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Active acoustic tracking suggests that soft sediment fishes can show site attachment: a preliminary assessment of the movement patterns of the blue-spotted flathead (Platycephalus caeruleopunctatus)

Active acoustic tracking suggests that soft sediment fishes can show site attachment: a... Background: It is generally considered that on relatively homogenous marine soft sediment habitats, such as sand, fish are unlikely to show site attachment. This poses challenges for management and the evaluation of the efficacy of marine protected areas, in which soft sediments often make up more than 70 % of habitats. The blue-spotted flathead is a commercially and recreationally targeted species found on soft sediments in coastal marine waters of south- eastern Australia. There are no published data on its movement patterns. Here, using active acoustic telemetry, we aim to (a) quantify movement and habitat use of blue-spotted flathead, (b) compare area usage to no-take sanctuary zone size and (c) obtain data to aid in the design of a large passive receiver array to be used in long-term comprehen- sive tracking of soft sediment fish. Results: Three of five blue-spotted flathead that were tagged exhibited strong site attachment and were detected close to their release points for the entire 60-day study period. The two other fish were not detected after 4 and 25 days and were likely to have moved out of the study area (search radius ≈ 3 km). For the three fish tracked over 60 days, the area used was compact (mean ± SE = 0.021 km ± 0.037) and two patterns of movement were appar- ent: (1) a small activity space used in its entirety each day (two fish) and (2) a larger activity space in which a separate area is utilised each day (one fish). Conclusions: Our study is the first to document the movement of blue-spotted flathead, and these preliminary results demonstrate two broad movement patterns shown by this species on soft sediments in Jervis Bay. Over the course of 60 days, a majority of fish in this study showed strong site attachment; however, a number of fish also made larger-scale movements. Finally, our study suggests that a tightly spaced, passive acoustic array would provide mean- ingful results for this species, although strategically placed receivers outside this array would be required to detect any longer range movements. site attachment and broader range movements [1]. An Background understanding of movement is particularly important as Soundly and effectively implementing and managing reserve effectiveness is dependent on the scale of move - marine protected areas (MPAs) requires knowledge of ment of species in relation to reserve size [2, 3]. Frequent species presence, abundance, size structure and also and large-scale movement of animals has been used to argue that MPAs are unlikely to have tangible benefits for *Correspondence: lcf775@uowmail.edu.au 1 wide ranging taxa [3]. For example, a spatial closure to School of Biological Sciences, University of Wollongong, Wollongong, NSW, Australia fishing such as a no-take sanctuary zone is thought to be Full list of author information is available at the end of the article © 2016 The Author(s). This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 3 of 11 less effective if the movement of the fish intended to be these data will provide a basis for the design of a large protected covers an area much larger than the area closed passive receiver array for long-term tracking of large to fishing [4]. If species display site attachment to areas numbers of soft sediment fish in a marine park (JBMP) well within reserve boundaries, then MPAs may have over appropriate spatial scales. The specific aims of this potential value; however, if significant numbers of indi - paper are to: (1) use active telemetry to examine blue- viduals have no site attachment and move between dif- spotted flathead movement patterns, behaviour and area ferent habitats or areas outside of the reserve boundaries, use, (2) compare movement to current no-take sanctuary then alternate management strategies may be more effec - zone size and (3) visualise patterns in activity space of tive [5]. blue-spotted flathead to better inform decision-making In many cases, particularly on marine soft sediments, on future tracking array design. little information on the habitat use and movement of fishes is available to inform MPA design and location. Methods Consequently, MPAs may not be of a suitable size or The study was undertaken in JBMP on the south coast in the correct location to provide effective protection. of NSW, Australia. Jervis Bay (Fig.  1) is approximately Understanding the habitats used, degree of site attach- 50  km and dominated by sub-tidal soft sediments (pre- ment and patterns of movement will substantially aid in dominately coarse sand). A mosaic of rocky intertidal, the design and management of MPAs, particularly where subtidal reefs and seagrass beds are scattered around the preferred fish habitat (such as spawning or aggregation edge of the bay. In addition, there are five designated no- grounds) can be identified [6]. Without such data, this take sanctuary zones within Jervis Bay where fishing is is impossible to assess or to infer the effectiveness of a not permitted; the remainder of the bay has zoning that marine reserve on soft sediments. allows for recreational fishing and limited forms of com - The homogeneous nature of marine soft sediments, mercial fishing (e.g. not trawling). The current zoning with little obvious structure or habitat differentiation, within the bay was implemented in 2002. appears to lead to a general assumption that fish will not On the 22 August 2011, blue-spotted flathead (n  =  5) show appreciable site attachment [7]. This is in compari - were line caught on sand at a depth of 10 m in the Hare son with reef-associated fishes which are often found to Bay no-take sanctuary zone (Fig.  1). The fish were then −1 show high levels of site attachment [8–11]. This assump - anaesthetised in seawater containing 60 mg L of Aqui- tion, however, is based on very little data, as relatively few S before a transmitter (Vemco V9 model; 21 mm length, studies look at the movement of demersal fish species on 9  mm diameter, 1.6  g in the water, battery life 80  days, open coastal marine soft sediments. This knowledge gap nominal ping interval 120 s) was inserted through a 1-cm appears incongruous with the fact that marine soft sedi- mid-ventral incision in the abdomen. Surgery lasted ments are the most common habitat on Earth [12], and <2 min and the incision was closed with one or two dis- comprises most of the habitat within near- and off-shore solving stitches tied with a double surgeon’s knot. Fish areas. Furthermore, although we have little data for the were then transferred to a holding tank and monitored effect of MPAs on soft sediment systems [13], marine for around 20  min, before releasing them at the site of soft sediments are often the major habitat type protected capture. within MPAs [7]. We actively tracked blue-spotted flathead for 12  days The blue-spotted flathead (Platycephalus caeruleopunc - over a 60-day period between 22 August and 20 Octo- tatus) is a common species found on marine sands in ber 2011, using a boat-based mobile receiver and direc- south-eastern Australia and is both commercially and tional hydrophone (Vemco VR100 and VH110). For the recreationally exploited [14]. Despite this, there are cur- first 4  days post-release, fish were tracked in daylight rently no published data on blue-spotted flathead move - hours, and we attempted to position each fish repeatedly ment patterns. This study sought to provide a preliminary throughout each day. Fish were then tracked on 8 ran- assessment of movement patterns within a temperate dom follow-up days in daylight hours, and we attempted MPA (Jervis Bay Marine Park—JBMP, NSW, Australia) to position the fish at least once on each of these days. to test the hypothesis that blue-spotted flathead would Previous trapping data in Jervis Bay suggested that not show any sign of site attachment (the consistent posi- blue-spotted flathead were not active at night. There - tioning of a fish within an area over the study period). fore, we decided not to track at night in this study and This study was carried out to inform the management of redirect the associated costs and effort to increase the the MPA, and more broadly, these preliminary data are study length. Fish were sequentially located, and after essential to aid in the design of marine reserves on soft we located the fish, which generally took between 10 and sediments and will go some way to filling a substantial 20  min to position to within 10 m, the position of the knowledge gap for this habitat. In terms of future studies, fish was recorded on a hand-held Garmin GPS 60 when Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 4 of 11 Fig. 1 Study location in Jervis Bay, NSW, Australia. Area where tagged fish were captured and released in Hare Bay no-take sanctuary zone is shown within the black square. All areas in shades of blue are marine sand; other major habitat types are indicated in the legend. Inset map: location of Jervis Bay in Australia. Subtidal reef features digitised preferentially from swath bathymetry, LADS, and ADS40 aerial imagery. Sources: NSW Department of Primary Industries, NSW Office of Environment and Heritage, Geoscience Australia. Mangrove, seagrass and saltmarsh boundaries as defined in [31] the signal strength was at its maximum (i.e. between and searching for the fish in circles of ever increasing size 70 and 90  dB). Previous range testing indicated that we out to maximum of 3 km. could reliably get to within 10  m of a tag to take a posi- tion. Subsequent searches commenced at the last known Data analysis position, and if the fish was not detected within 30 min, Positional data were visualised to evaluate movement we then searched for the next fish. Once several loca - patterns and site attachment. To estimate the activ- tions were recorded for each fish, a broader search pat - ity space for each fish, we used a fixed kernel method tern was implemented to try and locate any undetected to produce 95  % kernel utilisation distributions (KUDs; fish. This involved returning to the last known position default grid size/search radius of 50  ×  50  m and extent Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 5 of 11 of 1) which were visualised as 95 % probability contours. We calculated KUDs for the first 4  days of tracking and the entire tracking period to assess both post-release and short-term space use. KUDs were produced using the ‘adehabitatHR’ package in the statistical software R [15] and plotted as 95  % probability contours in the ZOATRACK interface [16]. To avoid fragmentation of Fig. 2 Daily presence–absence of five acoustically monitored blue- estimated activity spaces, Kie’s rule-based ad hoc method spotted flathead (P. caeruleopunctatus) in study area. Active tracking [17] was used to estimate a suitable smoothing parameter was undertaken on 12 days between August 22 and 20 October 2011 (h). The smoothing parameter was sequentially increased on days 1–4, 15, 18, 24, 25, 27, 36, 59 and 60 or decreased if required from the reference smoothing (h ) value by 0.10 increments, until the smallest continu- ref ous (rather than a number of discrete) 95  % KUD prob- ability contour that did not cut off any obvious paths The exception was F1 which used a much larger area between two subsequent detections was attained. We than the other fish and used a separate area on each of assumed uniform use of space within the 95 % probability the 4 days (Fig. 3; Table 1). F1 also moved a much further contour as the tracking strategy employed did not allow distance from tagging location, 534  m compared with a true estimate of core area use within the activity space. between 108 and 149  m for all other fish (209  ±  82  m; To indicate activity level, we used a minimum activ- mean  ±  SE). Activity over the 4  days was similar for all −1 ity index (MAI m  h ) [18] which was calculated by the fish with a MAI over the first 4  days ranging from 22.11 −1 −1 distance between two points divided by the time elapsed to 44.96 m h (29.34 m h  ± 4.15; mean ± SE, Table 1). between observations, averaged across all points for each Over 60 days, residency for the five fish averaged 74 % fish. The nature of the data collection meant that this (SE ± 14 %) suggesting strong site attachment (Table  1). was only possible for the first 4 days of intense tracking. Two fish (F2 and F3) appeared to move outside the no- A residency index (RI), as a proportion of total tracking take sanctuary zone after the first 4  days of intensive days detected, was calculated to give an estimate of site tracking, as searches well beyond the no-take sanctuary attachment. We make the assumption that where fish zone failed to detect these fish. Fish F2 did move back are not detected for two tracking days in succession they into the sanctuary zone, and was subsequently detected have left the study area. We also assume that fish remain on 2 days (days 24 and 25) to the south of the study area in the study area between two tracking days where they (Fig.  4). Despite extensive searches of the no-take sanc- are detected (e.g. if a fish is detected on day 18 of track - tuary zone and surrounding areas covering a minimum ing and then again on the next tracking day, day 24, we of 3-km radius around release point, we did not detect assume the fish stayed in the study area between those either fish again during the study. The three remaining days).We used displacement (D) given as distance in fish (including F5 which had the largest activity space metres from the release point to the final position after over the first 4  days) showed strong site attachment and 60 days and furthest distance (FD) from first release posi - were still being detected in Hare Bay sanctuary zone after tion (calculated for 4 and 60 days) to indicate straight line 60  days when the study concluded. The activity space distance that fish moved from the release point over the (95  % KUD) for the three fish remaining after 60  days study period. An additional file shows a detailed quanti - (0.121 km  ± 0.037; mean ± SE) was compact and much tative summary of movement pattern metrics including smaller than the ≈5.50  km of soft sediments within final h values and proportion of h reference (see Addi- Hare Bay sanctuary zone. F1 and F4 were detected on tional file 1: Table S1). all of the 12-day tracking which was undertaken, and F5 was detected on all but one tracking day (Fig.  2). Again, Results F1 covered the greatest amount of area, which was 2–4 All five of the tagged blue-spotted flathead (F1–F5) times greater than F4 and F5. Fish F1 also moved the fur- were active after tagging and detected on each of the thest distance from the tagging location over the 60 days first 4  days of post-tagging, and moved over a scale of (541 m), although its displacement at the end of the study 10–100s of metres within a day (Figs.  2, 3). The activity was only 108  m from the release point, compared with space (95  % KUD) over this time was generally compact 305 and 240 m for F4 and F5, respectively (Table 1). with a mean of 0.046 km  ± 0.025 (±SE). Most fish (F2– The three sfi h that were detected for the full 60-day F5) were continually reusing the same areas within their study length within the main study area each used a rela- activity space, with each animal’s positions being inter- tively small area but showed different movement patterns mingled through time over the 4  days (Fig.  3; Table  1). within their activity space (Fig.  4). F5 repeatedly used the Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 6 of 11 Fig. 3 Four-day activity space (95 % KUD) of five blue-spotted flathead (F1–F5). Calculated with positions obtained using active acoustic tracking over initial 4 days of continuous tracking between 22 and 25 August 2011. D1–D4 indicate tracking day for F1 (daily positions of F2–F5 were inter- mingled within their respective activity spaces) same area within its activity space. F4 used two areas rela- though the pattern of use varied greatly among individ- tively evenly within its activity space. F1 used the largest ual fish. The remaining two fish appeared to make much activity space and was detected in a separate area on each larger-scale movements. Tagged fish were only detected day it was tracked, but over the long-term revisited parts on soft sediments for the whole study period, and we did of its range visited earlier. Hence, for these fish, there was not detect blue-spotted flathead moving onto adjacent sea - consistency in terms of the usage of relatively small areas, grass or reef habitats despite these areas being searched. Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 7 of 11 Table 1 Quantitative summary of movement patterns of blue-spotted flathead over 4 and 60 days Fish ID Total length (mm) 4 days 60 days RI 95 % KUD FD MAI 95 % KUD D FD F1 400 0.11 534 30.55 0.211 108 541 1 F2 225 0.014 108 23.95 – – – 0.5 F3 402 0.015 109 44.96 – – – 0.33 F4 195 0.013 145 22.11 0.1 305 330 1 F5 432 0.010 149 25.11 0.051 240 240 0.92 Mean 331 0.046 209 29.34 0.121 218 370 0.75 SE 50 0.025 82 4.15 0.037 45 69 0.14 Fish total length (mm), furthest distance (FD) in metres from release point for the first 4 days and over 60 days. Displacement (D) in metres is distance from release 2 −1 point at study end. Activity space (km ) based on 95 % kernel utilisation distribution (KUD). Minimum activity index (MAI m h ) calculated by dividing the distance between two points by the time elapsed between observations. Residency index (RI) is given as a proportion of tracking days detected that fish on soft sediments would likely move over larger Discussion distances than those on hard substrata [7], blue-spot- This study demonstrated that a number of movement ted flathead in our study also exhibited short-term site patterns are exhibited by tagged blue-spotted flathead attachment comparable to many temperate reef fishes (Platycephalus caeruleopunctatus) found on soft sedi- (e.g. [11, 22]). In addition, blue-spotted flathead MAI of ments in Jervis Bay. Over a daily timescale, all fish in our −1 22.11–44.96 m h (mean ± SE = 29.34 ± 4.15) is much study used small relatively compact areas each day when lower than the reef-associated luderick (Girella tricuspi- actively tracked across daylight hours. Over periods of −1 data, 165.4  ±  74.87  m  h ; mean  ±  SE) assessed within up to 60 days, blue-spotted flathead in our study showed the same embayment and with the same tracking tech- two broad movement patterns; three out of five tagged nique [23]. fish showed strong site attachment and were detected on Two fish were lost from the study after 4 and 25  days. each day of tracking within the Hare Bay no-take sanctu- This was despite extensive searches of at least 3 km from ary zone. The remaining two fish appear to have moved their last recorded positions. The underlying reason for much larger distances of more than 3 km away from tag- this is unclear but could conceivably include capture, ging location. Given the perception that soft sediment tag failure, predation, or movement out of the study site. fishes are unlikely to show site attachment [7], and obser - Our observations suggest that blue-spotted flathead are vations that blue-spotted flathead can be strong active robust and survive surgery well; they recover readily from swimmers (Fetterplace personal observation from baited anaesthetic and, lacking a swim-bladder, are unaffected underwater video; see  data and materials section), it is by barotrauma. Previous tagging effects studies have particularly interesting that the majority of tagged fish in indicated that ‘tagging-induced’ mortality tends to occur our study showed such strong site attachment. The ability within the first 24 h after release [24]. Four out of five of of blue-spotted flathead to target many types of prey [19] our tagged fish were detected moving up to 25 days after coupled with the expected ambush predation by flathead surgery. This suggests that mortality from surgery in our species in general [20] could explain why blue-spotted study was unlikely. We would argue instead that the two flathead generally utilise relatively small areas over a day. fish that were not detected for the entire study simply Why some individuals continue to show this compact moved out of the study area. Capture is unlikely, at least space use over periods of 60 days and others move away in the study area, due to the study area being in a no-take is not clear. sanctuary zone. As these two fish may in fact have trav - Intriguingly, the movement patterns of the oceanic elled outside of tracking range, it follows that some part blue-spotted flathead assessed in this study are consistent of the population moves much greater distances than with those for estuarine dusky flathead (Platycephalus the averages estimated here. Why they moved remains fuscus) found in southern Australia [21]. Dusky flathead unclear and as our study is preliminary with a small sam- were found to be largely sedentary, often remaining in ple size it not possible to estimate exactly what portion of one section of Gippsland Lakes for months. A small the blue-spotted flathead population makes these larger- number of dusky flathead, however, were recorded mov - scale movements or how large these movements may be. ing up to 30 km over a few days. The use of active track - The larger-scale movements shown by two fish do not ing in our study provided high-resolution movement and appear to be driven by size, as both small and larger fish space-use patterns over a much smaller scale (10–100s left the study area and conversely both small and larger of metres). Unexpectedly, and contrary to suggestions Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 8 of 11 Fig. 4 Sixty-day activity space (95 % KUD) of three blue-spotted flathead (F1, F4, F5). Calculated with positions obtained using active acoustic track - ing over 60 days between 22 August and 20 October 2011. D1–D60 indicate tracking day for F1 and F4. Daily positions for F5 were intermingled within its activity space. The final 2 days of detections for F2 are also shown towards the southern edge of the figure fish also showed site attachment. As it is not possible to been reported to seasonally migrate in order to spawn, distinguish the sexes of blue-spotted flathead based on based on indirect evidence such as aggregation sight- markings or size (they are not known to show sexual size ings and the capture of spawning females around the dimorphism), it is more difficult to assess whether these mouths of estuaries [28]. While blue-spotted flathead are movements may be related to the sex of the fish. Many thought to spawn year round [26], there are no published fish make seasonal migrations at specific times of year evidence to support this and no evidence of migration (e.g. [25]), and the closely related dusky flathead have movements to date. Further investigation is required to Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 9 of 11 determine whether or not the larger movements shown expanding the duration and area of coverage, the cur- by some of our tagged fish are just roaming movements rent study has a number of implications for design of a over scales greater than our study size or are linked to large-scale tracking array. As a large tracking array can spawning movements. be expensive and time-consuming to install, our data We did not catch any blue-spotted flathead on, or provide guidance to best place passive receivers to cover detect tagged fish blue-spotted flathead moving onto this movement most efficiently. Our results indicate that seagrass or surrounding reef, suggesting that they are using a tightly spaced passive acoustic array for investiga- exclusively soft sediment fish. Our movement data sup - tion of the movement of this species is feasible and would ports findings of recent baited remote underwater video yield meaningful results. However, given the potential (BRUV) studies where no blue-spotted flathead were wider ranging movements of this species, using multiple recorded on reef within Jervis Bay (Rees, Davis and approaches would be useful to provide a more compre- Knott, unpublished data and Coleman et  al. [27]). How- hensive understanding of their movement patterns. At the ever, other BRUV studies have found very small numbers current study site, the entrance to Jervis Bay has now been of blue-spotted flathead on reef habitat; for example in gated and an array of receivers placed around the edge of Batemans Marine Park, Kelaher et al. [28] recorded blue- the bay. These extra receivers (also part of other ongoing spotted flathead on five out of 384 drops over 5 years; this studies) should provide a good idea of visitation to other raises the possibility that blue-spotted flathead occasion - sections of Jervis Bay and also detect if fish leave Jervis Bay. ally venture into edge areas of reef and seagrass habitats or reside there in very low numbers. Conclusions Many studies on the effectiveness or impacts of MPAs Our study, the first to document the movement of blue- have focused on changes in abundances and diversity, spotted flathead, provides clear evidence of short-term without taking into account critical information on site attachment and compact space use by part of the movement patterns of the species within them [2, 29]. blue-spotted flathead population in Jervis Bay. We also This is often because this information is not available or highlight the benefit of using active tracking as a first because while potentially very useful, quantifying the step in understanding the movement of unstudied spe- movement patterns and observing the natural behaviour cies. The area used by tagged fish showing site attach - of marine fish in the field is difficult to achieve. Without ment over a 60-day study period was much smaller than knowledge of the basic movement patterns of a species, no-take sanctuary zones on soft sediments in Jervis Bay it is difficult to predict effectiveness of spatial protection Marine Park. However, our results also suggest that measures such as MPAs [6]. Our study indicates that no- part of the population is also non-resident. While these take sanctuary zones protecting soft sediment habitats results suggest that blue-spotted flathead may respond in JBMP appear large enough to adequately encompass positively to protection provided by the no-take sanctu- the expected short-term movement of blue-spotted flat - ary zones in place, further tracking on a larger number head exhibiting site attachment. However, our data sug- of fish is needed to determine exactly what proportion of gest that two movement patterns are likely to exist within the population shows site attachment and if it continues the population, one that is highly site attached, and thus over the long term. Lastly, our results demonstrate that if would potentially benefit from MPAs, and one that tends we are to effectively manage fish found on soft sediments to roam, and thus may not benefit as much. If these pre - we need to revisit the current view that fish on this habi - liminary data are found to be representative of longer- tat are unlikely to show site attachment. term patterns of movement and activity space use by a large part of the blue-spotted flathead population, then it Additional file is likely that the Hare Bay no-take sanctuary zone is suf- ficiently large to provide protection for a large number of Additional file 1: Table S1. Detailed quantitative summary of move - blue-spotted flathead. If this is the case, we would suggest ment pattern metrics including final h values. that comparably sized zones on soft sediments in other areas of temperate Australia may also be appropriate. Though it is beyond the scope of this study, investigat - Abbreviations KUD: kernel utilisation distribution; JBMP: Jervis Bay Marine Park; MPA: marine ing what portion of the blue-spotted flathead population protected area; RI: residency index; D: displacement; FD: furthest distance; h: would need to show site attachment for spatial closures smoothing parameter; BRUV: baited remote underwater video. like MPAs to be effective will require tagging of much Authors’ contributions larger numbers of fish and deserves further attention. LF designed the study, conducted fieldwork, analysed data, drafted the manu- As this investigation was a preliminary assessment script and created the figures and graphics; AD designed the study, provided for movement of blue-spotted flathead with a view to materials and field resources and assisted in drafting of the manuscript; Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 10 of 11 NK designed the study, conducted fieldwork, provided materials and field 7. Caveen AJ, Sweeting CJ, Willis TJ, Polunin NVC. Are the scientific founda- resources and assisted in analyses and drafting of the manuscript; JN created tions of temperate marine reserves too warm and hard? Environ Conserv. the figures and graphics; MT provided materials and field resources, assisted 2012;39(3):199–203. in analyses and assisted in drafting of the manuscript. All authors read and 8. Ferguson AM, Harvey ES, Knott NA. Herbivore abundance, site fidelity and approved the final manuscript. grazing rates on temperate reefs inside and outside marine reserves. J Exp Mar Biol Ecol. 2016;478:96–105. Author details 9. Lee KA, Huveneers C, Macdonald T, Harcourt RG. Size isn’t everything: School of Biological Sciences, University of Wollongong, Wollongong, NSW, movements, home range, and habitat preferences of eastern blue Australia. Fish Thinkers Research Group, 11 Riverleigh Avenue, Gerroa, NSW gropers (Achoerodus viridis) demonstrate the efficacy of a small marine 2534, Australia. Fisheries NSW, New South Wales Department of Primary reserve. Aquat Conserv Mar Freshw Ecosyst. 2015;25(2):174–86. Industries, PO Box 5106, Wollongong, NSW, Australia. Port Stephens Fisher- 10. Harasti D, Lee KA, Gallen C, Hughes JM, Stewart J. Movements, home ies Institute, New South Wales Department of Primary Industries, Taylors range and site fidelity of snapper (Chrysophrys auratus) within a temper - Beach Rd, Taylors Beach, NSW, Australia. Jervis Bay Marine Park, New South ate marine protected area. PLoS ONE. 2015;10(11):e0142454. Wales Department of Primary Industries, 4 Woollamia Road, Huskisson, NSW, 11. Willis TJ, Parsons DM, Babcock RC. Evidence for long-term site fidelity of Australia. snapper (Pagrus auratus) within a marine reserve. NZ J Mar Freshw Res. 2001;35(3):581–90. Acknowledgements 12. Wilson WH. Competition and predation in marine soft-sediment com- We wish to thank the staff at Jervis Bay Marine Park that assisted in tracking of munities. Annu Rev Ecol Syst. 1991;21:221–41. flathead, in particular Ian Osterloh, Adrian Ferguson, Mark Fackerell, Matt Carr, 13. Sciberras M, Jenkins SR, Kaiser MJ, Hawkins SJ, Pullin AS. Evaluating the Matt Rees and Marie Claire Demers. We also thank Duane Byrnes for providing biological effectiveness of fully and partially protected marine areas. valuable GIS assistance and Margie Andreason for proof reading a number of Environ Evid. 2013;2(1):1–31. drafts. We acknowledge the efforts of two anonymous reviewers who helped 14. McGrouther M. Bluespotted Flathead, Platycephalus caeruleopunctatus to improve the initial manuscript substantially. McCulloch, 1922. Australian Museum Website. Published 15 September 2012. http://australianmuseum.net.au/eastern-blue-spotted-flathead- Competing interests platycephalus-caeruleopunctatus-mcculloch-1922. Accessed 20 June The authors declare that they have no competing interests. 2016. 15. R Development Core Team R. A language and environment for statisti- Availability of data and materials cal computing. R Foundation for Statistical Computing, Vienna. ISBN: The data set supporting the conclusions of this article is available in 3-900051-07-0. 2010. http://wwwR-project.org. the Zoatrack repository (publisher: the Atlas of Living Australia), DOI: 16. Dwyer R, Brooking C, Brimblecombe W, Campbell H, Hunter J, Watts M, 10.4226/68/5701CE37BD10D, DOI URL: http://dx.doi.org/10.4226/68/5701CE3 Franklin C. An open Web-based system for the analysis and sharing of 7BD10D [30]. Blue-spotted flathead baited underwater footage can be found animal tracking data. Anim Biotelem. 2015;3(1):1. at: https://vimeo.com/162039402. 17. Kie JG. A rule-based ad hoc method for selecting a bandwidth in kernel home-range analyses. Anim Biotelem. 2013;1(1):1–12. Ethics approval 18. 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Additional student funding was provided by the abundant benthic ambush-feeding teleosts in coastal waters of south- University of Wollongong. western Australia. J Mar Biol Assoc UK. 1998;78(02):587–608. 21. Hindell JS. Determining patterns of use by black bream Acanthopagrus Received: 16 April 2016 Accepted: 5 July 2016 butcheri (Munro, 1949) of re-established habitat in a south-eastern Aus- tralian estuary. J Fish Biol. 2007;71(5):1331–46. 22. Edgar GJ, Barrett NS, Morton AJ. Patterns of fish movement on eastern Tasmanian rocky reefs. Environ Biol Fishes. 2004;70(3):273–84. 23. Ferguson AM, Harvey ES, Taylor MD, Knott NA. A herbivore knows its patch: luderick, Girella tricuspidata, exhibit strong site fidelity on shallow References subtidal reefs in a temperate marine park. PLoS ONE. 2013;8(5):e65838. 1. Topping D, Lowe C, Caselle J. Home range and habitat utilization of adult 24. Thorstad EB, Næsje TF, Fiske P, Finstad B. Eec ff ts of hook and release California sheephead-Semicossyphus pulcher (Labridae), in a temperate on Atlantic salmon in the River Alta, northern Norway. Fish Res. no-take marine reserve. Mar Biol. 2005;147(2):301–11. 2003;60(2–3):293–307. 2. Gerber LR, Botsford LW, Hastings A, Possingham HP, Gaines SD, 25. Hirose T, Minami T. Spawning grounds and maturation status in adult Palumbi SR, Andelman S. Population models for marine reserve flathead flounder (Hippoglossoides dubius); off Niigata Prefecture, Sea of design: a retrospective and prospective synthesis. Ecol Appl. Japan. Fish Sci. 2007;73(1):81–6. 2003;13(sp1):47–64. 26. Rowling K, Hegarty A, Ives M. Bluespotted flathead (Playtcephalus 3. Kramer D, Chapman M. Implications of fish home range size caeruleopunctatus). In status of fisheries resources in NSW 2008/09, NSW and relocation for marine reserve function. Environ Biol Fishes. Industry & Investment, Cronulla; 2010, p. 47–9. 1999;55(1–2):65–79. 27. Coleman MA, Bates AE, Stuart-Smith RD, Malcolm HA, Harasti D, Jordan 4. Gaines SD, White C, Carr MH, Palumbi SR. Designing marine reserve net- A, Knott NA, Edgar GJ, Kelaher BP. Functional traits reveal early responses works for both conservation and fisheries management. Proc Natl Acad in marine reserves following protection from fishing. Divers Distrib. Sci. 2010;107(43):18286–93. 2015;21(8):876–87. 5. Palumbi SR. Marine reserves and ocean neighborhoods: the spatial scale 28. Kelaher BP, Coleman MA, Broad A, Rees MJ, Jordan A, Davis AR. Changes of marine populations and their management. Annu Rev Environ Resour. in fish assemblages following the establishment of a network of 2004;29(1):31–68. no-take marine reserves and partially-protected areas. PLoS ONE. 6. Grüss A, Kaplan DM, Guénette S, Roberts CM, Botsford LW. Consequences 2014;9(1):e85825. of adult and juvenile movement for marine protected areas. Biol Conserv. 29. Claudet J, Osenberg CW, Benedetti-Cecchi L, Domenici P, García-Charton 2011;144(2):692–702. J-A, Pérez-Ruzafa Á, Badalamenti F, Bayle-Sempere J, Brito A, Bulleri F, Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 11 of 11 Culioli J-M, Dimech M, Falcón JM, Guala I, Milazzo M, Sánchez-Meca J, 31. Williams RJ, West G, Morrison D, Creese RG. Estuarine resources of NSW. Somerfield PJ, Stobart B, Vandeperre F, Valle C. Marine reserves: size and In: The NSW comprehensive coastal assessment toolkit. CD ROM. ISBN: 0 age do matter. London: Wiley-Blackwell; 2008. p. 481–9. 7347 5805 7. NSW Department of Planning; 2007. 30. Fetterplace LC, Taylor MD, Knott NA. Data from: ‘Jervis Bay Marine Park: active tracking of blue-spotted flathead’. http://ZoaTrack.org; 2016. 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Active acoustic tracking suggests that soft sediment fishes can show site attachment: a preliminary assessment of the movement patterns of the blue-spotted flathead (Platycephalus caeruleopunctatus)

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Springer Journals
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Copyright © 2016 by The Author(s)
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Life Sciences; Animal Systematics/Taxonomy/Biogeography; Conservation Biology/Ecology; Terrestial Ecology; Bioinformatics; Freshwater & Marine Ecology
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2050-3385
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10.1186/s40317-016-0107-6
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Abstract

Background: It is generally considered that on relatively homogenous marine soft sediment habitats, such as sand, fish are unlikely to show site attachment. This poses challenges for management and the evaluation of the efficacy of marine protected areas, in which soft sediments often make up more than 70 % of habitats. The blue-spotted flathead is a commercially and recreationally targeted species found on soft sediments in coastal marine waters of south- eastern Australia. There are no published data on its movement patterns. Here, using active acoustic telemetry, we aim to (a) quantify movement and habitat use of blue-spotted flathead, (b) compare area usage to no-take sanctuary zone size and (c) obtain data to aid in the design of a large passive receiver array to be used in long-term comprehen- sive tracking of soft sediment fish. Results: Three of five blue-spotted flathead that were tagged exhibited strong site attachment and were detected close to their release points for the entire 60-day study period. The two other fish were not detected after 4 and 25 days and were likely to have moved out of the study area (search radius ≈ 3 km). For the three fish tracked over 60 days, the area used was compact (mean ± SE = 0.021 km ± 0.037) and two patterns of movement were appar- ent: (1) a small activity space used in its entirety each day (two fish) and (2) a larger activity space in which a separate area is utilised each day (one fish). Conclusions: Our study is the first to document the movement of blue-spotted flathead, and these preliminary results demonstrate two broad movement patterns shown by this species on soft sediments in Jervis Bay. Over the course of 60 days, a majority of fish in this study showed strong site attachment; however, a number of fish also made larger-scale movements. Finally, our study suggests that a tightly spaced, passive acoustic array would provide mean- ingful results for this species, although strategically placed receivers outside this array would be required to detect any longer range movements. site attachment and broader range movements [1]. An Background understanding of movement is particularly important as Soundly and effectively implementing and managing reserve effectiveness is dependent on the scale of move - marine protected areas (MPAs) requires knowledge of ment of species in relation to reserve size [2, 3]. Frequent species presence, abundance, size structure and also and large-scale movement of animals has been used to argue that MPAs are unlikely to have tangible benefits for *Correspondence: lcf775@uowmail.edu.au 1 wide ranging taxa [3]. For example, a spatial closure to School of Biological Sciences, University of Wollongong, Wollongong, NSW, Australia fishing such as a no-take sanctuary zone is thought to be Full list of author information is available at the end of the article © 2016 The Author(s). This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 3 of 11 less effective if the movement of the fish intended to be these data will provide a basis for the design of a large protected covers an area much larger than the area closed passive receiver array for long-term tracking of large to fishing [4]. If species display site attachment to areas numbers of soft sediment fish in a marine park (JBMP) well within reserve boundaries, then MPAs may have over appropriate spatial scales. The specific aims of this potential value; however, if significant numbers of indi - paper are to: (1) use active telemetry to examine blue- viduals have no site attachment and move between dif- spotted flathead movement patterns, behaviour and area ferent habitats or areas outside of the reserve boundaries, use, (2) compare movement to current no-take sanctuary then alternate management strategies may be more effec - zone size and (3) visualise patterns in activity space of tive [5]. blue-spotted flathead to better inform decision-making In many cases, particularly on marine soft sediments, on future tracking array design. little information on the habitat use and movement of fishes is available to inform MPA design and location. Methods Consequently, MPAs may not be of a suitable size or The study was undertaken in JBMP on the south coast in the correct location to provide effective protection. of NSW, Australia. Jervis Bay (Fig.  1) is approximately Understanding the habitats used, degree of site attach- 50  km and dominated by sub-tidal soft sediments (pre- ment and patterns of movement will substantially aid in dominately coarse sand). A mosaic of rocky intertidal, the design and management of MPAs, particularly where subtidal reefs and seagrass beds are scattered around the preferred fish habitat (such as spawning or aggregation edge of the bay. In addition, there are five designated no- grounds) can be identified [6]. Without such data, this take sanctuary zones within Jervis Bay where fishing is is impossible to assess or to infer the effectiveness of a not permitted; the remainder of the bay has zoning that marine reserve on soft sediments. allows for recreational fishing and limited forms of com - The homogeneous nature of marine soft sediments, mercial fishing (e.g. not trawling). The current zoning with little obvious structure or habitat differentiation, within the bay was implemented in 2002. appears to lead to a general assumption that fish will not On the 22 August 2011, blue-spotted flathead (n  =  5) show appreciable site attachment [7]. This is in compari - were line caught on sand at a depth of 10 m in the Hare son with reef-associated fishes which are often found to Bay no-take sanctuary zone (Fig.  1). The fish were then −1 show high levels of site attachment [8–11]. This assump - anaesthetised in seawater containing 60 mg L of Aqui- tion, however, is based on very little data, as relatively few S before a transmitter (Vemco V9 model; 21 mm length, studies look at the movement of demersal fish species on 9  mm diameter, 1.6  g in the water, battery life 80  days, open coastal marine soft sediments. This knowledge gap nominal ping interval 120 s) was inserted through a 1-cm appears incongruous with the fact that marine soft sedi- mid-ventral incision in the abdomen. Surgery lasted ments are the most common habitat on Earth [12], and <2 min and the incision was closed with one or two dis- comprises most of the habitat within near- and off-shore solving stitches tied with a double surgeon’s knot. Fish areas. Furthermore, although we have little data for the were then transferred to a holding tank and monitored effect of MPAs on soft sediment systems [13], marine for around 20  min, before releasing them at the site of soft sediments are often the major habitat type protected capture. within MPAs [7]. We actively tracked blue-spotted flathead for 12  days The blue-spotted flathead (Platycephalus caeruleopunc - over a 60-day period between 22 August and 20 Octo- tatus) is a common species found on marine sands in ber 2011, using a boat-based mobile receiver and direc- south-eastern Australia and is both commercially and tional hydrophone (Vemco VR100 and VH110). For the recreationally exploited [14]. Despite this, there are cur- first 4  days post-release, fish were tracked in daylight rently no published data on blue-spotted flathead move - hours, and we attempted to position each fish repeatedly ment patterns. This study sought to provide a preliminary throughout each day. Fish were then tracked on 8 ran- assessment of movement patterns within a temperate dom follow-up days in daylight hours, and we attempted MPA (Jervis Bay Marine Park—JBMP, NSW, Australia) to position the fish at least once on each of these days. to test the hypothesis that blue-spotted flathead would Previous trapping data in Jervis Bay suggested that not show any sign of site attachment (the consistent posi- blue-spotted flathead were not active at night. There - tioning of a fish within an area over the study period). fore, we decided not to track at night in this study and This study was carried out to inform the management of redirect the associated costs and effort to increase the the MPA, and more broadly, these preliminary data are study length. Fish were sequentially located, and after essential to aid in the design of marine reserves on soft we located the fish, which generally took between 10 and sediments and will go some way to filling a substantial 20  min to position to within 10 m, the position of the knowledge gap for this habitat. In terms of future studies, fish was recorded on a hand-held Garmin GPS 60 when Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 4 of 11 Fig. 1 Study location in Jervis Bay, NSW, Australia. Area where tagged fish were captured and released in Hare Bay no-take sanctuary zone is shown within the black square. All areas in shades of blue are marine sand; other major habitat types are indicated in the legend. Inset map: location of Jervis Bay in Australia. Subtidal reef features digitised preferentially from swath bathymetry, LADS, and ADS40 aerial imagery. Sources: NSW Department of Primary Industries, NSW Office of Environment and Heritage, Geoscience Australia. Mangrove, seagrass and saltmarsh boundaries as defined in [31] the signal strength was at its maximum (i.e. between and searching for the fish in circles of ever increasing size 70 and 90  dB). Previous range testing indicated that we out to maximum of 3 km. could reliably get to within 10  m of a tag to take a posi- tion. Subsequent searches commenced at the last known Data analysis position, and if the fish was not detected within 30 min, Positional data were visualised to evaluate movement we then searched for the next fish. Once several loca - patterns and site attachment. To estimate the activ- tions were recorded for each fish, a broader search pat - ity space for each fish, we used a fixed kernel method tern was implemented to try and locate any undetected to produce 95  % kernel utilisation distributions (KUDs; fish. This involved returning to the last known position default grid size/search radius of 50  ×  50  m and extent Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 5 of 11 of 1) which were visualised as 95 % probability contours. We calculated KUDs for the first 4  days of tracking and the entire tracking period to assess both post-release and short-term space use. KUDs were produced using the ‘adehabitatHR’ package in the statistical software R [15] and plotted as 95  % probability contours in the ZOATRACK interface [16]. To avoid fragmentation of Fig. 2 Daily presence–absence of five acoustically monitored blue- estimated activity spaces, Kie’s rule-based ad hoc method spotted flathead (P. caeruleopunctatus) in study area. Active tracking [17] was used to estimate a suitable smoothing parameter was undertaken on 12 days between August 22 and 20 October 2011 (h). The smoothing parameter was sequentially increased on days 1–4, 15, 18, 24, 25, 27, 36, 59 and 60 or decreased if required from the reference smoothing (h ) value by 0.10 increments, until the smallest continu- ref ous (rather than a number of discrete) 95  % KUD prob- ability contour that did not cut off any obvious paths The exception was F1 which used a much larger area between two subsequent detections was attained. We than the other fish and used a separate area on each of assumed uniform use of space within the 95 % probability the 4 days (Fig. 3; Table 1). F1 also moved a much further contour as the tracking strategy employed did not allow distance from tagging location, 534  m compared with a true estimate of core area use within the activity space. between 108 and 149  m for all other fish (209  ±  82  m; To indicate activity level, we used a minimum activ- mean  ±  SE). Activity over the 4  days was similar for all −1 ity index (MAI m  h ) [18] which was calculated by the fish with a MAI over the first 4  days ranging from 22.11 −1 −1 distance between two points divided by the time elapsed to 44.96 m h (29.34 m h  ± 4.15; mean ± SE, Table 1). between observations, averaged across all points for each Over 60 days, residency for the five fish averaged 74 % fish. The nature of the data collection meant that this (SE ± 14 %) suggesting strong site attachment (Table  1). was only possible for the first 4 days of intense tracking. Two fish (F2 and F3) appeared to move outside the no- A residency index (RI), as a proportion of total tracking take sanctuary zone after the first 4  days of intensive days detected, was calculated to give an estimate of site tracking, as searches well beyond the no-take sanctuary attachment. We make the assumption that where fish zone failed to detect these fish. Fish F2 did move back are not detected for two tracking days in succession they into the sanctuary zone, and was subsequently detected have left the study area. We also assume that fish remain on 2 days (days 24 and 25) to the south of the study area in the study area between two tracking days where they (Fig.  4). Despite extensive searches of the no-take sanc- are detected (e.g. if a fish is detected on day 18 of track - tuary zone and surrounding areas covering a minimum ing and then again on the next tracking day, day 24, we of 3-km radius around release point, we did not detect assume the fish stayed in the study area between those either fish again during the study. The three remaining days).We used displacement (D) given as distance in fish (including F5 which had the largest activity space metres from the release point to the final position after over the first 4  days) showed strong site attachment and 60 days and furthest distance (FD) from first release posi - were still being detected in Hare Bay sanctuary zone after tion (calculated for 4 and 60 days) to indicate straight line 60  days when the study concluded. The activity space distance that fish moved from the release point over the (95  % KUD) for the three fish remaining after 60  days study period. An additional file shows a detailed quanti - (0.121 km  ± 0.037; mean ± SE) was compact and much tative summary of movement pattern metrics including smaller than the ≈5.50  km of soft sediments within final h values and proportion of h reference (see Addi- Hare Bay sanctuary zone. F1 and F4 were detected on tional file 1: Table S1). all of the 12-day tracking which was undertaken, and F5 was detected on all but one tracking day (Fig.  2). Again, Results F1 covered the greatest amount of area, which was 2–4 All five of the tagged blue-spotted flathead (F1–F5) times greater than F4 and F5. Fish F1 also moved the fur- were active after tagging and detected on each of the thest distance from the tagging location over the 60 days first 4  days of post-tagging, and moved over a scale of (541 m), although its displacement at the end of the study 10–100s of metres within a day (Figs.  2, 3). The activity was only 108  m from the release point, compared with space (95  % KUD) over this time was generally compact 305 and 240 m for F4 and F5, respectively (Table 1). with a mean of 0.046 km  ± 0.025 (±SE). Most fish (F2– The three sfi h that were detected for the full 60-day F5) were continually reusing the same areas within their study length within the main study area each used a rela- activity space, with each animal’s positions being inter- tively small area but showed different movement patterns mingled through time over the 4  days (Fig.  3; Table  1). within their activity space (Fig.  4). F5 repeatedly used the Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 6 of 11 Fig. 3 Four-day activity space (95 % KUD) of five blue-spotted flathead (F1–F5). Calculated with positions obtained using active acoustic tracking over initial 4 days of continuous tracking between 22 and 25 August 2011. D1–D4 indicate tracking day for F1 (daily positions of F2–F5 were inter- mingled within their respective activity spaces) same area within its activity space. F4 used two areas rela- though the pattern of use varied greatly among individ- tively evenly within its activity space. F1 used the largest ual fish. The remaining two fish appeared to make much activity space and was detected in a separate area on each larger-scale movements. Tagged fish were only detected day it was tracked, but over the long-term revisited parts on soft sediments for the whole study period, and we did of its range visited earlier. Hence, for these fish, there was not detect blue-spotted flathead moving onto adjacent sea - consistency in terms of the usage of relatively small areas, grass or reef habitats despite these areas being searched. Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 7 of 11 Table 1 Quantitative summary of movement patterns of blue-spotted flathead over 4 and 60 days Fish ID Total length (mm) 4 days 60 days RI 95 % KUD FD MAI 95 % KUD D FD F1 400 0.11 534 30.55 0.211 108 541 1 F2 225 0.014 108 23.95 – – – 0.5 F3 402 0.015 109 44.96 – – – 0.33 F4 195 0.013 145 22.11 0.1 305 330 1 F5 432 0.010 149 25.11 0.051 240 240 0.92 Mean 331 0.046 209 29.34 0.121 218 370 0.75 SE 50 0.025 82 4.15 0.037 45 69 0.14 Fish total length (mm), furthest distance (FD) in metres from release point for the first 4 days and over 60 days. Displacement (D) in metres is distance from release 2 −1 point at study end. Activity space (km ) based on 95 % kernel utilisation distribution (KUD). Minimum activity index (MAI m h ) calculated by dividing the distance between two points by the time elapsed between observations. Residency index (RI) is given as a proportion of tracking days detected that fish on soft sediments would likely move over larger Discussion distances than those on hard substrata [7], blue-spot- This study demonstrated that a number of movement ted flathead in our study also exhibited short-term site patterns are exhibited by tagged blue-spotted flathead attachment comparable to many temperate reef fishes (Platycephalus caeruleopunctatus) found on soft sedi- (e.g. [11, 22]). In addition, blue-spotted flathead MAI of ments in Jervis Bay. Over a daily timescale, all fish in our −1 22.11–44.96 m h (mean ± SE = 29.34 ± 4.15) is much study used small relatively compact areas each day when lower than the reef-associated luderick (Girella tricuspi- actively tracked across daylight hours. Over periods of −1 data, 165.4  ±  74.87  m  h ; mean  ±  SE) assessed within up to 60 days, blue-spotted flathead in our study showed the same embayment and with the same tracking tech- two broad movement patterns; three out of five tagged nique [23]. fish showed strong site attachment and were detected on Two fish were lost from the study after 4 and 25  days. each day of tracking within the Hare Bay no-take sanctu- This was despite extensive searches of at least 3 km from ary zone. The remaining two fish appear to have moved their last recorded positions. The underlying reason for much larger distances of more than 3 km away from tag- this is unclear but could conceivably include capture, ging location. Given the perception that soft sediment tag failure, predation, or movement out of the study site. fishes are unlikely to show site attachment [7], and obser - Our observations suggest that blue-spotted flathead are vations that blue-spotted flathead can be strong active robust and survive surgery well; they recover readily from swimmers (Fetterplace personal observation from baited anaesthetic and, lacking a swim-bladder, are unaffected underwater video; see  data and materials section), it is by barotrauma. Previous tagging effects studies have particularly interesting that the majority of tagged fish in indicated that ‘tagging-induced’ mortality tends to occur our study showed such strong site attachment. The ability within the first 24 h after release [24]. Four out of five of of blue-spotted flathead to target many types of prey [19] our tagged fish were detected moving up to 25 days after coupled with the expected ambush predation by flathead surgery. This suggests that mortality from surgery in our species in general [20] could explain why blue-spotted study was unlikely. We would argue instead that the two flathead generally utilise relatively small areas over a day. fish that were not detected for the entire study simply Why some individuals continue to show this compact moved out of the study area. Capture is unlikely, at least space use over periods of 60 days and others move away in the study area, due to the study area being in a no-take is not clear. sanctuary zone. As these two fish may in fact have trav - Intriguingly, the movement patterns of the oceanic elled outside of tracking range, it follows that some part blue-spotted flathead assessed in this study are consistent of the population moves much greater distances than with those for estuarine dusky flathead (Platycephalus the averages estimated here. Why they moved remains fuscus) found in southern Australia [21]. Dusky flathead unclear and as our study is preliminary with a small sam- were found to be largely sedentary, often remaining in ple size it not possible to estimate exactly what portion of one section of Gippsland Lakes for months. A small the blue-spotted flathead population makes these larger- number of dusky flathead, however, were recorded mov - scale movements or how large these movements may be. ing up to 30 km over a few days. The use of active track - The larger-scale movements shown by two fish do not ing in our study provided high-resolution movement and appear to be driven by size, as both small and larger fish space-use patterns over a much smaller scale (10–100s left the study area and conversely both small and larger of metres). Unexpectedly, and contrary to suggestions Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 8 of 11 Fig. 4 Sixty-day activity space (95 % KUD) of three blue-spotted flathead (F1, F4, F5). Calculated with positions obtained using active acoustic track - ing over 60 days between 22 August and 20 October 2011. D1–D60 indicate tracking day for F1 and F4. Daily positions for F5 were intermingled within its activity space. The final 2 days of detections for F2 are also shown towards the southern edge of the figure fish also showed site attachment. As it is not possible to been reported to seasonally migrate in order to spawn, distinguish the sexes of blue-spotted flathead based on based on indirect evidence such as aggregation sight- markings or size (they are not known to show sexual size ings and the capture of spawning females around the dimorphism), it is more difficult to assess whether these mouths of estuaries [28]. While blue-spotted flathead are movements may be related to the sex of the fish. Many thought to spawn year round [26], there are no published fish make seasonal migrations at specific times of year evidence to support this and no evidence of migration (e.g. [25]), and the closely related dusky flathead have movements to date. Further investigation is required to Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 9 of 11 determine whether or not the larger movements shown expanding the duration and area of coverage, the cur- by some of our tagged fish are just roaming movements rent study has a number of implications for design of a over scales greater than our study size or are linked to large-scale tracking array. As a large tracking array can spawning movements. be expensive and time-consuming to install, our data We did not catch any blue-spotted flathead on, or provide guidance to best place passive receivers to cover detect tagged fish blue-spotted flathead moving onto this movement most efficiently. Our results indicate that seagrass or surrounding reef, suggesting that they are using a tightly spaced passive acoustic array for investiga- exclusively soft sediment fish. Our movement data sup - tion of the movement of this species is feasible and would ports findings of recent baited remote underwater video yield meaningful results. However, given the potential (BRUV) studies where no blue-spotted flathead were wider ranging movements of this species, using multiple recorded on reef within Jervis Bay (Rees, Davis and approaches would be useful to provide a more compre- Knott, unpublished data and Coleman et  al. [27]). How- hensive understanding of their movement patterns. At the ever, other BRUV studies have found very small numbers current study site, the entrance to Jervis Bay has now been of blue-spotted flathead on reef habitat; for example in gated and an array of receivers placed around the edge of Batemans Marine Park, Kelaher et al. [28] recorded blue- the bay. These extra receivers (also part of other ongoing spotted flathead on five out of 384 drops over 5 years; this studies) should provide a good idea of visitation to other raises the possibility that blue-spotted flathead occasion - sections of Jervis Bay and also detect if fish leave Jervis Bay. ally venture into edge areas of reef and seagrass habitats or reside there in very low numbers. Conclusions Many studies on the effectiveness or impacts of MPAs Our study, the first to document the movement of blue- have focused on changes in abundances and diversity, spotted flathead, provides clear evidence of short-term without taking into account critical information on site attachment and compact space use by part of the movement patterns of the species within them [2, 29]. blue-spotted flathead population in Jervis Bay. We also This is often because this information is not available or highlight the benefit of using active tracking as a first because while potentially very useful, quantifying the step in understanding the movement of unstudied spe- movement patterns and observing the natural behaviour cies. The area used by tagged fish showing site attach - of marine fish in the field is difficult to achieve. Without ment over a 60-day study period was much smaller than knowledge of the basic movement patterns of a species, no-take sanctuary zones on soft sediments in Jervis Bay it is difficult to predict effectiveness of spatial protection Marine Park. However, our results also suggest that measures such as MPAs [6]. Our study indicates that no- part of the population is also non-resident. While these take sanctuary zones protecting soft sediment habitats results suggest that blue-spotted flathead may respond in JBMP appear large enough to adequately encompass positively to protection provided by the no-take sanctu- the expected short-term movement of blue-spotted flat - ary zones in place, further tracking on a larger number head exhibiting site attachment. However, our data sug- of fish is needed to determine exactly what proportion of gest that two movement patterns are likely to exist within the population shows site attachment and if it continues the population, one that is highly site attached, and thus over the long term. Lastly, our results demonstrate that if would potentially benefit from MPAs, and one that tends we are to effectively manage fish found on soft sediments to roam, and thus may not benefit as much. If these pre - we need to revisit the current view that fish on this habi - liminary data are found to be representative of longer- tat are unlikely to show site attachment. term patterns of movement and activity space use by a large part of the blue-spotted flathead population, then it Additional file is likely that the Hare Bay no-take sanctuary zone is suf- ficiently large to provide protection for a large number of Additional file 1: Table S1. Detailed quantitative summary of move - blue-spotted flathead. If this is the case, we would suggest ment pattern metrics including final h values. that comparably sized zones on soft sediments in other areas of temperate Australia may also be appropriate. Though it is beyond the scope of this study, investigat - Abbreviations KUD: kernel utilisation distribution; JBMP: Jervis Bay Marine Park; MPA: marine ing what portion of the blue-spotted flathead population protected area; RI: residency index; D: displacement; FD: furthest distance; h: would need to show site attachment for spatial closures smoothing parameter; BRUV: baited remote underwater video. like MPAs to be effective will require tagging of much Authors’ contributions larger numbers of fish and deserves further attention. LF designed the study, conducted fieldwork, analysed data, drafted the manu- As this investigation was a preliminary assessment script and created the figures and graphics; AD designed the study, provided for movement of blue-spotted flathead with a view to materials and field resources and assisted in drafting of the manuscript; Fetterplace et al. Anim Biotelemetry (2016) 4:15 Page 10 of 11 NK designed the study, conducted fieldwork, provided materials and field 7. Caveen AJ, Sweeting CJ, Willis TJ, Polunin NVC. Are the scientific founda- resources and assisted in analyses and drafting of the manuscript; JN created tions of temperate marine reserves too warm and hard? Environ Conserv. the figures and graphics; MT provided materials and field resources, assisted 2012;39(3):199–203. in analyses and assisted in drafting of the manuscript. All authors read and 8. Ferguson AM, Harvey ES, Knott NA. Herbivore abundance, site fidelity and approved the final manuscript. grazing rates on temperate reefs inside and outside marine reserves. J Exp Mar Biol Ecol. 2016;478:96–105. Author details 9. Lee KA, Huveneers C, Macdonald T, Harcourt RG. Size isn’t everything: School of Biological Sciences, University of Wollongong, Wollongong, NSW, movements, home range, and habitat preferences of eastern blue Australia. Fish Thinkers Research Group, 11 Riverleigh Avenue, Gerroa, NSW gropers (Achoerodus viridis) demonstrate the efficacy of a small marine 2534, Australia. Fisheries NSW, New South Wales Department of Primary reserve. Aquat Conserv Mar Freshw Ecosyst. 2015;25(2):174–86. Industries, PO Box 5106, Wollongong, NSW, Australia. Port Stephens Fisher- 10. Harasti D, Lee KA, Gallen C, Hughes JM, Stewart J. Movements, home ies Institute, New South Wales Department of Primary Industries, Taylors range and site fidelity of snapper (Chrysophrys auratus) within a temper - Beach Rd, Taylors Beach, NSW, Australia. Jervis Bay Marine Park, New South ate marine protected area. PLoS ONE. 2015;10(11):e0142454. Wales Department of Primary Industries, 4 Woollamia Road, Huskisson, NSW, 11. Willis TJ, Parsons DM, Babcock RC. Evidence for long-term site fidelity of Australia. snapper (Pagrus auratus) within a marine reserve. NZ J Mar Freshw Res. 2001;35(3):581–90. Acknowledgements 12. Wilson WH. 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Animal BiotelemetrySpringer Journals

Published: Jul 28, 2016

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