Adjusting survival estimates for premature transmitter failure: a case study from the Sacramento-San Joaquin Delta

In telemetry studies, premature tag failure causes negative bias in fish survival estimates because tag failure is interpreted as fish mortality. We used mark-recapture modeling to adjust estimates of fish survival for a previous study where premature tag failure was documented. High rates of tag failure occurred during the Vernalis Adaptive Management Plan’s (VAMP) 2008 study to estimate survival of fall-run Chinook salmon (Oncorhynchus tshawytscha) during migration through the San Joaquin River and Sacramento-San Joaquin Delta, California. Due to a high rate of tag failure, the observed travel time distribution was likely negatively biased, resulting in an underestimate of tag survival probability in this study. Consequently, the bias-adjustment method resulted in only a small increase in estimated fish survival when the observed travel time distribution was used to estimate the probability of tag survival. Since the bias-adjustment failed to remove bias, we used historical travel time data and conducted a sensitivity analysis to examine how fish survival might have varied across a range of tag survival probabilities. Our analysis suggested that fish survival estimates were low (95% confidence bounds range from 0.052 to 0.227) over a wide range of plausible tag survival probabilities (0.48–1.00), and this finding is consistent with other studies in this system. When tags fail at a high rate, available methods to adjust for the bias may perform poorly. Our example highlights the importance of evaluating the tag life assumption during survival studies, and presents a simple framework for evaluating adjusted survival estimates when auxiliary travel time data are available.     

An experimental evaluation of the effects of geolocator design and attachment method on between‐year survival on whinchats Saxicola rubetra

Data from location logging tags have revolutionised our understanding of migration ecology, but methods of tagging that do not compromise survival need to be identified. We compared resighting rates for 156 geolocator‐tagged and 316 colour ringed‐only whinchats on their African wintering grounds after migration to and from eastern Europe in two separate years. We experimentally varied both light stalk length (0, 5 and 10 mm) and harness material (elastic or non‐elastic nylon braid tied on, leg‐loop ‘Rappole’ harnesses) in the second year using a reasonably balanced design (all tags in the first year used an elastic harness and 10 mm light stalk). Tags weighed 0.63 g (0.01 SE), representing 4.1% of average body mass. There was no overall significant reduction in between‐year resighting rate (our proxy for survival) comparing tagged and untagged birds in either year. When comparing within tagged birds, however, using a tied harness significantly reduced resighting rate by 53% on average compared to using an elastic harness (in all models), but stalk length effects were not statistically significant in any model considered. There was no strong evidence that the fit (relative tightness) or added tag mass affected survival, although tied tags were fitted more tightly later in the study, and birds fitted with tied tags later may have had lower survival. Overall, on a precautionary principle, deploying tags with non‐elastic tied harnesses should be avoided because the necessary fit, so as not to reduce survival, is time‐consuming to achieve and does not necessarily improve with experience. Geolocator tags of the recommended percentage of body mass fitted with elastic leg‐loop harnesses and with short light stalks can be used without survival effects in small long‐distance migrant birds.     

Survival of Pelagic Juvenile Loggerhead Turtles in the Open Ocean

Abstract: We deployed pop-up archival transmitting tags on 15 loggerhead sea turtles (Caretta caretta) that had been lightly hooked in the United States pelagic longline fishery and on 10 loggerheads that we dip-netted off the surface to serve as controls in the North Atlantic Ocean. We received data from tags of 10 lightly hooked turtles and 7 control turtles. We used data transmitted by the tags in a known-fate model to estimate annual survival rates and determine if there were differences in survival between the 2 groups. The best model indicates there is no difference in survival between the lightly hooked and control turtles, and the estimated annual survival rate was 0.814 (95% CI = 0.557-0.939). Our results suggest that when all fishing gear is removed lightly hooked turtles may not suffer any additional mortality relative to control turtles.     

No effect of passive integrated transponder tagging method on survival or body condition in a northern population of Black‐capped Chickadees (Poecile atricapillus)

Passive integrated transponder (PIT) tags allow a range of individual‐level data to be collected passively and have become a commonly used technology in many avian studies. Although the potential adverse effects of PIT tags have been evaluated in several species, explicit investigations of their impacts on small (<12 g) birds are limited. This is important, because it is reasonable to expect that smaller birds could be impacted more strongly by application of PIT tags. In this study, we individually marked Black‐capped Chickadees (Poecile atricapillus), a small (circa 10 g) passerine, at the University of Alberta Botanic Garden to evaluate potential lethal and sublethal effects of two PIT tagging methods: attachment to leg bands or subcutaneous implantation. We used a Cox proportional hazards model to compare the apparent survival of chickadees with leg band (N = 79) and implanted PIT tags (N = 77) compared with control birds that received no PIT tags (N = 76) over the subsequent 2 years based on mist net recaptures. We used radio‐frequency identification (RFID) redetections of leg band PIT tags to evaluate sex‐specific survival and increase the accuracy of our survival estimates. We also used a generalized linear regression model to compare the body condition of birds recaptured after overwintering with leg band PIT tags, implanted PIT tags, or neither. Our analysis found no evidence for adverse effects of either PIT tagging method on survival or body condition. While we recommend carefully monitoring study animals and evaluating the efficacy of different PIT tagging methods, we have shown that both leg band and subcutaneously implanted PIT tags ethical means of obtaining individualized information in a small passerine.     

Growth, survival, and tag retention of steelhead trout (Oncorhynchus mykiss) and its application to survival estimates

Steelhead trout, Oncorhynchus mykiss, are known to expel acoustic tags which can negatively bias population survival estimates. Tag burden may also affect the development and behavior of smolts, thereby impacting the results of population and behavioral studies. We monitored the growth, condition, and tag expulsion rate of steelhead trout in similar-sized individuals and used these data to adjust survival rates from an acoustic telemetry study conducted in the Sacramento River. Eighty fish were surgically implanted with tags—40 with cylindrical tags of 9 mm diameter and 21 or 24 mm length (V9, Vemco Ltd) and 40 with a 7 mm diameter and 20 mm length tag (V7, Vemco Ltd)—to examine the impact of tag size on peritoneal retention and survival rate of juvenile steelhead trout. A total of 20 % (16/80) of all tags were expelled by smolts during the 143-day study. Ten V9 tags were expelled between day 18 and day 66. Six V7 tags were expelled between day 21 and day 143. A statistical difference was found for retention rate by surgeon even though the surgeons were of equal experience and received the same training. There were no significant differences in the tag retention rate in relation to the tag/body weight ratio, or in growth (weight or fork length) among the control, V7 or V9 treatment groups over the duration of the study. All individuals survived throughout the experiment. Two methods were used to adjust the survival estimates of an acoustic telemetry data set from the Sacramento River based on the tag retention study. First, a simple individual censorship approach in Program MARK was utilized and next ATLAS, a software program designed to compensate for bias in survival estimates caused by tag failures was used. The results of the adjusted survival estimates were not significantly different from the unadjusted rates suggesting that it may be more important to focus on improving surgical techniques to reduce tag expulsion rather than adjusting survival estimates dependent on the study. The surgical techniques utilized in this study did not have significant impacts on the growth rates of either of the tag treatment groups compared to the control. However, tag retention was an issue regardless of the size and weight of the implanted tag and the size of the steelhead.     

Testing assumptions of mark–recapture theory in the coral reef fish Lutjanus apodus

This study tested assumptions of the Cormack–Jolly–Seber capture–mark–recapture (CMR) model in a population of the tropical snapper Lutjanus apodus in the central Bahamas using a combination of laboratory and field studies. The suitability of three different tag types [passive integrated transponder (PIT) tag, T-anchor tag and fluorescent dye jet-injected into the fins] was assessed. PIT tags were retained well, whereas T-anchor tags and jet-injected dye were not. PIT tags had no detectable effect on the rates of growth or survival of individuals. The capture method (fish trapping) was found to provide a representative sample of the population; however, a positive trap response was identified and therefore the assumption of equal capture probability was violated. This study illustrates an approach that can be used to test some of the critical assumptions of the CMR theory and it demonstrates that CMR methods can provide unbiased estimates of growth and mortality of L. apodus provided that trap response is explicitly modelled when estimating survival probability.     

Why do Argos satellite tags stop relaying data?

1. Satellite tracking of animals is very widespread across a range of marine, freshwater, and terrestrial taxa. Despite the high cost of tags and the advantages of long deployments, the reasons why tracking data from tags stop being received are rarely considered, but possibilities include shedding of the tag, damage to the tag (e.g., the aerial), biofouling, battery exhaustion, or animal mortality.
2. We show how information relayed via satellite tags can be used to assess why tracking data stop being received. As a case study to illustrate general approaches that are broadly applicable across taxa, we examined data from Fastloc‐GPS Argos tags deployed between 2012 and 2019 on 78 sea turtles of two species, the green turtle (Chelonia mydas) and the hawksbill turtle (Eretmochelys imbricata).
3. Tags transmitted for a mean of 267 days (SD = 113 days, range: 26–687 days, median = 251 days). In 68 of 78 (87%) cases, battery failure was implicated as the reason why tracking data stopped being received. Some biofouling of the saltwater switches, which synchronize transmissions with surfacing, was evident in a few tags but never appeared to be the reason that data reception ceased.
4. Objectively assessing why tags fail will direct improvements to tag design, setup, and deployment regardless of the study taxa. Assessing why satellite tags stop transmitting will also inform on the fate of tagged animals, for example, whether they are alive or dead at the end of the study, which may allow improved estimates of survival rates.

     

Differential survival of male and female partially resistant horn flies (Diptera: Muscidae) on steers treated with permethrin

Males and females from a heterozygous, resistant strain (SR) of the horn fly, Haematobia irritans (L.), were tested for survival until reproductive maturity on steers with ear tags in outdoor, screened enclosures and on steers sprayed on the neck in an indoor isolation room. After 6 d, female SR flies on outdoor steers with one tag had 10 times greater survival than males; almost no SR flies on steers with two ear tags survived. Survival of male and female SR flies on steers sprayed on the neck was reduced during the first 24 h, but not thereafter. Lower survival of males compared with females on treated steers reflected differential survival of the sexes during exposure to treated cloths in a laboratory bioassay. Hair samples from neck, back, rump, and lower legs of steers with ear tags in outdoor pens were tested for toxicity to the SR flies. These bioassays indicated high localization of insecticide on the neck of steers with ear tags.     

Detection and apparent survival of PIT‐tagged stream fish in winter

Environmental fluctuations exert strong control on behavior, survival, and fitness of stream biota. Technical improvements increasingly allow for tracking the response of large numbers of individuals to environmental fluctuations, for instance, by remote detection of animals equipped with PIT (passive integrated transponder) tags. PIT tags were implanted into 393 juvenile and adult brown trout Salmo trutta L. and European sculpin Cottus gobio L. in a boreal stream subjected to considerable ice formation. With weekly trackings over 6 months, we quantified apparent survival and detection probability in relation to biological, environmental, and methodological factors. Individuals with a higher physical condition in autumn showed a higher apparent survival; this pattern was consistent across all species and age classes. Detection probability decreased with increasing thickness of the surface ice layer; this effect was most pronounced for juvenile trout and benthic‐living sculpin, both tagged with smaller‐sized tags. Detection probability was reduced in structurally complex habitats. Our study demonstrates that apparent survival and particularly detection probability may show pronounced spatiotemporal variation. In order to compare results from different sampling occasions and sites, a good knowledge of the study site and of the regulating factors is crucial.     

Tag retention and mortality of adult Atlantic salmon Salmo salar gastrically tagged with different sized telemetry transmitters

A batch of 1 sea winter pre‐spawning adult Salmo salar from the Bush river in Northern Ireland, U.K., were gastrically tagged with large (13 mm diameter) and small (9 mm diameter) dummy acoustic telemetry tags alongside untagged control fish. Survival differed between control and tagged fish and the estimated probability of survival by the end of the study for control fish with no tag was 0·94, small tags was 0·90 and large tags was 0·72. Tag loss through regurgitation was slightly higher for fish tagged with larger tags than for fish tagged with smaller tags and the estimated probability of tag loss for fish with a small tag was 0·10 and for large tags was 0·14.     

Effects of tag type on red snapper Lutjanus campechanus tag retention,growth and survival under hatchery conditions

Management of red snapper stocks is one of the most controversial issues in the Gulf of Mexico fisheries. Fish tagging programs can help to address many management issues if an appropriate tag is selected. In the study three types of tags were tested to determine tag retention and effects on red snapper growth and survival. Hatchery‐produced juveniles (43.7 ± 5.6 g) were either not tagged or given: internal anchor tags anchored in the abdominal cavity; passive integrated transponders (PIT) tags injected into the body cavity; or metal self‐piercing fish tags clamped around the dentary bone. Fish were stocked at 25 m^?3 in three replicate circular tanks/tag type, and fed a commercial feed for a 150 days period. Fish survival did not differ by treatment. Average weight gains after 150 days were similar. Tag retention differed by treatment (P = 0.0002), with 100% retention of PIT tags, 98.4 ± 2.8% retention of jaw tags, and 90.4 ± 1.6% retention of internal anchor tags. Loss of internal anchor tags occurred primarily within the first month. Inflammation was frequently observed at the internal anchor tag insertion site. Irritation and cell proliferation was sometimes observed in areas around the jaw tags.     

Survival of the African white‐backed vulture Gyps africanus in north‐eastern South Africa

Old World vultures are in decline across their entire range. Although critical for the formulation of effective conservation measures, neither survival nor movement patterns of African vultures are adequately known. This paper presents survival and movement data on the African white‐backed vultures (Gyps africanus) from South Africa. Survival estimates were modelled on resightings of tagged vultures. Birds were captured en masse and resighted between November 2005 and December 2010. A total of 93 adult and subadult birds were fitted with uniquely numbered patagial tags, which were resighted 3707 times(mean of 39.8 resightings per bird). The programme MARK was used to estimate survival. The best model was one where survival and recaptures varied only with time (e.g. year). However, owing to the fading (illegibility) of tags in later years, the relationship with time is probably spurious. The second best model was one where survival and recaptures varied with age and time. Annual survival estimates increased from 85.2% in second‐year birds to 99.9% in adults. This corresponds well with the survival of two other Gyps vultures that have been studied to date and underscores the point that additional mortality of adults in these long‐lived species will result in rapid population declines.     

A synthesis of tagging studies examining the behaviour and survival of anadromous salmonids in marine environments

This paper synthesizes tagging studies to highlight the current state of knowledge concerning the behaviour and survival of anadromous salmonids in the marine environment. Scientific literature was reviewed to quantify the number and type of studies that have investigated behaviour and survival of anadromous forms of Pacific salmon (Oncorhynchus spp.), Atlantic salmon (Salmo salar), brown trout (Salmo trutta), steelhead (Oncorhynchus mykiss), and cutthroat trout (Oncorhynchus clarkii). We examined three categories of tags including electronic (e.g. acoustic, radio, archival), passive (e.g. external marks, Carlin, coded wire, passive integrated transponder [PIT]), and biological (e.g. otolith, genetic, scale, parasites). Based on 207 papers, survival rates and behaviour in marine environments were found to be extremely variable spatially and temporally, with some of the most influential factors being temperature, population, physiological state, and fish size. Salmonids at all life stages were consistently found to swim at an average speed of approximately one body length per second, which likely corresponds with the speed at which transport costs are minimal. We found that there is relatively little research conducted on open-ocean migrating salmonids, and some species (e.g. masu [O. masou] and amago [O. rhodurus]) are underrepresented in the literature. The most common forms of tagging used across life stages were various forms of external tags, coded wire tags, and acoustic tags, however, the majority of studies did not measure tagging/handling effects on the fish, tag loss/failure, or tag detection probabilities when estimating survival. Through the interdisciplinary application of existing and novel technologies, future research examining the behaviour and survival of anadromous salmonids could incorporate important drivers such as oceanography, tagging/handling effects, predation, and physiology.     

Advances in tagging syngnathids, with the effects of dummy tags on behaviour of Hippocampus guttulatus

Artificial marking and tagging techniques have been used to study movement, population dynamics, behaviour, ecology, survival and growth of at least 25 syngnathid species. External necklace-style tags and injection of visible implant elastomer have been the most used techniques, uniquely identifying hundreds of individual syngnathids to study population dynamics, mortality, behaviour, ecology and growth in at least 13 and 12 species, respectively. Only two studies, both on larger syngnathid species, have tested the use of internal or electronic tags. This new case study reveals that dummy tags, weighing up to 6% of individual body mass, have minimal effect on normal ex situ behaviour of the long-snouted seahorse Hippocampus guttulatus, a smaller syngnathid. In paired aquarium trials, tags did not affect movement, holdfast use or general behavioural state, and only had a short-term effect (1 day) on vertical orientation. Tagged H. guttulatus gained more mass during the 5 day trials, a result which warrants further exploration but indicates that tags did not reduce feeding. This study shows promise for using electronic tagging to study H. guttulatus and similarly sized syngnathids in the wild.     

External and internal tags for the green sea urchin

Two internal and three external tags were tested on the green sea urchin. Desirable qualities of a tag were high sea urchin survival, retention for at least a few months, detection on the sea floor by divers, identification of individuals, quick application, and low cost. These objectives were met by an external nylon screw tag visible to divers and two internal aluminum tags detectable with an underwater metal detector. Successful tags were inserted through a hole drilled in the test and were tested in the laboratory and field. All internal tags were retained for the full duration of the 4-month trial and did not retard growth or affect survival. Divers could identify individual urchins with nylon screw tags, but the tag retention rate was lower.     

A Model for Censored Survival Data When There is an Intervening Event

A model is discussed for incorporating information from a time-dependent covariable (an intervening event) and covariables independent of time into the analysis of survival data. In the model, it is assumed that individuals are potentially subject to two paths to failure, one including the intervening event and the other not. Additional assumptions are that failure times associated with the two paths are independent and that the time to failure subsequent to the intervening event is dependent on the intervening event time. Allowing the underlying hazard rates for the model to follow a WEIBULL form, use of the model and methods for fitting and hypothesis testing are illustrated by application to a follow-up study involving industrial workers where disability retirement was the intervening event. Extensions of the model to accommodate grouped survival data are presented.     

A method adapting microarray technology for signature-tagged mutagenesis of Desulfovibrio desulfuricans G20 and Shewanella oneidensis MR-1 in anaerobic sediment survival experiments

Signature-tagged mutagenesis (STM) is a powerful technique that can be used to identify genes expressed by bacteria during exposure to conditions in their natural environments. To date, there have been no reports of studies in which this approach was used to study organisms of environmental, rather than pathogenic, significance. We used a mini-Tn10 transposon-bearing plasmid, pBSL180, that efficiently and randomly mutagenized Desulfovibrio desulfuricans G20 in addition to Shewanella oneidensis MR-1. Using these organisms as model sediment-dwelling anaerobic bacteria, we developed a new screening system, modified from former STM procedures, to identify genes that are critical for sediment survival. The screening system uses microarray technology to visualize tags from input and output pools, allowing us to identify those lost during sediment incubations. While the majority of data on survival genes identified will be presented in future papers, we report here on chemotaxis-related genes identified by our STM method in both bacteria in order to validate our method. This system may be applicable to the study of numerous environmental bacteria, allowing us to identify functions and roles of survival genes in various habitats.     

A Field Evaluation of an External and Neutrally Buoyant Acoustic Transmitter for Juvenile Salmon: Implications for Estimating Hydroturbine Passage Survival

Turbine-passed fish are exposed to rapid decreases in pressure which can cause barotrauma. The presence of an implanted telemetry tag increases the likelihood of injury or death from exposure to pressure changes, thus potentially biasing studies evaluating survival of turbine-passed fish. Therefore, a neutrally buoyant externally attached tag was developed to eliminate this bias in turbine passage studies. This new tag was designed not to add excess mass in water or take up space in the coelom, having an effective tag burden of zero with the goal of reducing pressure related biases to turbine survival studies. To determine if this new tag affects fish performance or susceptibility to predation, it was evaluated in the field relative to internally implanted acoustic transmitters (JSATS; Juvenile Salmon Acoustic Telemetry System) used widely for survival studies of juvenile salmonids. Survival and travel time through the study reach was compared between fish with either tag type in an area of high predation in the Snake and Columbia rivers, Washington. An additional group of fish affixed with neutrally-buoyant dummy external tags were implanted with passive integrated transponder (PIT) tags and recovered further downstream to assess external tag retention and injury. There were no significant differences in survival to the first detection site, 12 river kilometers (rkm) downstream of release. Travel times were also similar between groups. Conversely, externally-tagged fish had reduced survival (or elevated tag loss) to the second detection site, 65 rkm downstream. In addition, the retention study revealed that tag loss was first observed in fish recaptured approximately 9 days after release. Results suggest that this new tag may be viable for short term (<8 days) single-dam turbine-passage studies and under these situations, may alleviate the turbine passage-related bias encountered when using internal tags, however further research is needed to confirm this.     

Growth,survival, and tag retention of steelhead trout (<Emphasis Type="Italic">O. mykiss</Emphasis>) surgically implanted with dummy acoustic tags

Recent advances in micro-electronics make the study of the migration of even small marine animals (>12 cm) over many 1000s of kilometres a serious possibility. Important assumptions in long-term studies are that rates of tag loss caused by mortality or tag shedding are low, and that the tagging procedure does not have an unacceptable negative effect on the animal. This paper reports results from a study to examine the retention of relatively large (24 × 8 mm) surgically-implanted dummy acoustic tags over a 7-month period in steelhead pre-smolts (O. mykiss), and the effects of implantation on growth and survival. Although there was some influence on growth to week 12, survival was high for animals > 13 cm FL. In the following 16-week period, growth of surgically implanted pre-smolts was the same as the control population and there was little tag loss from mortality or shedding. Currently available acoustic tags can be implanted in salmonid fish ≥12 cm FL, although combined losses from mortality and tag shedding were 33–40% for animals in the 12 and 13 cm FL size classes. By 14 cm FL, combined rates of tag loss (mortality plus tag shedding) for surgically implanted tags dropped to <15% and growth following surgery was close to that of the controls. Our results suggest that studies of ocean migration and survival over periods of many months are now feasible even for animals as small as salmon smolts. Surgically implanted salmon smolts are therefore good candidates for freshwater and coastal ocean-tracking studies on relatively long time scales (months). On such time scales, even relatively small salmon smolts may move thousands of kilometers in the ocean.     

Long‐term survival of humpback whales radio‐tagged in Alaska from 1976 through 1978

Invasive tags designed to provide information on animal movements through radio or satellite monitoring have tremendous potential for the study of whales and other cetaceans. However, to date there have been no published studies on the survival of tagged animals over periods of years or decades. Researchers from the National Marine Mammal Laboratory and the Woods Hole Oceanographic Institution tracked five humpback whales with implanted radio tags in southeastern Alaska in August 1976 and July 1977, and tracked two humpback whales in Prince William Sound, Alaska, in June 1978. All seven of these individually identified humpback whales were resighted at least 20 yr after first being tagged, and five of the seven have been observed for more than 30 yr; some of them are among the most resighted humpback whales in the North Pacific. Photos of tagging sites taken during and subsequent to tagging operations show persistent but superficial scarring and no indication of infection. These pioneering field studies demonstrated both long‐term survival of the whales and the short‐term effects of deploying radio tags, which at the time were larger and more invasive than those typically used today.