseo expresar mi agradecimiento al Dr. Sanchíz, director de esta tesis, por sugerirme el tema de investigación y por su valiosa colaboración, tolerancia y acertadas críticas durante el desarrollo del trabajo. E1 Dr. Castanet CLaboratoire d'Anatomie Comparée, IJniversité Paris VII) revisó amablemente los resultados esqueletocronológicos. Aumerosas personas me acompañaron en los viajes de recolección de material. Especialmente agradezco la colaboración de Begoña Arano y Asunción Rodríguez Espeso en Portugal; Juan Carlos Arrechea y Jesús Dorda en 14arruecos, y Marisa Esteban en el sur de Espa4ia. Las siguientes personas colectaron material adicional para este estudio: Begoña E►rano, Ignacio Doadrio, Jesús Dorda, Carmen Díaz-Paniagua, Marisa Esteban, Mario García-Paris, Paloma Garzón, Rafael Márquez y Carolina Martín- Albaladejo. Rosa María Gómez-Prieto colaboró pacientemente en el trabajo de laboratorio, y mecanografió las tablas. Luis García-Hidalgo, Jose Enrique Gonzalez y Miguel Tejedo me ayudaron a abordar l.a estadística. Antonio Madrigal, Manolo Merino y Rafael Merino, pasaron a tinta las figuras. Angel Sanz, con su dominio de las técnicas fotagráficas, mejoró la resolució:n de las preparaciones histológicas. Pilar Herrero, Carmen Díaz-Paniagua, Rosa María Gómez, Mercedes Hervás y Teresa Aparicio me animaron constantemente en el curso de la investigación. Juan Carlos Arrechea me acompañó en innumerables y tediosos muestreos semanales por la provincia de Madrid todos estos años. Realizó los últimos retoques en las figuras y me ayudó a comprobar las referencias internas del texto. E1 personal del Museo Nacional de Ciencias Naturales de Madrid facilitó mi estancia en el Centro y el desarrollo del trabajo. Esi;e trabajo ha sido parcialmente financiado con una beca predoctoral del CSIC en el Museo Nacional de Ciencias Naturales de Madrid y una beca de intercambio entre el CSIC y el CNRS en el Laboratoire d'Anatomie Comparée de la Université de Paris VII. La investigación se incluye en el proyecto CSIC- CAICYT 2].1 (1985-1988). IHD ICE 1. IHTRODUCCIOH 1 2. li[ATERIALES, TECHICAS AHALITICAS Y I^[ETODOLOGIA 12 2 .1. l[ATERIALES 13 2 .2 . TECH ICAS DE ^[iJESTRBO 15 2.3. TBCHICAS HISTOLOGICAS 17 2.3.1. Tínciones de cartílago y hueso 1? 2.3.2. Preparación de esqueletos desarticulados 20 2.3.3. Técnicas de esqueletocronología 21 2.4 . PROTOCOLO DB L05 S%PER Il[fiHT05 DB CREC Il[ IBHTO Y DESARROLLO LARYARIO 25 2.4.1. Bsperinento para determinar la temperatura óptiaa de crecíaiento 25 2.4.2. Bsperimento para deter^inar la edad a que se alcan?-A cada estadio de desarrollo 25 2.5. DBFIHICIOB Dfi LA BIOXfiTRIA fi%TERBA 26 2.5.1. Hedidas esternas de renacuajos 26 2.5.2. l[edidas egternas de adultos 27 2.6. AHALISIS CIIAHTITATIVOS 30 3. RESULTADOS Y DISCUSIOH 31 3.1. TRABAJOS PRELIxIHARBS 32 3.1.1. Pelobates cultri^. TEIIPEBATURA OPTIli[A DB CREC Il[ IfiHTO LARVAB IO 32 3.12. Pelobates cultri,^,ç. BDAD A QIIE SE ALCAHZA CADA BSTADIO DE DBSARROLLO DBL PERIODO LARVARIO A TEPIPERATURA OPTU[A 36 3.1.3. TALLA fiH CADA BSTADIO DE DESARROLLO DE Pelobates cultri^ Y Pel^iy^^unctatus 40 3.1.4. BS"TRUCTURA POR SB%OS Y BDADES DE LAS POBLACIOHES POSTIi[fiTAI[ORFICAS DBL GBHERO Pelobates AHALIZADAS 46 .1. Pelobates cultri^ 46 .2. Pelobates varaldii 56 .3. Pelobates fuscus y P. s9riacus 60 3.1.5. TAl[AfFO DB LOS EJBI[PLARES !lDULTOS 62 .1. Dimorfismo ^^^1 en Pelobates cultr^^ y P. varaldii 62 .2. Diferencias morfométricas intraespecíficas en P_elobates cultri^ 66 .3. Tamaño medio de los adultos de las especies estudiadas 70 3.1.6. RBLACIOH fiFTRfi fiDAD Y TAl[A4í0 BH Pelobates cultriTnes y P. varaldii 72 3.2. DESARROLLO DE LOS SISTExAS ESQ►ELETICOS DE Pelobates cult_^ Y Pel^ es punctatus 77 3.2.1. CRA^fEO 7? .1. Descripción del cráneo adulto de Pelobates cultripes ?? .2. Secuencia de asificación del cráneo de P. cultrip^, 77 .3. Comparación con Pelgjytes nunctatus y otras especies de Anuros 95 3.2.2. COLUl^[IfA VERTEBRAL 118 .1. Descripción de la columna adulta de Pelobates cultri^ 118 .2. Desarrollo de la columna vertebral de Pelobates cultr1p^ç 118 .3. Secuencia de asificación de la columna vertebral de P. cultri,^ 120 .4. ^[alformaciones de la columna vertebral de P. cultri^ 129 .5. Comparación con Pelody}es ^^nctatus y las restantes especies de Anuros 136 3.2.3. XIEXBRO AHTERIOR Y CIBTiiRA ESCAPIILAR 150 .1. Descripción del miembro anterior y de la cintura escapular adulta de Pelo ^tes cultr ^ 150 .2. Secuencia de osificación del miembro anterior y cintura escapular de P. cultrines 150 .3. Comparación con Pel^^,punctatus y otras especies de Anuros 170 3.2.4. XIfiIIBRO POSTBRIOR Y CIHTURA PfiLVIAHA 195 .1. Descripción del miembro posterior y de la cintura pelviana de Pelobates sstl^fg^ 195 .2. Secuencia de osificación del aieabro posterior y cintura pelviana de P. cultrip^ 195 .3. Comparación con PB_l^y.#^_^unctatus y otras especies de Anuros 214 4. COHCLUSIOHES GBHfiRALfiS 246 5. B IBL IOGRAF IA 254 6. APfiIfDICB 2^g 1. IBTRODIICCIOIf - 2 - 1. IHTRODIICCIOH Los caracteres osteológicos juegan un papel fundamental en la macrosistemáticá de los anuros. Lynch <1973) utilizó^ 38 caracteres en su análisis evolutivo del orden; vefntitrés de ellos, casi las dos terceras partes, eran osteológicos. A1 mismo tiempo, el esqueleto es casi el único sistema orgánico disponible para estudios paleontológicos y evolutivos del pasado. Por otra parte, podemos afíadir la utilidad del sistema esquelético para estudios de biología evolutiva; existen tendencias recientes en esta disciplina que hipotetizan alteraciones en la duración relativa de los acontecimientos del desarrollo Cheterocronías) como una de las causas fundamentales del cambio morfológico (Alberch et al., 1979). Este trabajo revisa y actualiza los conocimientos existentes sobre la osificación del esqueleto de los anuros. Paralelamente examina el proceso de condrificación de los miembros anteriores y posteriores, especialmente en lo referente al carpo y al tarso, y de sus respectivas cinturas. Su aportación más original reside en el estudio del proceso de osificación con una visión muy amplia del desarrollo, pues considera parte importante del mismo no sólo todos los estadios larvarios, sino también todas las clases de edad de las etapas juvenil y adulta. A1 mismo tiempo, determina la variabilidad de caracteres osteológicos de importancia sistemática, y establece homologías atendiendo al origen embriológico de los elementos. De hecho, el método de trabajo permite utilizar ejemplares preservados en alcohol o formol largo tiempo y, específicamente, integrar datos procedentes de series ontogénicas fósiles. Los estudios de este tipo son necesarios, pues los parámetros básicos para dilucidar la participación de procesos heterocrónicos en los cambios morfológicos todavía no se han cuantificado en el esqueleto completo de ningún anuro. Tales parámetros son: momentos de inicio y finalización de la osificación, tamaño de las piezas al comienzo de la misma, y tasa de crecimiento de los elementos esqueléticos. Por lo cual, es evidente la necesidad de conocer con detalle la cronología de la diferenciación de todos los elementos esqueléticos de algunas especies clave, medir en ellas el mayor ^número de parámetros del desarrollo y precisar su variabilidad - 3 - intraespecífica. Esto nos proporcionaría un marco de referencia para determina.r qué parámetros y procesos del desarrollo del esqueleto tienen utilidad taxonómica y filogenética, así como identificar caracteres que resulten convergentes en la etapa adulta. Hasta el momento se ha investigado el proceso de osificación del cráneo de dieciacho especies de anuros, de las 3.438 existentes <Duellman & Trueb, 1986). Estas especies pertenecen a ocho de las veintidós famílias reconocidas (Duellman & Trueb, 1986). Para nuestros propósitos tres de estos trabajos son importantes, ya que se realizaron con un número suficiente de ejemplares y se estudió toda la etapa larvaria: - Kemp & Hoyt <1969> sobre Rana ^piens. Estos autores además de realizar una descripción completa y detallada de la cronología de la osificación de todos los huesos de cráneo, describen dos alteraciones de la secuencia de osificación normal en renacuajos tratados con tiroxina en estadios tempranos del desarrollo. Entre los inconvenientes a la hora de utilizar sus datos en nuestros estudios camparativos, podemos segalar que utilizaron una tabla de desarrollo específica de Rana pipiens, la de Taylor & Kollros (1946>; además, la descripción del progreso de la osificación se interrumpe al fin de la metamorfosis, y por último, los animales fueron criados en cautividad. Gaudin C1978) sobre Bufo boreas. En este trabajo se comparan las secuencias de osificación de renacuajos de campo de dos localidades distintas con renacuajos de laboratorio. Se pone de manifiesto que la osificación en los animales de laboratorio se retrasa respecto a los del campo. Se reseñan todos los huesos y se describe el avance de la osif.icación en la. etapa postmetamórfica, dividiendo los ejemplares en clases de tamaño. Se utiliza la tabla de desarrollo de Limbaud 8^ Volpe (19:i7), muy parecida a la de Gosner C1960>.
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