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Orientation and Migration in Talitrus Saltator

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Orientation and Migration in Talitrus Saltator CORE Metadata, citation and similar papers at core.ac.uk Provided by Florence Research Revista Chilcna de Historia Natural 69: 553-563, 1996 Orientation and migration in Talitrus saltator Orientación y migraci6n en Talitrus saltator FELICITA SCAPINI, MARIO FALLACI and MARCO C. MEZZETTI Dipartimento di Biologia animalc e Genetica. Universita di Firenze. via Romana 17,50125 Firenze. ltalia E-mail: scapini @dbag.unifi.it ABSTRACT Talitrus saltator a beach dwelling crustacean amphipod, has been for long the subject of orientation studies. as it shows orientation perpendicular to the shoreline in response to stressful factors of environmental dryness or submersion. Sun orienta- tion permits zonal recovery in case of abrupt displacement from their burrows during the day, and is inherited with genetic differences among populations from differently oriented shores. Moreover, other orientation mechanisms are used in addition to or instead of the sun compass to maintain zonation. A learning capability has also been shown. which can modify the orientation after individual experience. Rhythmic components in the orientational responses have been demonstrated. which would be related to daily migrations in nature. In the laboratory under constant conditions. sandhoppers show rhythmic changes in their responses to simple \'isual stimuli and differences have been shown among populations from Mediterranean and Atlantic shores. A study is also presented aimed at answering the still open question of whether and how sandhoppers are oriented in nature during the foraging movements. which they perform at night under suitable meteorological conditions. Spontaneously active sandhoppers were captured with pit-fall cross traps positioned on the eulittoral and shortly after tested in orientation arenas. Changes in sandhopper orientation resulted during the night. which were related to their zonation and activity. The orientational cues were \isual. to both sky and landscape factors. On the whole. the results throw new light on the ecological meaning of sandhoppers orientation and the conditions in which learning of direction finding may occur in nature. Ke)· words: Talitrus saltatorethoccology. orientation. rhythms. genetics. RESUMEN Talitrus saltator crustáceo anfipodo cuya residencia habitual son las playas de arena. ha sido por largo tiempo objeto de cstudios de orientación debido a que se orienta pcrpendicularmente a la orilla del mar en respuesta a factores de estrés tales como desecación del ambiente o inmersi6n por efccto del oleaje. Durante cl dia la orientaci6n solar le perrnite efectuar la recuperación zonal en cl caso de un repentino traslado del área habitual de cntcrramiento. Este mecanismo de orientaci6n se hereda y prescnta diferencias geneticas entre poblaciones provenientes de playas con distintas orientaciones. Se presentan ademásotros mccanismos de orientaci6n en adici6n o en vez de la orientaci6n solar para mantener la zonaci6n. Tambien se ha cornprohado una capacidad de aprendizaje que puede modificar la orientaci6n como consecuencia de la experiencia individual. SL· ha dcmonsrrado la existencia de componentes ritrnicos en las respucstas de orientaci6n que podrian relacionarse con las migracioncs diarias ocurrentes en la playa. Bajo condiciones constantes de laboratorio, Ios anfipodos mucstran cambios rítmicos en sus rcspuestas a estimulos visuales simples. habiendose demostrado diferencias entre poblaciones costeras del Mediterráneo ) del Atlántico. El presente es un estudio que aspira a dar una respuesta a la pregunta aún abierta sobre la posihilidad y modo de orientaci6n de Ios anfipodos durante su busqueda de alimento. actividad que efectuan de noche cuando las condicioncs meteornl6gicas son apropiadas. Se capturaron anfipodos con trampas ubicadas a ras de la arena para luego ser cstudiados en cámaras de orientaci6n. Los camhios de orientaci6n que Ios talitridos manifestaron durante la noche se relaciona- ron con la zonaci6n y el estado de actividad. Las referencias para la orientación fueron visuales. cielo y paisaje. En sintesis. Ios resultados entregan nuevos antecedentes sobre el significado ecol6gico de la orientaci6n de Ios anfipodos talitridos y de las condiciones en las cuales el aprendiLaje para recuperar una dirccci6n podrfa darse en la naturaleza. Palabras clave: Talitrussaltatoretoccologia, orientaci6n. ritmos. genetica. INTRODUCTION stage and juveniles are released from the brood-pouch of the female at an adult-like The sandhopper Ta!itrus saltator (Montagu) autonomous stage. Their typical habitat is the is a sandy-beach dweller very common on eulittoral above the high tide marks, with Mediterranean and European Atlantic coasts. slight differences in the zonation of activity Its life style is roughly the same during its between adults and juveniles, being juveniles entire life cycle; it develops without larval more active at water's edge. The research (Received I December 1994; accepted 28 July 1996) 554 SCAPINI ET AL. done on this spec1es has revealed many directional light, substrate slope, earth's interesting points that make it a case-study, magnetic field) or to two cues in opposition. particularly for orientation and the genetic Tests have been repeated at different times of determination of orientation. We shall here day. present a brief review of this research con- The genetics of sun orientation has been sidering the many aspects of eco-ethology of studied by testing the inexperienced proge- the species, highlighting new findings that nies of different populations, reared in the await further clarification, namely the tim- laboratory under artificial light conditions; ing and plasticity of orientation behaviour. the learning capability by comparing the A field study and laboratory tests were orientation of individuals from the same performed in order to investigate how and to population which had had different ex- what extent the behaviour of sandhoppers periences (in the rearing containers or in is plastic, i.e. susceptible to both short term previous tests). modifications by changing external condi- The genetic structure of natural popu- tions and/or internal factors, and to long Iations has been analyzed with isoenzyme term ones as far as endogenous inheritable electrophoresis and a dendrogram has been information is concerned. This latter would calculated based on UPGMA clustering of be the basis of populations' adaptation and genetic distance coefficients (Nei). diversity. Spontaneous activity and migration in nature, its timing and zonation have been studied from the captures of sandhoppers in MATERIAL AND METHODS cross pit-fall traps positioned along a transect in the eulittoral zone of the beach. The traps The experiments reported in the paper have have been controlled hourly throughout a been conducted on talitrid amphipods of the 24 hour cycle, or every 2 hours for a period species Talitms sa/rotor (Montagu) from of 72 consecutive hours and repeating the Mediterranean (Italy, Fig. I) and eastern survey each month for one year. Climatic Atlantic (France, Wales) populations. To test and microclimatic conditions have been the orientation of sandhoppers the follow- registered simultaneously. ing methods have been used: I) releases in a Statistical analysis of the circular distribu- glass bowl, 40 cm in diameter, according to tions has been performed according to Pardi & Papi (I 953), in which the sandhop- Batschelet ( 1981 ). Linear simple and multiple per directional choises have been photo- regression analyses have been applied to the graphically recorded from underneath at data (Statgraphics 5.0, 1991 STSC Inc.). given intervals of time, when they hopped and crawled in the bowl; 2) releases from the centre of a circular arena, 40 cm in diameter, RESULTS and captures with pit-fall traps positioned at its rim, underlying 5° each. This method was Orientation in sandhoppers first introduced by Hartwick ( 1976) and modified by Scapini & Pardi ( 1979) to test When a sandhopper is transferred from its sandhoppers singly; 3) releases on sand and burrow to arid conditions, it runs or hops recording of the path of each sandhopper by seawards, while, when immersed in seawater using the marks of the hops. or released on a water saturated substrate, Orientation tests were performed on the it crawls or runs landwards (Scapini 1979). beach 1) with all cues available, 2) by exclud- This behaviour has been defined as zonal ing single cues (landscape vision, beach recovery. Sandhoppers have at their disposal slope) and 3) by putting cues in opposition, an abundance of mechanisms to maintain after transferring sandhoppers to a differently zonation by orienting along the sea-land axis, oriented shore. Laboratory experiments have as reviewed by Pardi & Scapini (1987). The been carried out in controlled conditions, in importance of astronomical cues, such as the order to test orientation to a single cue (sun, sun and moon azimuths and the pattern of black boundary at the horizon, artificial polarization of the light of the sky, were first ECO-ETHOLOGY OF SANDHOPPERS 555 Fig. 1: Localities on the Italian coasts where the Tulitrus saltator populations mentioned in the following figures were caught. SRO-San Rossore (Pisa); CAS-Castiglione della Pescaia, BAL-Bocca t.l"Albegna, GIA-Giannella, FEN-Feniglia, BUR-Burano (Grosseto); NET-Nettuno (Latina); BRU- Brussa (Venezia); GOR-Goro (Ferrara); TFA-Torre Fantine, LES-Lesina, ROD-Rodi Garganico, SIP- Siponto (Foggia). At SRO and FEN. two or three points were sampled. Localidades
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