Spedicato, M.T.; Lembo, G.; Marmulla, G. (eds.) 133 Aquatic telemetry: advances and applications. Proceedings of the Fifth Conference on Fish Telemetry held in Europe. Ustica, Italy, 9-13 June 2003. Rome, FAO/COISPA. 2005. 295p. Movements of juvenile and adult spider crab (Maja squinado) in the Ría da Coruña (N-W Spain)

C. Bernárdez*, E.González-Gurriarán, B. García-Calvo, A. Corgos, & J. Freire

Dpto. Bioloxía Animal, B. Vexetal e Ecoloxía. Universidade da Coruña. Campus da Zapateira, s/n. E-15071 A Coruña (Spain). *Corresponding Author, e-mail: [email protected]

Key words: Maja squinado, movements, maturity, ultrasonic telemetry, Ría da Coruña, North-West Spain.

Abstract

In July 2002, 12 juvenile and 12 adult (recently terminal moulted) spider crabs were tagged using ultrasonic pingers (Sonotronics, USA) in the Ría da Coruña (N-W Spain). Both adults and juveniles were released in the same sandy shallow bottoms (5-10 m) in which they were caught. This area, as known from previous studies, is a typical juvenile area where spider crabs carry out their terminal moult before the autumn migration to deep bot- toms. The tracking was carried out daily for a period of two weeks, although not all the crabs were located every day. Juvenile crabs were found to move small distances, performing non-directional movements near the coast, in protected areas up to 10 m deep. Two of the adult crabs performed small-scale movements in the outer (more exposed) part of the estuary, at depths up to 20 m. The rest of the adult crabs remained in the juvenile area, per- forming the same non-directional movements as the juveniles. Four male adult crabs were recaptured by com- mercial fisheries in the following months, three of them at more than 40 m deep, in the central channel of the estuary, probably while performing their seasonal descent migration. A female carrying eggs was recaptured in December near the coast.

Introduction in this period, leading to migratory movements to deeper waters (Latrouite and Le Foll, 1989; The spider crab, Maja squinado (Decapoda, González-Gurrirán and Freire, 1994; Hines et al., Majidae) (recently splitted into Maja squinado 1995). The adults inhabit deep areas where they are (Herbst, 1788), inhabiting Mediterranean coasts, the target of a tanglenet fishery (González- and Maja brachydactyla Balss, 1922, in the Gurriarán et al., 1993; González-Gurriarán and Atlantic coasts of Europe (Neumann, 1996)), is a Freire, 1994). Movements and habitat use of this species that lives at depths ranging from subtidal species is highly important for the fishery, given level to more than 150 m (González-Gurriarán that catches are directly related to the activity et.al., unpublished data). There is strong evidence rhythms and migrations. Although many studies that spatial segregation exists between juveniles have been carried out analyzing migration move- and adults; these habitat differences are variable ments using mark-recapture methods (Camus, according to the season (Corgos et al., 2002). In the 1983; Kergariou, 1976; Le Foll, 1993; González- Ría da Coruña (N-W Spain), juveniles (carapace Gurriarán et al., 2002), little information is avail- lenght < 120-140 mm) inhabit predominantly shal- able on the animal’s activity in the coastal shallow low waters characterized by mixed hard-and-soft areas in the months prior to migration (González- bottoms. One or two months after the terminal Gurriarán and Freire, 1994). moult that takes place in their second year, through The objective of this study was to describe patterns which they attain maturity, females start their of movement and habitat use in the summer period gonad maturation (González-Gurriarán et al., when shallow areas, determined by previous stud- 1998). Some behavioural changes are also defined ies, are shared by juveniles and young adults 134

(recently moulted to maturity). Local movements shallow sandy bottom area in the inner part of the between habitats or areas within a habitat as well as Ría da Coruña (43º 21’ N, 8º 22’ W), between the 8th aggregation patterns are aspects of interest as yet to and the 10th July 2002. Twelve juvenile (with cara- be established. pace length between 84 and 135.5 mm) and twelve The study used ultrasonic telemetry, previously adult (recently moulted to maturity; carapace shown to be an efficient tool to obtain accurate data length between 112 and 173 mm) spider crabs were on crab small-scale movements and migrations tagged using Sonotronics IBT-96-2 and CHP-87-S (Maynard and Webber, 1987; Wolcott and Wolcott, pingers, respectively. The first has a weight of 2.5 2002). It has also been successfully used in the spi- g in water and dimensions of 28 by 9.5 mm. The der crab (Freire and González-Gurriarán, 1998; battery lasts for 60 days, and the operating fre- González-Gurriarán and Freire, 1994; González- quency is 69 or 76.8 kHz. Under ideal oceano- Gurriarán et al., 2002; Hines et al., 1995). graphic conditions the pingers can be detected from distances up to 500 m (Sonotronics’ technical specifications). The pingers used for the tagging of Materials and methods adult crabs weighted 8 g in water, and their dimen- sions were 67 by 18 mm. The battery lasts for 7 In July 2002, 24 spider crabs were tagged using months and the transmission frequency is 75 kHz. ultrasonic pingers in the Ría da Coruña (NW Spain) They can be detected from distances up to 3000 m (Fig. 1). The tagging took place in Bastiagueiro, a in ideal conditions.

Fig. 1 – Ría da Coruña (NW Spain), showing tagging area (*) and oceanographic data collection stations (4, 3b and 3c). 135

The spider crabs were captured using pots. Algae Some recaptures were obtained from commercial and other epiphytes were cleaned off the dorsal fisheries in the following months. Data such as part of the carapace by scrubbing them with a fishing date, position and depth were taken. In brush. Then the area was dried using alcohol and the only case of a recaptured female, dissection acetone in a cloth. Tags were then attached to the was performed to determine gonad and egg mat- clean, dry carapace on the dorsal side, using quick uration stage and number of sperm masses epoxy. Adult crabs’ tags, which are bigger, could (González-Gurriarán et al., 1998). be labelled by writing UDC´s (Universidade da Oceanographic data, recorded monthly, were pro- Coruña) telephone number and the word “reward” vided by the Instituto Español de Oceanografía on its surface. Smaller tags used for juveniles (Centro Costero de A Coruña). These included allowed no writing on their surface. Thus, the temperature and salinity in three sampling sta- labelled part of a T-bar anchor tag was also tions in the tagging area and outer part of the attached to the carapace, using the same quick estuary, from 20 to 40 m depth. Two data loggers, epoxy. The tagging process was as quick as possi- that recorded temperature data every two hours, ble (<20 minutes), in order to avoid long air expo- were placed in the tagging area, at fixed depths of sure. 5 and 10 m. A Sonotronics USR-5W receiver with a Sonotronics DH-2 directional hydrophone was used to locate the tagged crabs on a daily basis Results (although some of the crabs were not located every day due to unfavourable weather and From a total of 24 tagged crabs, four were never oceanographic conditions, crab loss, lack of time, located (lost after release) and one crab was locat- etc…), using a small boat equipped with echo- ed only once. These five crabs were all adults. sounder and GPS. The telemetry experiment Mean number of position fixes per crab was 10 started the 8th July 2002 and ended the 24th July (range: 4-12). During the tracking period, both 2002. juvenile and adult spider crabs performed small- Distance between consecutive locations and scale movements within the inner part of the estu- movement direction in relation to the north was ary (Fig. 2). The mean distance between locations estimated from GPS positions of located crabs. was 187 m for juveniles and 179 m for adults The distance between locations was assumed to (range: 94-517 and 114-250, respectively) and the be the minimum distance travelled by the animal mean speed 154 mday-1 for juveniles and 159 between observations. Speed (mday-1) was calcu- mday-1 for adults (range: 58-272 and 100-250, lated as distance between locations in consecu- respectively). These movements were carried out tive days. Distance between locations, speed and in shallow sandy bottoms. Mean depth was 6.5 m mean depth was calculated omitting the release for juveniles and 9.5 m for adults. observation. The statistic r was computed, which Figure 3 shows the distance between crabs during is equal to the mean vector length and represents the experiment. Crabs moved up to 187 mday-1, a measurement of concentration of the different and average distance between crabs was 80 m movement angles. The length will range from 0 (range: 17-106 m). to 1; larger number indicates that the observa- One juvenile crab showed significant directionality tions are clustered more closely around the mean (SW) in its movements (r=0.57, p=0.03). Two than lower numbers. The Rayleigh test was per- crabs that were lost for more than one week during formed to determine if the movement of each ani- the tracking were excluded from the directionality mal presented a significant directional orienta- analysis. The rest of the crabs did not show signif- tion. The test calculates the probability of the null icant directionality in their movements (tests of hypothesis that the data are distributed in a uni- Rayleigh, p>0.05; Figs. 2 and 4); r statistic ranged form manner (Batschelet, 1981). between 0.08 and 0.4. 136

Fig. 2 – Examples of movements of two juvenile crabs (#225 and #358).

In late August, one month after the end of the track- was recaptured in late December 2002 in coastal ing, three adult males were recaptured by commer- shallow waters. cial fisheries. One of them was recaptured in the Oceanographic data (Fig. 5, a and b) show a stable inner part of the estuary, at a depth of 17 m. The pattern of both temperature and salinity during the other two were recaptured in the central channel, late spring-summer period (May to September). outside the estuary, at a depth of 50 m. Another Mean temperatures show an oscillation of 2 ºC and male was recaptured outside the estuary in May salinity initiates a slow decrease in September, 2003, at a depth of 45 m. A female carrying eggs related to the first autumn rains.

Fig. 3 – Average distance (m) between crabs (both juve- Fig. 4 – Average angles of movements in juveniles and nile (solid line) and adult (dotted line) from release to adults. Vector length is proportional to directionality of the end of the experiment. Thinner lines show the con- movement (statistic r; radius of the circumference=1). fidence interval. Triangles represent release dates. 137

muscle and hepato-pancreas condition. The first stage is characterized by movements that follow the same pattern as those of juveniles. We found, for both juveniles and adults, that the average dis- tance between animals was smaller than the aver- age distance between consecutive locations. Taking into account the potential movement of the crabs (up to 187 mday-1), the average distance between crabs during tracking (80 m) would point to an aggregative pattern as suggested in previous studies (Corgos et al., 2002). An increase in movement scale and directionality was found in the second phase, prior to migration (González-Gurriarán and Freire, 1994). Right before the descent autumn migration, adults move progressively to the outer part of the shallow area, leaving it coordinately in September (males) and October (females) (Corgos et al., 2002). Our results show recaptures in the central channel out- side the estuary, at a depth of 50 m, suggesting the beginning of movements to deeper areas. Seasonal migrations have been described in decapods such as blue crab (Skajaa et al., 1998; Wolcott and Wolcott, 2002), snow crab (Maynard and Webber, 1987), king crab (Fotheringham, 1975) and swim- ming crabs (Venema and Creutzberg, 1973). Fig. 5 – a, Bottom temperature at stations 4, 3b and 3c The beginning of the descent migration, which tends (20, 35 and 40 m depth, respectively), and temperature at 5 and 10 m in the tagging area. b, Bottom salinity at to occur in late summer or autumn, has been found stations 4, 3b and 3c. to be strongly related to oceanographic parameters together with ontogenetic aspects (Venema and Creutzberg, 1973; Latrouite and Le Foll, 1989; González-Gurriarán and Freire, 1994; Tankersley et Discussion al., 1998; Watson et al., 1999; González-Gurriarán et al., 2002; Wolcott and Wolcott, 2002). The period In the Ría da Coruña, juvenile spider crabs show a in which our tracking was performed corresponds to clear aggregative pattern in shallow areas, proba- the first stationary phase described by González- bly to avoid predation risk (Corgos et al., 2002). Gurriarán and Freire (1994). Mating might take During late summer, both juveniles and recently place in deep areas (González-Gurriarán et al., moulted to maturity adults carry out small-scale, 1998) where a lek mating system has been suggest- non-directional movements. This kind of short, ed by Corgos et al. (2002). random movement in shallow areas has also been The possibility of fertilizing eggs with sperm observed in other crabs, such as Cancer pagurus from previous matings would give the females (Skajaa et al., 1998), and in Homarus americanus the opportunity to spawn two o three times with- (Watson et al., 1999). out remating (González-Gurriarán et al., 1998). González-Gurriarán and Freire (1994), describe a The hypothesis of a late winter-early spring two-phase movement pattern for these young ascent migration towards shallow coastal areas, adults, while gradually increasing their gonad, where higher temperatures would favour egg 138

development, was suggested by González- González-Gurriarán, E., Fernández, L., Freire, J., Gurriarán et al. (2002). Spawning migrations Muiño, R. & Parapar, J. 1993. Reproduction of have also been described for Callinectes sapidus the spider crab Maja squinado (Brachyura: (Carr et al., 2002; Tankersley et al., 1998; Majidae) in the southern Galician coast (NW Wolcott and Wolcott, 2002). Male spider crab Spain). ICES, Shell. Comm. C.M. 1993/K:19, 15 pp. recaptures still occur in spring in deep areas, thus González-Gurriarán, E., Fernández, L., Freire, J. & supporting the hypothesis of a just-female ascent Muiño, R. 1998. Mating and role of seminal migration. Breeding female recaptures in shallow receptacles in the reproductive biology of the spi- waters occur from December on, as it is the case der crab Maja squinado (Decapoda, Majidae). J. in our study, supposedly after performing both Exp. Mar. Biol. Ecol., 220: 269-285. migrations. González-Gurrirán, E. & Freire, J. 1994. Movement patterns and habitat utilization in the spider crab Maja squinado (Herbst) (Decapoda, Majidae) Acknowledgements measured by ultrasonic telemetry. J. Exp. Mar. Biol. Ecol., 184: 269-291. Oceanographic data were provided by I.E.O. González-Gurriarán, E., Freire, J. & Bernárdez, C. Maja through Dr. M. Varela. This study was funded by 2002. Migratory patterns in the spider crab squinado using electronic tags and telemetry. J. the Spanish Ministerio de Ciencia y Tecnología, Crust. Biol., 22 (1): 91-97. Dirección General de Investigación (Project Hines, A.H., Wolcott, T.G., González-Gurriarán, E., REN2000-0446/MAR). González-Escalante, J.L. & Freire, J. 1995. Movement patterns and migrations in crabs: telemetry of juvenile and adult behaviour in References Callinectes sapidus and Maja squinado. J. Mar. Biol. Ass. UK, 75: 27-42. Batschelet, E. 1981. Circular statistics in Biology. Kergariou, G. 1976. Premiers résultats obtenus par New York, Academic Press, 371 pp. marquage de l’araignée de mer, Maia squinado, Camus, P. 1983. Résultats d’une opération de marquage déplacements, mortalité par pêche. ICES, Shell. d’araignée de mer (Maja squinado, Herbst) adult en Benth. Comm., C.M. 1976/K:14, 6 pp. baie d’Audierne (Bretagne Sud). ICES, Shell. Latrouite, D. & Le Foll, D. 1989. Donnees sur les Comm. C.M. 1983/K:29, 11 pp. migrations des crabes torteau Cancer pagurus et Carr, S. D., Tankersley, R. A., Hench, J. L., Forward, les arignees de mer Maja squinado. Océanis R. B. Jr. & Luettich, R. A. Jr. 2002. Field obser- 15:133-142. vations of ebb-transport of the blue crab Le Foll, D. 1993. Biologie et exploitation de l’araignée Callinectes sapidus near a barrier island inlet de mer Maja squinado Herbst en Manche Ouest. using ultrasonic telemetry. Open Sciences 2002, Université de Bretagne Occidentale. IFREMER, RI Honolulu, Hawaii. DRV 93-030. Thése de Doctorat, 524 pp. Corgos A., Bernárdez, C., Verísimo P. & Freire, J. Maynard, D.R. & Webber, D.M. 1987. Monitoring 2002. Population dynamics of Maja squinado in the movements of snow crab (Chionoecetes the Ría de A Coruña (Galicia, NW Spain), using opilio) with ultrasonic telemetry. Proceedings of mark-recapture experiments. 8th Colloquium Oceans, 87: 962-966. Crustacea Decapoda Mediterranea, 2-6 September Neumann, V. 1996. Comparative investigations of the 2002, Corfu, Greece. systematics and taxonomy of European Maja Fotheringham, N. 1975. Structure of seasonal migra- species (Decapoda, Brachyura, Majidae). tions of the littoral hermit crab Clibanarius vitta- Crustaceana, 69: 821-852. tus. J. Exp. Mar. Biol. Ecol., 18: 47-53. Skajaa, K., Fernoe, A., Loekkeborg, S. & Haugland, Freire, J. & González-Gurriarán, E. 1998. New E.K. 1998. Basic movement pattern and chemo- approaches to the behaviour ecology of decapod oriented search towards baited pots in edible crab crustaceans using telemetry and electronic tags. (Cancer pagurus L.). Hydrobiologia, 371-372: Hydrobiologia, 371/372: 123-132. 143-153. 139

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