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Orden STOMATOPODA Manual Revista IDE@-SEA, nº 84 (30-06-2015): 1-10. ISSN 2386-7183 1 Ibero Diversidad Entomológica @ccesible www.sea-entomologia.org/IDE@ Clase: Malacostraca Orden STOMATOPODA Manual CLASE MALACOSTRACA Orden Stomatopoda Pere Abelló1 & Guillermo Guerao2 1 Institut de Ciències del Mar (CSIC), Passeig Marítim 37-49, 08003 Barcelona (España) [email protected] 2 Passeig Fabra i Puig 344, 08031 Barcelona (España) 1. Breve definición del grupo y principales caracteres diagnósticos Los estomatópodos o galeras son crustáceos marinos bentónicos de talla relativamente grande y hábitos depredadores, restringidos principalmente a las aguas someras tropicales, aunque algunas especies habitan el talud continental hasta profundidades de unos 1500 m (Manning, 1991). Su cuerpo es alargado y una de sus principales características morfológicas es la presencia del segundo par de apéndices torá- cicos modificados en forma de pinza subquelada prensil de morfología similar a la de las mantis religio- sas. Se conocen en la actualidad un total de casi 500 especies pertenecientes en la actualidad a un único suborden, los Unipeltata, constituido por siete superfamilias: Bathysquilloidea, Erythrosquilloidea, Eurys- quilloidea, Gonodactyloidea, Lysiosquilloidea, Parasquilloidea y Squilloidea (Ahyong & Schram, 2013). En la actualidad se reconocen un total de 17 familias de estomatópodos. El registro fósil nos indica que el origen de los estomatópodos se encuentra en el Devónico. Este orden se encuentra actualmente en pleno estudio, tanto a nivel biológico, fisiológico y ecológico como a nivel de sistemática y filogenia. Diagnosis: El pleon (abdomen) está más desarrollado que el cefalotórax, de apariencia reducida (no cubre todos los segmentos torácicos); los ojos suelen ser pedunculados y lobulados; las anténulas pre- sentan tres flagelos; la escama antenal o escafocerito tiene dos artejos, el basal generalmente pequeño; los toracópodos o apéndices torácicos tienen protopodios tri-segmentados (pre-coxa, coxa y basis) y endópodos (o rama interna) con cuatro segmentos; las branquias, dendrobranquiadas, se originan en las bases de los pleópodos. 1.1. Morfología El cuerpo está constituido por tres tagmas: céfalon o cabeza, pereion o tórax y pleon o abdomen. El céfa- lon está constituido por cinco somitos (excluyendo el acron), el pereion por ocho y el pleon por seis (ex- cluyendo el telson). El telson está bien desarrollado y se encuentra ocasionalmente fusionado con el sexto somito abdominal formando un pleotelson. Posteriormente al acron o somito oftálmico se presentan los cinco somitos cefálicos (con anténulas, antenas y apéndices bucales: mandíbulas, maxílulas y maxilas). Los cinco primeros segmentos torácicos se denominan a veces maxilípedos, mientras que los tres restantes constituyen los toracópodos o patas marchadoras. El caparazón está bien desarrollado y cubre dorsalmente el céfalon (excepto el acron o región pre- segmental y el somito antenular) y la mitad anterior del pereion. Los ojos son pedunculados y compuestos y presentan la córnea lobulada. El ojo compuesto de los es- tomatópodos es muy complejo, y en algunas especies tropicales ha sido considerado como el ojo más complejo y diverso de la biosfera (Marshall, 1988; Bok et al., 2013). Revista IDE@-SEA, nº 84 (30-06-2015): 1-10. ISSN 2386-7183 2 Ibero Diversidad Entomológica @ccesible www.sea-entomologia.org/IDE@ Clase: Malacostraca Orden STOMATOPODA Manual Fig. 1. Morfología externa de la galera, Squilla mantis; (a) vista dorsal; (b) vista lateral. (original de G. Guerao); publicado en Abelló & Guerao (2004). Los ojos de los estomatópodos presentan distintos tipos de fotoreceptores sensibles a la intensidad de luz y a longitudes de onda que van desde el ultravioleta hasta casi prácticamente el infrarrojo. Además, algunos fotoreceptores son sensibles a la polarización de la luz. La estructura física y molecular de estos fotoreceptores es extremadamente compleja. La visión en color es importante en la comunicación entre individuos, así como en el comportamiento reproductor y territorial. El ojo típico de un estomatópodo se presenta estructurado en tres lóbulos, con dos hemisferas, dor- sal y ventral, separadas por una banda media de varias filas de omatidios especializados. Los omatidios de ambas hemisferas poseen básicamente la estructura típica de los ojos compuestos de aposición pre- sentes en los crustáceos. Las principales modificaciones y especializaciones características y únicas de los estomatópodos se presentan en la banda media. La orientación de los omatidios permite obtener una imagen a partir de tres áreas distintas e independientes del ojo. Adicionalmente a la visión solapada de los tres campos visuales, cada ojo puede presentar movimientos independientes dirigidos a distintas áre- as del campo visual presente en cada momento. En algunas familias, los ojos pueden presentar distintas Revista IDE@-SEA, nº 84 (30-06-2015): 1-10. ISSN 2386-7183 3 Ibero Diversidad Entomológica @ccesible www.sea-entomologia.org/IDE@ Clase: Malacostraca Orden STOMATOPODA Manual áreas especializadas, con alta sensibilidad a longitudes de onda diferenciadas o con estructuras dirigidas al aumento de la agudeza visual. En las retinas de los estomatópodos se han detectado un mínimo de once tipos distintos de rodopsi- nas (pigmentos visuales), el máximo número detectado en cualquier otro grupo zoológico. Adicionalmen- te, los fotoreceptores poseen filtros para minimizar ruido y optimizar la sensibilidad a longitudes de onda específicas. El somito antenular suele estar cubierto por la placa rostral, articulada con el caparazón. Las anté- nulas son trisegmentadas y terminan en tres flagelos. Las antenas están formadas por un protopodio anterior o pedúnculo bisegmentado, por un exopodio bisegmentado y por un endopodio trisegmentado provisto de un flagelo. El segmento terminal del exopo- dio es ancho y constituye el escafocerito. Los restantes somitos cefálicos y los cuatro primeros somitos torácicos están cubiertos dorsalmente por el caparazón. Éste presenta normalmente tres surcos en su superficie: uno cervical y dos gástricos paralelos. Las mandíbulas son grandes y presentan un proceso incisivo y un proceso molar. Puede o no existir un palpo mandibular. Las maxílulas constan de un protopodio bisegmentado con un endito en cada segmento, basal y coxal. El endopodio está reducido a un pequeño segmento (palpo). No existe exopodio. Las maxilas son las más externas. Están formadas por cuatro segmentos; los dos primeros (basa- les) presentan enditos más o menos desarrollados. Los cinco primeros pares de apéndices torácicos (o toracópodos) son subquelados y se utilizan pa- ra la predación, alimentación y excavación de madrigueras. Están constituidos por siete artejos, aunque suele haber alguna fusión intersegmental. El segundo par es mucho más largo y fuerte que el resto; el propodio y el dáctilo forman una poderosa pinza subquelada (el dáctilo o segmento terminal se cierra sobre el propodio), de apariencia similar a las pinzas de las mantis religiosas, de donde les viene su nom- bre en inglés, mantis shrimps. Los tres últimos pares de apéndices torácicos son birrámeos y se utilizan en la locomoción. El pro- topodio está formado por tres segmentos. La rama externa está constituida por un único segmento y constituye el endopodio, contrariamente a lo que pudiera parecer. La rama interna, bisegmentada, es el exopodio. Las posiciones relativas del exopodio y el endopodio se han invertido durante el desarrollo. Los gonoporos se localizan en el sexto segmento del pereion en las hembras y en el octavo en los machos. Los machos presentan además un par de estructuras tubulares copulatorias o penes en la base del último par de apéndices torácicos. Existen seis pares de apéndices abdominales o pleópodos. Los cinco primeros suelen llevar bran- quias y tienen también una función natatoria. Se trata de apéndices birrámeos, anchos y aplastados. Constan de un protopodio basal provisto de un endopodio y un exopodio, cada uno con dos segmentos poco conspicuos. Cada segmento distal del endopodio lleva en el apéndice interno o estilamblis una es- tructura denominada retináculo mediante la cual se une a su homólogo. Los movimientos de cada par de pleópodos se ajustan de esta manera y pueden moverse al unísono. En los machos, los endopodios del primer y segundo par de pleópodos están modificados a modo de petasma, aunque el segundo sólo muy ligeramente. El sexto par de apéndices abdominales lo constituyen los urópodos. Éste se encuentra usualmente separado del telson, pero puede haber en ciertos casos una fusión entre los mismos, formándose un pleotelson. Los urópodos consisten típicamente en un protopodio que termina en un exopodio bisegmen- tado y un endopodio unisegmentado. Una proyección ventral de la base del urópodo constituye el proceso bifurcado. El segmento proximal del exopodio está provisto de una serie de espinas móviles. Los urópo- dos forman junto con el telson un amplio abanico natatorio. En la superficie anteroventral del telson se encuentra el poro anal. 1.2. Historia natural Hábitat: Los estomatópodos son crustáceos bentónicos que habitan en madrigueras excavadas en el fango o en oquedades y grietas en zonas rocosas y arrecifes de coral. Reproducción: Son especies ovíparas que cuidan la puesta. Las hembras ovígeras transportan los em- briones en los maxilípedos, aglutinados en una secreción procedente de la glándula del cemento, muy evidente como franjas blanquecinas presentes en hembras maduras entre los tres últimos pares
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