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Cephalopod movement occurs during all phases of the life history. Environmental parameters have a large influence on movement at the various life cycle stages. Chemical tagging of paralarvae through maternal transfer may prove a viable technique for tracking this little understood life stage.
Cephalopod movement occurs during all phases of the life history. Environmental parameters have a large influence on movement at the various life cycle stages. Chemical tagging of paralarvae through maternal transfer may prove a viable technique for tracking this little understood life stage.
Cephalopod movement occurs during all phases of the life history. Environmental parameters have a large influence on movement at the various life cycle stages. Chemical tagging of paralarvae through maternal transfer may prove a viable technique for tracking this little understood life stage.
migration patterns Jayson M. Semmens Gretta T. Pecl Bronwyn M. Gillanders Claire M. Waluda Elizabeth K. Shea Didier Jouffre Taro Ichii Karsten Zumholz Oleg N. Katugin Stephen C. Leporati Paul W. Shaw Received: 2 January 2007 / Accepted: 1 February 2007 / Published online: 16 March 2007 Springer Science+Business Media B.V. 2007 Abstract Cephalopod movement occurs during all phases of the life history, with the abundance and location of cephalopod populations strongly inuenced by the prevalence and scale of their movements. Environmental parameters, such as sea temperature and oceanographic processes, have a large inuence on movement at the various life cycle stages, particularly those of oceanic squid. Tag recapture studies are the most common way of directly examining cephalopod movement, particularly in species which are heavily shed. Electronic tags, however, are being more commonly used to track cephalopods, providing detailed small- and large-scale move- ment information. Chemical tagging of paralarvae through maternal transfer may prove to be a viable technique for tracking this little under- stood cephalopod life stage, as large numbers of individuals could be tagged at once. Numerous indirect methods can also be used to examine cephalopod movement, such as chemical analyses of the elemental and/or isotopic signatures of J. M. Semmens (&) G. T. Pecl S. C. Leporati Marine Research Laboratories, Tasmanian Aquaculture and Fisheries Institute, University of Tasmania, Private Bag 49, Hobart, TAS 7001, Australia e-mail: Jayson.semmens@utas.edu.au B. M. Gillanders Southern Seas Ecology Laboratories, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, SA 5005, Australia C. M. Waluda Biological Sciences Division, British Antarctic Survey, High Cross, Madingley Road, Cambridge CB3 0ET, UK E. K. Shea Delaware Museum of Natural History, P.O. Box 3937, Wilmington , DE 19807, USA D. Jouffre Unite OSIRIS, IRD, BP 1386, Dakar, Senegal T. Ichii National Research Institute of Far Seas Fisheries, 2-12-4 Fukuura, Kanazawa-ward, Yokohama-City 236-8648, Japan K. Zumholz Leibniz-Institute of Marine Sciences, IFM- GEOMAR, Du sternbrooker Weg 20, D-24105 Kiel, Germany O. N. Katugin Pacic Fisheries Research Centre, 4 Shevchenko Alley, Vladivostok 690950, Russia P. W. Shaw School of Biological Sciences, Royal Holloway University of London, Egham TW20 0EX, UK 1 3 Rev Fish Biol Fisheries (2007) 17:401423 DOI 10.1007/s11160-007-9048-8 cephalopod hard parts, with growing interest in utilising these techniques for elucidating migra- tion pathways, as is commonly done for sh. Geographic differences in parasite fauna have also been used to indirectly provide movement information, however, explicit movement studies require detailed information on parasite-host specicity and parasite geographic distribution, which is yet to be determined for cephalopods. Molecular genetics offers a powerful approach to estimating realised effective migration rates among populations, and continuing developments in markers and analytical techniques hold the promise of more detailed identication of mi- grants. To date genetic studies indicate that migration in squids is extensive but can be blocked by major oceanographic features, and in cuttlesh and octopus migration is more locally restricted than predictions from life history parameters would suggest. Satellite data showing the location of shing lights have been increas- ingly used to examine the movement of squid shing vessels, as a proxy for monitoring the movement of the squid populations themselves, allowing for the remote monitoring of oceanic species. Keywords Cephalopods Movement Migration Environmental variability Introduction Movement of cephalopod populations takes place during all phases of the life history, from the passive drifting of egg masses (ODor and Balch 1985) and paralarvae to the daily vertical migra- tions of many oceanic species (Roper and Young 1975) and large-scale migrations (thousands of kilometres) of adult animals from feeding to spawning grounds (ODor 1992). Cephalopods are short lived, and exhibit a high level of plasticity in their life history strategies where changes in environmental conditions have a large part to play in inuencing population size, abun- dance and distribution. One of the major factors inuencing the active movement of cephalopods is likely to be the availability and movement of food supplies (Rodhouse and Nigmatullin 1996), and many cephalopod populations have evolved their migratory strategies to maximise high feed- ing rates. Movement patterns can inuence the response of cephalopods to environmental change, and the abundance and location of cephalopod populations and associated sheries is strongly inuenced by the prevalence and scale of their movements (Boyle and Boletzky 1996). Those populations which undertake the largest migrations are prone to the largest uctuations in population abundance (ODor 1992; Dawe et al. 2000). Particularly in ommastrephid squid, annual uctuations in catch can vary between 2 and 5 orders of magnitude for an individual species (FAO 2000). This review seeks to clarify our understanding of cephalopod movement and migration and determine where future research investment will be most benecial in terms of increasing our understanding and providing effective tools for resolving movement patterns. We appraise meth- ods for resolving movement and migration pat- terns of cephalopod species and examine the strong relationship between movement in cepha- lopods and environmental parameters. Methods for resolving movement/migrations patterns/processes The various methods used to examine cephalopod movement/migration will be explored in this section, using relevant examples, along with potential methods currently being developed or to date only used for other groups, such as sh. These methods are divided into ve broad cate- gories: tag recapture methods, which involve marking and releasing animals; electronic tags; chemical tags; natural tags (which includes para- sites, molecular genetics and elemental signa- tures) and tracking shing eets. Tag-recapture External tags External tags which are attached to the study animal and identify individuals or groups of 402 Rev Fish Biol Fisheries (2007) 17:401423 1 3 animals, along with instructions for reporting, are the most widely used technique to examine the movement patterns of cephalopods (see Nagasa- wa et al. 1993 for an excellent review and description of the types of external tags com- monly used), and other marine organisms in general. This wide-spread use is due to their relatively low cost and ease of application, which allows for large number of individuals to be tagged. There are several potential disadvantages to using these tags, however, including damage to the tagged animal from the tag rubbing against the muscle/skin, (e.g., Kanamaru and Yamashita 1966; Sauer et al. 2000), tag loss (e.g., Kanamaru and Yamashita 1966; Tsuchiya et al. 1986, in Nagasawa et al. 1993), and the relatively large tag and applicator meaning that in general juveniles can not be tagged. Recapture rates are variable and inuenced by factors such as level and extent of shing (e.g., Mokrin 1988; Markaida et al. 2005), tag colour (e.g., Nagasawa et al. 1993; Sauer et al. 2000), tag type (e.g., Mokrin 1993; Nagasawa et al. 1993) and tag placement (e.g., Nagasawa et al. 1993; Sauer et al. 2000). Report- ing rates for external tags can also be highly variable for a wide variety of reasons such as, sher apathy, resentment of research/manage- ment and insufcient rewards. Extensive studies of the annual spawning migrations of the ommastrephid squid, Todarodes pacicus, one of the largest single species ceph- alopod sheries in the world, has seen over 1 million squid tagged in the Sea of Japan and the North Pacic with various types of external tags (see Nagasawa et al. 1993; Kidokoro, pers. comm.), most commonly T-bar (or anchor) tags with a pentagonal pennant (Shevtsov 1973, 1978; Skalkin 1973; Mokrin 1988, 1993, pers. comm.; Nagasawa et al. 1993). Such large numbers are required as low recapture rates are common for ommastrephids shed in large offshore shing zones (e.g., 0% for Illex illecebrosus in New- foundland, Hurley and Dawe 1981; 2.5% for T. pacicus, Nagasawa et al. 1993), whereas recap- ture rates are generally much higher in inshore waters (e.g., 1932% I. illecebrosus, Hurley and Dawe 1981; 8% Dosidicus gigas in the Gulf of California, Markaida et al. 2005; 10% Sepioteu- this australis in Tasmania, Australia, Pecl et al. 2006). Interestingly, several studies suggest that recaptures rates may be size and/or sex dependent (e.g., D. gigas, Markaida et al. 2005; S. australis, Pecl unpublished data). As well as high release numbers, spatial cov- erage of the full extent of the shery is important for tag-recapture studies if the migration routes of the various species/populations are to be correctly dened (e.g., T. pacicus, Nakamura and Mori 1998; Mori and Nakamura 2001). This spatial coverage is only possible if the shery operates throughout the migratory route, for example the movement of D. gigas into and out of the Gulf of California is unknown due to the much lower level of shing effort in the Southern Gulf and Pacic Ocean (Markaida et al. 2005). The Japanese T. pacicus tag-recapture pro- grams demonstrate a major advantage of external tags, with their relatively low cost and ease of application making them well suited to creating long-term data sets to examine temporal trends in migration patterns. Analysis of long-term tag- recapture data for autumn spawning T. pacicus, where 20,00030,000 individuals have been tagged annually since the 1950s (Nagasawa et al. 1993; Kidokoro pers. comm.) suggest that within the Sea of Japan, the southward migratory route to the spawning area changes in response to climate shifts caused by variations in the southward meandering of the Subarctic Front (Kidokoro et al. 2004). Unlike ommastrephid squids, coastal cephalo- pod species such as octopods, loliginids and sepi- oids, generally undertake relatively small scale (tens to hundreds of kilometres) migrations, between spawning and feeding grounds (e.g., Loligo forbesi, Holme 1974; Waluda and Pierce 1998, Sepia ofcinalis, Wang et al. 2003; Royer et al. 2006, Octopus vulgaris, Sobrino et al. 2002). Long-term tag-recapture studies of the loliginid squids Uroteuthis edulis and Heterololigo bleekeri in the Sea of Japan between 19741985 and 1960 1989 respectively (see Nagasawa et al. 1993), showed that despite favouring open coastal waters, these squid do not move beyond the continental shelf (Natsukari and Tashiro 1991). Tag-recapture studies of Loligo reynaudii on the southeast coast of South Africa showed that loliginids can move relatively large distances within a spawning area, Rev Fish Biol Fisheries (2007) 17:401423 403 1 3 with an average movement of around three km/day and a maximum of almost 15 km/day during their spawning period of a month or more (Sauer et al. 2000), which may be in response to changing environmental conditions (see Roberts and Sauer 1994) and food availability. External tagging experiments on octopuses present additional challenges to those for squid and cuttlesh (see examples in Nagasawa et al. 1993; Bakhayokho 1986), as their dexterity and strength allows them to easily remove external tags (e.g., O. vulgaris, FAO 1979; Tsuchiya et al. 1986, in Nagasawa et al. 1993). Despite this, Domain et al. (2000, 2002) successfully tagged 4245 O. vulgaris on the Senegalese shelf using Petersen discs attached to the base of an arm, including individuals as small as 200 g (Domain et al. 2002), with 1,019 recaptured (24%) by the commercial line-shing eet. The percentage of recaptures varied from 0% to 49% depending on the tagging location along the shelf, with low recapture rates where few professional shermen were present (Domain et al. 2002), again high- lighting the link between the success of tag- recapture programs and the spatial coverage of the shery (Markaida et al. 2005). Despite the success of the tagging method, all recaptured individuals were captured in the area they were released (Domain et al. 2002), suggesting that O. vulgaris may not undertake migrations on the Senegalese coast, unlike the seasonal inshore- offshore migrations made by this species in the Mediterranean Sea (Mangold 1983), however, octopus were only at large for 100 days maximum and all stages of the life cycle were not examined. External tags attached to the base of the arms have proved effective for tagging Enteroctopus doeini in Japan (see Nagasawa et al. 1993) and North America (e.g., Hartwick et al. 1984a, b; Robinson and Hartwick 1986; Hartwick et al. 1988). Tag-recapture studies and sheries sam- pling demonstrated that E. doeini undertakes two seasonal inshore-offshore migrations per year in the northern Sea of Japan (Kanamura and Yamashita 1966, 1969, in Nagasawa et al. 1993; Kanamura and Yamashita 1967, in Mottet (1975)). A similar pattern was not found for E. doeini in British Columbia, Canada (Hartwick et al. 1984b, 1988), and electronic or natural tagging techniques may be more applicable for elucidating an inshore/offshore migratory pattern (see below). Tattooing and branding Tattooing and branding techniques have been trialled on several cephalopods, predominately octopus, but with limited success (see Nagasawa et al. 1993). Although these approaches over- come the problem of tag removal and allow for juveniles to be tagged, they are often technically difcult to undertake and can cause severe stress and damage to the animal. However, Tsuchiya et al. (1986, in Nagasawa et al. 1993) compared various external tagging techniques with tattooing for O. vulgaris, and found tattooing dye spots in the mantle musculature to be the most successful, in terms of tags retention and minimising damage/ stress to the animal. Cuttlesh (Sepia latimanus) have also been tagged using a similar technique, namely injecting red latex between the skin and cuttlebone. How- ever there were few recaptures (two from 450 individuals tagged), marks lasted only a short time (2 months) and wounds led to mortality (Oka 1990b, in Nagasawa et al. 1993). Forsythe (pers. comm.), however, successfully tagged juve- nile cultured Sepia ofcinalis (approximately 10 cm ML) using uorescent injectable micro- spheres, with at least one individual growing through to maturity, with its tag still visible after six months; however these animals were never released from captivity. Similarly, visible implant elastomer (VIE) tags (Northwest Marine Tech- nology, Shaw Island, USA, www.nmt.us), which are uorescent silicon tags injected under the skin, have been used successfully to tag and recapture Sepioteuthis sepioidea juveniles as small as 19 mm ML in the eld (Replinger and Wood in press), and may have potential for tag-recapture of paralarval cephalopods, as small sh have been tagged and recaptured using VIE tags (e.g., 8 mm TL reef sh, Frederick 1997). Electronic tags These are generally electronic devices that are implanted or attached to the animal and have 404 Rev Fish Biol Fisheries (2007) 17:401423 1 3 either a unique code or frequency, which is transmitted to a recording device, or record envi- ronmental information as the animal swims around. There are several different classes of tag (see below). Acoustic transmitters Acoustic telemetry, where an aquatic organism is tagged with a high-frequency transmitter that can be heard by a mobile or xed hydrophone linked to a receiver that provides either a presence/ absence record, a geographical position, or the relative direction of the animal, has been used since the 1950s to study behaviour and physiol- ogy, particularly in sh (Arnold and Dewar 2001; Sibert 2001). Development of this technology has increased rapidly in recent years matching that of electronics and technology in general, but also in response to a trend towards spatial management of sheries worldwide (Sibert 2001). The advan- tage of this type of tag is that individuals do not need to be recaptured to gain movement infor- mation, and unlike tag-recapture studies, you gain much more detailed information rather than just a capture and recapture position. A disadvantage of these tags is that they have typically been quite large, often 20100 mm, however, the technology is getting smaller all the time, with the new generation of tags being signicantly smaller (15 mm and 0.5 g in water, Vemco, Halifax, Canada, www.vemco.com). Cost is also a disad- vantage, as releasing large numbers of acoustic tags is an expensive exercise and the receiving equipment, which also can be relatively expen- sive, is also required. As such, these studies may be done in conjunction with tag-recapture pro- grams (e.g., S. australis, Pecl et al. 2006) with traditional tag-recapture providing minimal data on a large number of animals and acoustic tagging providing much more detailed information on fewer individuals. A further disadvantage comes where moored automatic tracking systems are not available or practical, meaning tagged animals have to be manually followed in vessels using directional hydrophones which can generally only be done for short periods of time. Three of the most interesting acoustic tagging studies on cephalopods use a Radio-Acoustic Positioning and Telemetry system (RAPT, Vem- co) to triangulate the position of tagged animals and track them in near real-time (see Klimley et al. 2001 for details on RAPT); this system has provided previously unavailable insights into the behaviour of inshore cephalopods. Loligo rey- naudii was tracked in South Africa using RAPT, revealing their mating system to be unexpectedly complex (Sauer et al. 1997). RAPT was used to examine small-scale movement of Sepia apama in Port Lincoln, South Australia, demonstrating that the cuttlesh spent more than 95% of the day resting, which suggests that these are sit and wait predators and make few excursions in pursuit of food (Aitken et al. 2005). Unique vertical move- ments of E. doeini tagged with depth (absolute pressure)-position tags in Northern Japan, which were observed to be out of sequence with the tides, were detected using RAPT (Rigby and Sakurai 2005). Observations using SCUBA re- vealed these octopuses were moving towards a gill net and scaling the net to remove sh, thus producing the unusual vertical movements. Recently, underwater curtains of acoustic listening stations (Vemco VR2 acoustic receivers, see Heupel et al. 2006 for a review of this technology) have been used to examine move- ment of cephalopods (Stark et al. 2005; Pecl et al. 2006). This technology is becoming increasingly popular for examining the movement of a wide variety of sh and shark species (see Heupel et al. 2006). Pecl et al. (2006) assessed the relationship between two S. australis spawning areas on the east coast of Tasmania, Australia separated by approximately 30 km. Although individuals were spawning over several months and travelled 100s of km within the spawning areas during this time, movement between the two areas or out of each area was not detected. Interestingly, these results contrast in some respects with the preliminary external tagging experiment carried out by Molts- chaniwskyj and Pecl (this volume), where 90% of S. australis individuals tagged with T-bar tags on the spawning grounds were later caught within 1 km of the tagging site, suggesting no movement. This appears to highlight one of the main advan- tages acoustic monitoring has over tag-recapture studies, that more detailed movements can be detected in a well designed study i.e., careful Rev Fish Biol Fisheries (2007) 17:401423 405 1 3 placement of receivers, not just the release and capture points. The use of acoustic curtains, may become more commonly used to examine med- ium to large-scale movements of cephalopods, with several countries establishing permanent arrays of these listening stations for tracking a wide range of marine animals, and users needing only to purchase tags (e.g., the Pacic Ocean Shelf Tracking (POST) project, www.post- coml.org). These arrays will, however, only extend as far as the continental shelf and so will be of greatest benet to inshore species, such as loliginids, sepioids and octopods. There have been relatively few acoustic telem- etry studies of oceanic squid, due to the difcul- ties of tracking in this environment, however, Nakamura (1991, 1993), Yatsu et al. (1999) and Yano et al. (2000) actively tracked mature female Ommastrephes bartramii in the North Pacic, D. gigas off Peru and Thysanoteuthis rhombus in Okinawa, Japan respectively, using depth-posi- tion tags. All species showed clear daily patterns of vertical movement, with D. gigas migrating to below 1000 m and O. bartramii and T. rhombus moving deeper during the night, perhaps to avoid predators (Nakamura 1993) or follow prey (Yano et al. 2000), with the vertical movement of T. rhombus matching that of the deep scattering layer. As active-tracking technology develops, these oceanic studies may become more common, with the potential for shing eets to automati- cally upload data from tagged squids, using vessel-mounted receivers. PIT tags Passive Internal Transponder (PIT) tags are elec- tromagnetically coded tags that are injected under the skin of various animals, including many sh species; particularly salmonids (see Prentice et al. 1990). The advantage of these tags is they are relatively small (typically 10 mm and 0.1 g), inex- pensive, and individuals can be identied automat- ically. However, the tag has to pass close by a reader for the animal to be identied, meaning the tagged animal needs to be recaptured and read with a portable tag reader (e.g., Roussel et al. 2000), or in some situations electronic gates can be established, such as in shways (Castro-Santos et al. 1996) or at-bed antennae can be used to monitor small-scale behaviour in streams (e.g., Armstrong et al. 1996). Only one published study has used these tags for cephalopods (Anderson and Babcock 1999). PIT tags were implanted in the base of the arm of eld (in lairs) and captive Octopus tetricus and captive Octopus maorum. All octopus retained the tags for the life of the study (captive: 14 months; eld: 13 weeks), with no signs of damage or deterioration from the tags, and individuals demonstrating normal growth. As such, PIT tagging could be a useful tool for providing long-term recognition of octopuses, with shing vessels and sh processors able to be provided with automatic detectors that record tag IDs as the animals are dropped into holding tanks/bins, eliminating the reporting problems associated with external tags. Recently, Hallprint (South Australia, www.hallprint.com) began pro- ducing PIT tags imbedded within conventional T- bar external tags, allowing for both vessels with/ without auto-readers to record tag numbers, with these tags perhaps suited to the large-scale tagging programs for T. pacicus, to increase the tag reporting rate. These tags and auto-readers have recently been trialled in the southern rock lobster shery in Tasmania, with the concept proving to be worth pursuing (see www.hall- print.com). A limitation of these tags is, however, the need to pass close to the tag reader and therefore aside from the utility mentioned above, they are likely to only be useful in eld situations for small-scale movement. Coded micro-wire tags Coded micro-wire tags (Northwest Marine Tech- nology) are a small length of stainless steel wire (1 0.25 mm), which is magnetised and injected into the animal. Tags are detected using a manual or automatic magnetic detector and an etched identifying number can be read with a micro- scope. The advantage of these tags is that they are relatively cheap (although the auto-readers are relatively expensive), cause minimum harm, and very small animals can be tagged (e.g., Arctic char Salvelinus alpinus 22 mm total length, Champi- gneulle et al. 1987). The disadvantage is the small 406 Rev Fish Biol Fisheries (2007) 17:401423 1 3 tag needs to be removed from the animal for its individual code to be read. These tags have not been used on cephalopods, but may have promise for tagging paralarvae/juveniles, and are widely used on small salmonids in the USA. Archival tags Archival (or data storage) tags record and store environmental data, typically light intensity for estimating geoposition, pressure for estimating depth, and water and body temperature (see Arnold and Dewar 2001 for a review). One major disadvantage of these tags is that they have to be recovered to download the data; they are also quite large (3080 mm) and relatively expensive. However, an advantage is that like acoustic tags they provide shery independent data and can provide a wide variety of ecological data over large distances and time-scales. New types of archival tags have been designed to overcome the recovery issue, with pop-up archival transmitting (PAT) tags (Wildlife computers, Washington, USA, www.wildlifecomputers.com) detaching from the animal at set times and up-loading summaries of their data to satellites once they are on the surface. Communicating history acoustic transponder (CHAT) tags (Vemco), download their data, and in the case of so-called business card tags data from other tags encountered along the way is also downloaded, to moored or mobile acoustic receivers (Voegeli et al. 2001, Vemco pers. comm.). However, the current size of these tags limits the use of this technology to large species (e.g., D. gigas, Gilly et al. 2006). Although, with the technology steadily getting smaller and automatic down-loading versions being developed, it is envisaged that these types of tags will become more common in cephalopod movement/migration studies, as they are for sh and sharks (see Arnold and Dewar 2001). Only three published studies have used archi- val tags on cephalopods (ODor et al. 2002; Jackson et al. 2005; Gilly et al. 2006). Interest- ingly, the rst two studies both employed hybrid acoustic/archival tags to maximise the amount/ type of data collected and the ability to recover the archival tags, although the latter was only partially successful. Most notably, Gilly et al. (2006) tagged D. gigas in the Gulf of California using 96 archival tags (Lotek, Ontario, Canada, www.lotek.com) and 10 PAT tags, with only one of the archival tags recovered. Of the 10 PAT tags, seven up-loaded data and provided temper- ature, depth and horizontal movement (deter- mined from light intensity) data. The PAT tag data demonstrated that during seasonal migra- tions D. gigas can travel almost 100 km in around 3 days, with the direction of the migration matching prevailing currents. Most interestingly, D. gigas spent most of the daylight hours and some of the night in a midwater hypoxic zone below 250 m, the oxygen minimum layer (OML), with Gilly et al. (2006) suggesting that the squid were foraging in this layer as it overlapped with the prey-rich deep scattering layer. Laboratory experiments showed that D. gigas has physiolog- ical adaptations that allow it to actively forage in the OML (Gilly et al. 2006) Chemical tags This type of tagging incorporates compounds (e.g., uorescent chemicals, elements and iso- topes) into calcied tissues. The advantages of this type of tagging are that you can mark large numbers of animals simultaneously with minimal handling, all life history stages can be marked, the compounds do not affect growth and mortality, and the marks can be detected for a considerable time. Disadvantages include the need to sacrice the organism to detect the mark, often there are a limited number of unique marks possible, and detection may be limited to analytical methods (at least for elemental and isotopic tags). Fluorescent markers There have been very few published studies on cephalopods using uorescent markers to exam- ine movement, however, squid statoliths have been routinely marked with the uorescent com- pounds tetracycline and calcein in order to validate their age (see review by Jackson (1994)). Statoliths of hatchling (Fuentes et al. 2000) and subadult (100 g) (Sakaguchi et al. 2000) O. vulgaris were successfully stained by Rev Fish Biol Fisheries (2007) 17:401423 407 1 3 immersion of the animals in solutions of alizarin complexone. Subadult octopus showed the uo- rescent mark on their statoliths when recaptured after 31 days (Sakaguchi et al. 2000). These stud- ies suggest that uorescent marking of statoliths could be a viable technique for mass labelling of paralarval cephalopods to examine movement throughout their life history. As an example of the potential of this type of tagging, Jones et al. (1999) marked >10 million embryos of a reef sh with tetracycline to determine how far the larvae disperse. They examined 5000 juveniles to obtain 15 marked individuals. The authors estimated that they had marked between 0.5% and 2% of all embryos and therefore 1560% of juveniles return to the natal population. Elements and isotopes Besides uorescent compounds, a range of ele- ments (e.g., Sr, rare earth elements, Ennevor and Beames 1993; Giles and Attas 1993; Ennevor 1994) and isotopes (e.g., enriched isotopes of Ba and Sr, Munro et al. unpublisheddata) have alsobeen used to successfully mark sh via immersion. Strontium is the element most commonly used to mark sh (Brown and Harris 1995), but Sr:Ca ratios of freshwater can be as much as that of marine waters or higher (Kraus and Secor 2004) and therefore potential exists to confuse natural Sr signatures with articial Sr marks. This problemmay not be as great for cephalopods found in marine environ- ments since Sr:Ca of marine waters is relatively constant. Only one study has used Sr to mark cephalopod statoliths, but the objective was to validate growth increments rather then determine movement (Hurley et al. 1985). Squid (Illex illece- brosus) statoliths were marked by feeding the squid shrimp that had been soaked in a SrCl 2 solution (Hurley et al. 1985). Other elements occurring in low concentrations (e.g., rare earth elements) have had variable success in producing marks in sh especially over short time scales (Munro et al. unpublished data), but have not been applied to cephalopods. Enriched stable isotopes can be used to create unequivocal marks that cannot be mistaken for a natural signature since there is no way that a wild sh could have a similar signature (Munro et al. unpublished data). Other advantages of enriched stable isotope marking include that it is stress-free to the sh, the isotopes are naturally occurring and are stable (i.e. non- radiogenic) and therefore pose no environmental or human health risks, the method can be applied to any life history stage, and standard equipment (e.g., laser ablationICP-MS) canbe usedtoanalyse the otoliths since the isotope ratios are shifted so drastically (Munro et al. unpublished data). Whilst enriched stable isotopes are expensive, small con- centrations (e.g. 15 lg/l Ba and 100 lg/l Sr) are required to produce clear shifts in isotopic ratios (Munro et al. unpublished data). The work of Munro et al. (unpublished data) was focused on ngerlings of golden perch (Macquaria ambigua), but there is potential for the technique to be used on larvae. Presumably, this technique could also be used to mark cephalopods, and then questions related to movement and metapopulation dynam- ics could be addressed. The previous techniques described all require marking of thousands of ngerlings via immer- sion. A potentially easier method may be to mark the mother in an effort to mark her subsequent offspring. Recently, a new technique for arti- cially mass marking sh larvae, transgenerational marking, has been described (Thorrold et al. 2006). Briey, sh larvae are marked after gravid females injected with enriched isotopes transfer the isotope spike to the embryonic otoliths of their offspring (Thorrold et al. 2006). Thorrold et al. (2006) demonstrated that both a benthic- spawning and a pelagic-spawning sh had unequivocal isotope signatures over a range of dose rates and that marked larvae were found for at least 90 days after a single injection. This method may also have potential for use on cephalopods, but further work is required to determine this. If the method is applicable to cephalopods, then potential exists to mark indi- viduals that form part of breeding aggregations (e.g., giant Australian cuttlesh, Sepia apama) and determine the degree of natal homing. Natural tags Natural tags are organisms, compounds or phys- ical marks (e.g., colour patterns) that naturally 408 Rev Fish Biol Fisheries (2007) 17:401423 1 3 occur in the animal of interest and there is variability in their occurrence between regions, seasons, depth etc. The main advantage of these types of tags is that all animals are tagged and thus all specimens that are captured, regardless of size, represent a recovery. Furthermore natural behaviour is guaranteed throughout the entire tagging-period. The disadvantage, however, is the information stored in natural tags can be difcult to read and/or interpret. Parasites Parasites have long been used as biological tags or markers in shes (e.g., Sindermann 1983; Wil- liams et al. 1992; MacKenzie and Abaunza 1998) and can be used to examine movement/migration, providing there is variability in degrees of host infection or parasite genetics across the study area, the parasite is easily found and recognized in the host, and the parasite does not inuence the hosts health or behaviour (Williams et al 1992). After examining 2000 individuals from 10 species of cephalopods in Galician waters, Gonza lez et al. (2003) identied three ecological groupings of parasite fauna, coastal, intermediate and neri- to-oceanic, suggesting that the ecological niche of a cephalopod species is more important in deter- mining its parasite fauna than its phylogeny. Similarly, distinct coastal and oceanic parasite communities have been identied for a number of Atlantic ommastrephid species (Gaevskaya 1977, in Gonza lez et al. 2003; Gaevskaya and Nigma- tullin 1978; Bargov 1982; Nigmatullin and Shu- khgalter 1990, in Gonza lez et al. 2003). Geographic differences in parasite fauna made stock discrimination possible in the oceanic squid O. bartramii (Bower and Margolis 1991) and T. rhombus (Bower and Miyahara 2005), and indi- rectly provided information on movement. How- ever, explicit movement and migration studies require more detailed information on parasite- host specicity and parasite geographic distribu- tion (Nagasawa and Moravec 2002). Variation in infection by dicyemid mesozoans is found within cephalopod families, species, and individuals, with these parasites found in tropical, subtropical, temperate and polar regions, making them poten- tially useful for large-scale migration studies of benthic or epibenthic species (Finn et al. 2005). Parasites that infect juvenile shes at their nursery grounds but not adults at their feeding or spawning grounds are used to understand recruit- ment migration in shes (Williams et al. 1992). However, there is an almost complete lack of information on parasites of cephalopod paralar- vae (Vidal and Haimovici 1999), with parasites identied in paralarvae of only two species (I. argentinus, Vidal and Haimovici 1999; Sthenoteu- this oualaniensis, Vecchione 1991), and the method of infection not known in either case. Much more research into the parasite fauna of cephalopods is needed before parasites can be reliably used as natural tags, however, there may be the potential for parasites to provide a tagging technique for paralarval cephalopods, which is currently lacking. Molecular approaches Molecular genetic methods can be used to estimate migration rates of individuals between and among populations, but the estimates pro- duced are fundamentally different from those produced by physical tagging studies or direct observation. Genetic methods cannot estimate absolute numbers for contemporary migration, but they do offer a very powerful approach for estimating relative rates of gene ow among populations averaged over long timescales, and so are relevant to long-term changes in popula- tion demographics and distributions. Theoreti- cally, gene ow rates of only a few individuals per generation (i.e. migration rates <1%) are high enough to maintain genetic homogeneity among populations. So the detection of signicant genetic differences between areas or populations indicates that very low rates of effective migration are occurring, and the degree of genetic differen- tiation gives an estimate of relative rates. Molec- ular genetic approaches have been applied to a wide variety of species for the estimation of population connectivity and the inference of migration rates (e.g., Palumbi 2003), although to date their application to cephalopods has been limited (see below). Rev Fish Biol Fisheries (2007) 17:401423 409 1 3 Recent developments in genetic marker meth- odologies (e.g., microsatellite DNA regions) and analytical procedures (coalescent-based ap- proaches and maximum likelihood e.g., MI- GRATE, Beerli and Felsenstein 2001, or Bayesian e.g., BAYESASS+, Wilson and Rannala 2003) to estimate long-term or contemporary migration (respectively), hold the prospect of more relevant estimates of migration rates gen- erated from genetic data. These developments also have taken the molecular genetic approach closer to the direct observation and identication of migration at the individual level. Improved methods of estimating migration, identifying clusters of individuals, and identifying the popu- lation of origin of individuals are now available (see review by Pearse and Crandall 2004). These techniques have unravelled the complex dynamics of migration, dispersal and the spatial and/or temporal integrity of populations (Buonaccorsi et al. 2005; Weetman et al. 2006), but their potential contribution to understanding cephalo- pod biology may be limited to those cephalopods with patchy distributions, association of individ- uals with small home ranges, and limited adult migration or juvenile dispersal, such as reef- associated squid and Nautilus, octopus species with patchy distributions, and cuttlesh species with restricted dispersal. The large majority of genetic studies of ceph- alopods conducted to date have involved squid and these studies have shown that where no major barriers to migration occur (such as land masses, major hydrographic fronts, deep water, etc), squid exhibit very low or no genetic differentiation among populations throughout their geographical range. Such a pattern is common to both myopsid (e.g., Sepioteuthis australis, Triantallos and Adams 2001; Loligo gahi, Shaw et al. 2004) and oegopsid squid (e.g., T. pacicus, Kim 1993; Katugin 2002; Ommastrephes bartramii, Katugin, 2002; Illex argentinus, Adcock et al. 1999; Berry- teuthis magister, Katugin 1999). Genetic homogeneity suggests that there is effective dispersal/movement and interbreeding across large geographical areas, at least on time- scales of tens of generations. Where barriers to adult and/or paralarval migration/movement do occur, populations exhibit signicant genetic divergence indicating breakdown of gene ow. For example, gene frequencies in loliginid squid are signicantly different among populations to either side of cold upwelling systems on the western coasts of both South America (L. gahi, Shaw et al. 2004) and southern Africa (Loligo reynaudii, Shaw et al., unpublished data), indicat- ing that the upwelling is a complete barrier to both adult and paralarval movement. Similarly, stretches of deep water are indicated as barriers to migration and dispersal of neritic species such as S. australis (Triantallos and Adams 2001) and L. forbesi (Shaw et al. 1999), which otherwise exhibit panmixia across large geographical ranges. Other barriers to gene ow in squid include physical subdivision of species ranges (Doryteuthis pealeii, Herke and Folz 2002), river umes (Loligo plei, Herke and Folz 2002), different water mass boundaries (separating the genetically distinct peripheral form of S. aus- tralis, Triantallos and Adams 2001), and semi- isolated marine basins (B. magister, Katugin 2000). Extensive population genetic studies of cuttle- sh are limited to two species, Sepia ofcinalis and S. apama. Both species exhibit the same genetic pattern of isolation-by-distance (IBD), where populations show increasing genetic differentia- tion with increasing geographical distance, indi- cating locally restricted gene ow (i.e. generation- on-generation movement) over surprisingly small scales (<300 km) for such mobile species (Pe rez Losada et al. 1999, 2002; Kassahnet al. 2003). Both species also exhibit evidence of phylogeographic discontinuities associated with specic geographi- cal features of either current (Almeria-Oran and Ionian/Aegean Sea hydrographic front systems in the Mediterranean, Pe rez Losada et al. 2002, 2007) or historical importance (Bass Strait, Kassahn et al. 2003; Gibraltar Strait and Siculo-Tunisian Strait, Pe rez Losada et al. 2007). The important point concerning such historical genetic patterns for the consideration of migration is the fact that historical subdivision of cuttlesh populations caused by sea level changes during Pleistocene glaciations are still evident in the genetic structur- ing of present day populations indicating the slow timescale on which gene ow is occurring in these species: effective population movement/migration 410 Rev Fish Biol Fisheries (2007) 17:401423 1 3 rates along coasts must be very low. Low sea levels during the Pleistocene period also resulted in the isolation of demersal deep-water B. magister pop- ulations within the Sea of Japan basin so that they have accumulated notable genetic differences (compared to conspecic populations from adja- cent areas of the Northwest Pacic) and evolved into a separate subspecies (Katugin 2000). The few genetic studies of octopus to date, all involving Octopus vulgaris, indicate genetic homogeneity over large areas (many 100s of km) but with some evidence of differentiation at large geographical scales, for example between the western and eastern Mediterranean (Casu et al. 2002; Maltagliati et al. 2002). The microsat- ellite DNA study by Murphy et al. (2002) also described differentiation at more local scales (100s of km), associated with discontinuities in population distributions. These results indicate that the extensive gene ow and widespread genetic homogeneity predicted from the posses- sion of a pelagic paralarval dispersal stage in O. vulgaris may not necessarily be realised in natural populations, and that gene ow and paralarval dispersal may be locally restricted. Molecular genetics offers a powerful approach to estimating realised effective migration rates among populations, and continuing developments in markers and analytical techniques hold the promise of more detailed identication of mi- grants. However the biological and life history characteristics of many cephalopod species mean that assignment and migration estimation meth- ods may have limited success, and careful consid- eration is needed in matching species with suitable genetic techniques. To date genetic studies indicate that migration/dispersal in squids is extensive but can be blocked by major ocean- ographic features, and in cuttlesh and octopus migration/dispersal is more locally restricted than predictions from life history parameters would suggest (e.g., Shaw et al. 1999, 2004; Murphy et al. 2002; Pe rez Losada et al. 2002, 2007). Elemental signatures: statoliths as environmental recorders Chemical analyses of the naturally occurring elemental and/or isotopic signatures of sh otoliths are widely used and have been a useful tool for reconstructing a shs life history, including eluci- dating migration pathways (e.g., Tsukamoto et al. 1998; Campana 1999; Elsdon and Gillanders 2005, 2006). Chemical investigations on cephalopod statoliths are few, but there is growing interest in this eld of research. One study has also analysed C and N isotopes of the beak, primarily to infer trophic relationships, but also to indicate life long residency in the same water mass or migration between different areas (Cherel and Hobson 2005; see also Takai et al. 2000 for example using muscle tissue). Such analyses are based on the assump- tions that once deposited the otolith/statolith material is not resorbed or reworked over time, the structure of interest continues to growthrough- out the life of the organism, and that the chemical and physical environment in which the organism is found is reected in the chemistry of the otolith/ statolith (Campana 1999). The chemical informa- tion together with the growth increments found in these structures can be coupled to provide unprec- edented life stage-specic information on the movements of shes/cephalopods. Calcied structures can provide an accurate chronology of exposure to particular environ- mental conditions, including salinity, temperature and composition of ambient water, therefore providing information on movement between different water masses (for reviews regarding sh and sampling and analyses issues see Campana 1999; Elsdon and Gillanders 2003a; Gillanders 2005a, b). Although a number of studies have investigated how otolith chemistry changes with different environmental parameters (e.g., Elsdon and Gillanders 2002, 2003b, 2004), relatively few studies have focused on statoliths (but see Ikeda et al. 1997; Zacherl et al. 2003a, b; Arkhipkin et al. 2004; Landman et al. 2004; Zumholz et al. 2006, this issue, in press a, b). A negative relationship between levels of barium in the statoliths of Sepia ofcinalis and environmental temperature was found in a laboratory study (Zumholz et al. in press a), and a negative relationship between water temperature and Sr:Ca in statoliths of eld caught squid has also been determined (Ikeda et al. 1998; Arkhipkin et al. 2004). These results suggest that changes in Sr:Ca and/or Ba:Ca (and potentially other ele- Rev Fish Biol Fisheries (2007) 17:401423 411 1 3 ments such as U:Ca) may indicate movement to colder or warmer waters. For example, an increase in Sr:Ca, Ba:Ca (and U:Ca) in the outer region of G. fabricii statoliths is suggestive of movement to cooler water (Zumholtz et al. in press b). It should be noted that some other eld studies have not been able to nd a relationship between statolith chemical composition and tem- perature (e.g., Ikeda et al. 2003) and therefore further laboratory experiments on other species are needed. Lack of a relationship also highlights the need to have data on water chemistry to properly interpret such analyses. Likewise, oxy- gen isotopes have also been used to infer move- ment (e.g., Landman et al. 2004). Besides relating elemental analyses to environ- mental parameters, which typically requires experimental work to validate such relationships, it is also possible to determine natal origins of sub-adults/adults. Such an analysis requires using known origin hatchlings as a baseline data set and then retrospectively determining the natal origin of adults by analysing the region of the statolith formed during its paralarval/early juvenile life (e.g., S. australis, Pecl unpublished data) (for examples using sh see Gillanders and Kingsford 1996; Gillanders 2002). Before such analyses can proceed it is rst necessary to demonstrate that spatial differences in elemental signatures of hatchlings are found, which may not necessarily be the case (e.g., Sepia apama, Gillanders unpub- lished data). Several other assumptions also exist, namely that all source populations have been sampled (this assumption is also necessary for molecular methods) and that the temporal stabil- ity of elemental signatures is known or alterna- tively that adults are matched to the appropriate year class of paralarvae/juveniles (see Gillanders 2005b for further details). Spatially resolved statolith analyses have great potential to provide information on life history, habitat use and migrations of cephalopods. How- ever, due to their lack of growth increments (Tait 1980) octopus statoliths, unlike other cephalopod statoliths, are of little use for time-specic trace element studies. Stylets (also known as vestigial shells) are a little known structure unique to the octopoda and are thought to represent a remnant shell. Their composition is currently unknown, however, the stylet microstructure of Octopus pallidus has been found to have distinct concen- tric regions, a visible pre-hatch nucleus and age- related growth (daily growth increments) (Dou- bleday et al. 2006). Due to this microstructure, stylets are likely to incorporate elements from the environment on a chronological basis and there- fore have the potential to address questions on the dispersal patterns of both juveniles, including paralarvae, and adults. For studies using trace elements to reconstruct the life-history of individuals the main restric- tion at present is the lack of experimental laboratory data examining the relationship be- tween statolith (or stylet) chemistry and a range of environmental parameters. Such information is required for each species of interest since there are likely to be different responses for different species. However, for those studies that only aim to assign adults to their natal origin, there is no need to understand why elements vary, only that they do and this allows individuals to be assigned back to their natal habitat. Tracking shing eets Since the late 1990s the use of satellite data showing the location of shing lights have been increasingly used to examine the distribution and abundance of squid sheries, which use lights to attract squid, operating in various regions around the globe. The advantage of this technique is it allows for remote monitoring of species, which migrate in oceanic waters, and as such can be difcult to study. The disadvantages are this technique monitors the movement of the shing eet, not that of the squid directly, and it is only applicable to those sheries that use light to attract squid, however, these sheries are the largest and therefore often of the most interest. The majority of these are far seas jig sheries for ommastrephid squid, with the shing eets con- sisting of as many as 20,000 vessels (Kiyofuji and Saitoh 2004) each using up to 300 kW for light production (Rodhouse et al. 2001). The emission of light from squid shing vessels is detectable using imagery from the United States Air Force Defence Meteorological Satel- 412 Rev Fish Biol Fisheries (2007) 17:401423 1 3 lite ProgramOperational Linescan System (DMSP-OLS) (Rodhouse et al. 2001; Maxwell et al. 2004) which allows the spatial distribution of shing eets to be observed at a high level of resolution, at around 2.7 km (Cho et al. 1999). DMSP-OLS data have been successfully ground- truthed by comparing the location of shing vessels with synoptic data from aerial surveys (Maxwell et al. 2004) and Platform Transmitter Terminals xed to shing vessels (Waluda et al. 2004). Using DMSP-OLS data eet distribution can be tracked on a daily basis if required, and several studies have found that changes in eet distribution can be indicative of the eet tracking squid as they undertake migrations (e.g., I. argentinus, Haimovici et al. 1998; Waluda et al 2002; T. pacicus, Kiyofuji and Saitoh 2004). Whilst high intensity shing lights are visible from space, ordinary deck lights from shing vessels are generally not detected using DMSP- OLS (Maxwell et al. 2004) at the current level of image resolution. Perhaps, as satellite resolution improves, those squid sheries using small boats with weaker lights and hand-jigging methods, such as the artisanal shery for Dosidicus gigas in the Gulf of California (Morales-Bojo rquez et al. 2001), or the shery targeting Loligo reynaudii off the coast of South Africa (Roberts and Sauer 1994) will be observed in satellite imagery in the future, thus greatly expanding the use of this method. The inuence of environmental variability on movement and migration The embryonic development of many cephalo- pods has been shown to be highly temperature dependent, with eggs generally developing faster in warmer waters (ODor et al. 1982; Boyle et al. 1995; Forsythe et al. 2001). Spawning regions and the distribution of egg masses have been shown to be linked to specic temperature zones (Roberts and Sauer 1994; Ichii et al. 2004; Bower and Ichii 2005), with the distribution of many paralarval and juvenile squid linked to local regional ocean- ography (Vecchione 1999; Dawe et al. 2000; Sakurai et al. 2000; Waluda et al. 2001b; Ander- son and Rodhouse 2002). Pelagically spawning ommastrephid squid appear to utilise water masses in which density increases with depth (due to decreasing temperature or increasing salinity), allowing neutrally buoyant egg masses to be held within the mesopelagic zone and transported to suitable areas for hatching (ODor and Balch 1985; Bakun and Csirke 1998). Even in the deep sea, brooding deep-sea octopus Graneledone spp. have been observed in association with cold seep regions (Drazen et al. 2003). Various studies have linked environmental conditions during the spawning and hatching of cephalopods with recruitment success or failure, with availability of thermal resources shown to inuence population size in the following season (Bower 1996; Sakurai et al. 2000; Yatsu et al. 2000; Waluda et al. 2001a; Waluda and Rodhouse 2006). Sea surface temperature is the most easily obtained, and therefore the most common param- eter used to assess links between cephalopod distribution and environment, providing an easily obtained proxy for oceanographic variability. Other factors such as rainfall, river discharge (Sobrino et al. 2002), water turbidity (Roberts and Sauer 1994; Faure et al. 2000; Scho n et al. 2002), solar ux, sea level pressure, wind speed, wind direction (Denis et al. 2002) sea level variability (Miyahara et al. 2005) and salinity (Laughlin and Livingston 1982) may also be important in inuencing the distribution and migration of various cephalopod populations. Local oceanographic processes during the early life history stages may be particularly important in retaining paralarval squid in areas of high food availability. Retention processes have been shown to favour the recruitment of many ceph- alopod species (Table 1). Oceanographic vari- ability has also been shown to alter the patterns and timing of the migrations of some coastal species, such as octopods, loliginids and sepioids, suggesting that migration patterns may be linked to specic water masses. For example, the strength of the Atlantic currents entering the English Channel and North Sea can inuence the timing of S. ofcinalis (Wang et al. 2003) and L. forbesi migrations (Holme 1974; Waluda and Pierce 1998; Robin and Denis 1999), with the migration of L. forbesi into the English Channel Rev Fish Biol Fisheries (2007) 17:401423 413 1 3 Table 1 The inuence of oceanographic processes on cephalopod migration and sheries Species Region Oceanographic process Effect References Octopus vulgaris Central East Atlantic Retention of eggs/ juveniles Increased retention with increased upwelling at Arguin Bank, Mauritania Faure et al. (2000) Dosidicus gigas Eastern Pacic Retention of eggs/juveniles Increased retention with increased upwelling off Peru and Central America Vecchione (1999) and Waluda et al. (2006) Illex illecebrosus Northwest Atlantic Retention of eggs/juveniles Increased retention via transport processes in the Gulf Stream and slope water Perez and ODor (1998) and Dawe et al. (2000) Illex argentinus Southwest Atlantic Retention of eggs/juveniles Retention related to mesoscale variability at conuence of Falkland (Malvinas) and Brazil Currents Brunetti and Ivanovic (1992), Leta (1992) and Waluda et al. (2001a) Doryteuthis opalescens Northeast Pacic Retention of eggs/juveniles Increased retention via tidally reversing currents and inshore entrainment in Southern California Bight Zeidberg and Hamner (2002) Octopus vulgaris Northeast Atlantic Retention of eggs/juveniles Increased retention with increased upwelling, Ria of Vigo, Spain Gonza lez et al. (2005) Loligo vulgaris Northeast Atlantic Retention of eggs/juveniles Increased retention with increased upwelling, Ria of Vigo, Spain Gonza lez et al. (2005) Illex argentinus Southwest Atlantic Frontal processes at the shelf edge Paralarval distribution related to strong thermal gradients between Brazil Current and the convergence front Leta (1992) and Bazzino et al. (2005) Todarodes pacicus Northwest Pacic Frontal processes at the shelf edge Paralarvae/juveniles transported via the Kuroshio Extension to Sea of Japan Sakurai et al. (2000) and Kawabata et al. (2006) Illex illecebrosus Northwest Atlantic Frontal processes at the shelf edge Enhanced shoreward transport of paralarvae/ juveniles due to increased meandering of the Gulf Stream Rowell et al. (1985), Vecchione and Roper (1986) and Dawe et al. (2000) Thysanoteuthis rhombus Northwest Pacic Frontal processes at the shelf edge Transport of juveniles through the Tsushima Strait via Tsushima current Bower and Miyahara (2005) Todarodes pacicus Northwest Pacic Fishery aggregations at frontal regions Fishery located at boundary between Kuroshio and Oyashio currents Cho et al. (1999) Illex argentinus Southwest Atlantic Fishery aggregations at frontal regions Fishery located at frontal regions between Falkland current and coastal shelf waters Waluda et al. (2001b) Ommastrephes bartramii North Pacic Fishery aggregations at frontal regions Increased catches associated with strong thermal gradient at Subarctic boundary Ichii et al. (2004), Bower and Ichii (2005) Ommastrephes bartramii Northwest Pacic Rings, meanders, streamers Fishery occurs in streamers shed from the Kuroshio, Oyashio and Tsushima-Tsugaru Current systems Sugimoto and Tameishi (1992) Pterygioteuthis giardi Southwest Pacic Rings, meanders, streamers Squid associated with a warm core eddy of East Australia Current, Tasman Sea Brandt (1983) Brachioteuthis riisei Southwest Pacic Rings, meanders, streamers Squid associated with a warm core eddy of East Australia Current, Tasman Sea Brandt (1983) Martialia hyadesi Southern Ocean Rings, meanders, streamers Squid associated with warm core ring at Northeast Georgia Rise, Antarctic Polar Frontal Zone Rodhouse et al. (1996) 414 Rev Fish Biol Fisheries (2007) 17:401423 1 3 occurring earlier in years of higher than average temperature (Sims et al. 2001). Variability in current systems may also shift squid from their usual feeding areas. In the south Atlantic, westward shifting of the Falkland Current has been shown to displace part of the L. gahi population to a region north of the usual feeding area (Arkhipkin et al. 2006), whereas in the north Atlantic a southward shift in the position of the Gulf Stream (coupled with increased meandering and thus improved paralarval retention) has been shown to favour high abundance of I. illecebrosus in Canadian waters, at the north of the species range (Dawe et al. 2000). Those cephalopods that undertake the longest migrations, spanning several thousands of kilo- metres are generally large oceanic ommastrephids associated with high-energy western boundary current systems. Large boundary current systems provide both a highly productive environment and an environment where appropriate tempera- tures are available at all life stages (ODor and Coelho 1993), however, they are environmentally unpredictable areas. As such, the adult squid migrate from these boundary current systems to areas with lower productivity but more environ- mentally stability in order to spawn (Rodhouse and Nigmatullin 1996). The migration of young stages of many species is facilitated by their association with current systems, with paralarval or juvenile squid often associated with frontal processes at the shelf edge (Sakurai et al. 2000) (see Table 1 for examples). Thus, juvenile squid take advantage of high-speed transport systems to assist their migration to feeding areas. Current systems such as the Gulf Stream provide a rapid transport system, of up to 1,000 km per week for juvenile squid (Rowell et al. 1985). The further poleward the young squid travel, the larger they grow, the longer they take to return to spawning grounds and the greater their fecundity. Cephalopod abundance and distribution can be strongly inuenced by upwelling, which are areas of high productivity, with populations apparently shifting to less common parts of their species range during periods of extreme warming and reduced upwelling (Ichii et al. 2002; Zeidberg and Hamner 2002; Waluda and Rodhouse 2006). Variability in upwelling has also been found to restrict the movement of L. reynaudii to spawning areas off the coast of South Africa (Scho n et al. 2002), and to inuence the distribution of Sthe- noteuthis pteropus around the coast of western Africa (Zuyev et al. 2002). Frontal regions (dened by large temperature changes over a small distance) may be important for the accumulation of large concentrations of cephalopods (see Table 1 for examples), with the inuence of mesoscale processes at fronts such as the formation of rings, meanders and streamers also shown to inuence the distribution of om- mastrephid squid (see Table 1 for examples). Large-scale environmental variability may change the distribution patterns of squid species, and effect shifts in timing of seasonal movements and migrations, probably by altering the avail- ability of thermal resources (Sims et al. 2001). The inuence of the El Nin o-La Nin a, Southern Oscillation Index (SOI), North Atlantic Oscilla- tion (NAO) and Trans-Polar Index (TPI) have been shown to inuence the movement and migration of cephalopod populations in various locations around the globe, and the inuence of these phenomena can be seen in locations far removed from the source of variability, mediated by teleconnections inuencing local oceanogra- phy (Waluda et al. 1999). Variability in environment related to the Southern Oscillation and TPI were shown to have a large inuence on cephalopod species of importance to sheries, particularly during the juvenile and adult phases of the life cycle respec- tively (Waluda et al. 2004). In the Northern Hemisphere, interannual variability in the timing of L. forbesi migrations has been shown to be related to variability in the strength of the NAO (Sims et al. 2001). Higher abundances of I. illecebrosus were shown to be associated with a negative NAO index (Dawe et al. 2000), and the summer biomass of Eledone cirrhosa in the Ligurian Sea was shown to be related to the NAO index during the previous winter (Orsi Relini et al. 2006). In the eastern Pacic, the inuence of El Nin o variability has been observed in sheries for Doryteuthis opalescens off Califor- nia (Jackson and Domeier 2003; Maxwell et al. 2004; Reiss et al. 2004; Zeidberg et al. 2006) and D. gigas off Central and South America (Ichii Rev Fish Biol Fisheries (2007) 17:401423 415 1 3 et al. 2002; Waluda et al. 2006). Warming (El Nin o) events are generally seen to lead to a reduction in squid abundance in common shery areas coupled with an increase in catches outside these areas (Pearcy 2002; Zeidberg et al. 2006), suggesting El Nin o events could potentially cause a shift in distribution rather than a decline in overall population size. Interest in the inuence of environmental parameters on cephalopod movement/migration at the various life cycle stages is increasing. Much work has been done examining environmental inuences on the abundance and distribution of cephalopods (particularly those which form sig- nicant sheries), and the development of models utilising some of the observations described here to predict abundance and distributions are becoming more common (Roberts et al. 1998; Waluda et al. 1999; Agnew et al. 2002; Denis et al. 2002; Ichii et al. 2002; Pierce and Boyle 2003; Reiss et al. 2004; Zuur and Pierce 2004; Miyahara et al. 2005; Sacau et al. 2005), enabling a better understanding of the processes inuenc- ing cephalopod population variability. Under- standing these patterns will become increasingly important, with increased ocean temperatures predicted as a consequence of global warming, which could disrupt or change migration patterns for some cephalopod species, as is predicted for migrating pacic salmon (Welch et al. 1998). Conclusion In summary, over recent years we have seen many exciting developments in terms of our increasing ability to examine movement and migration of cephalopods, and understanding the inuence of environmental processes on this movement. Still many challenges lie ahead and it is hoped that this review can spark some interest in tackling the key questions that remain. One of the major questions that needs addressing is how to tag and track paralarval cephalopods, as our knowledge of this life stage is far less than that of adults. This is particularly important; as it is this stage which is most affected by environmental inuences, par- ticularly temperature and food uctuations. Natural tags, such as parasites or elemental signatures, hold promise for tracking paralarval cephalopods, as each animal caught is a recap- ture, however, large numbers of paralarvae need to be examined rst to dene the parasitic and elemental characteristics of various populations before these techniques could be used routinely. Perhaps, chemical tagging of paralarvae through maternal transfer will prove to be a successful technique, as large numbers of offspring could be tagged at once. However, extensive laboratory studies are rst required to determine if this technique is viable for cephalopods. Acknowledgements This review came about as a result of discussions held at the Cephalopod Movement and Migration Workshop held as part of the 2006 CIAC Conference in Hobart. The authors would like to thank all the participants for their valuable contribution to the discussion. We are also grateful to Hideaki Kidokoro, Ken Mori, Yoshikazu Nakamura, Kazuya Nagasawa, and Mitsuo Sakai for providing information on recent tagging experiments in Japan. Thanks also to Stephanie Semmens for editing the manuscript. Funding to JMS and GTP to study cephalopod movement and migration comes from Australian Research Council Linkage grants and Postdoctoral Fellowships (C00107233 and LP0347556 respectively). KZ was funded by the Deutsche Forschungesgemeinschaft (DFG PI 203/11-1, 11-2, 11-3, HA 2100/9-1). Travel to CIAC by PWS was supported by the Royal Society of London. References Adcock GJ, Shaw PW, Rodhouse PG, Carvalho GR (1999) Microsatellite analysis of genetic diversity in the squid Illex argentinus during a period of intensive shing. Mar Eco Prog Ser 187:171178 Agnew DJ, Beddington JR, Hill SL (2002) The potential use of environmental information to manage squid stocks. 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