Biases associated with the use of underwater visual census techniques to quantify the density and size-structure of fish populations

Estimates of reef fish densities made by divers visually censusing 5 m wide strip transects were compared with capture-resight estimates calculated independently using data on the resighting frequencies of fish marked with colour-coded tags. The difference in density estimates between methods varied between species but with patterns consistent at the three eastern Tasmanian sites studied. Densities of the two most abundant species, the wrasses Notolabrus tetricus and Notolabrus fucicola, showed good agreement between methods. By contrast, populations of the two monacanthids Meuschenia australis and Meuschenia freycineti were underestimated by an order of magnitude in strip transects relative to capture-resight, while populations of the open-water latrid Latridopsis forsteri were overestimated.For all common fish species observed in strip transects, variation in density estimates between divers was extremely low compared to variation between sites and between months. Variation in density estimates between different days within a week and between 50 m blocks located 50 m apart was also negligible; however, residual error associated with variation by a single diver within a block and day was high. Biases associated with changing detectability of fishes in open versus densely vegetated habitats were assessed by algal removal experiments, and were found to be negligible or non-significantly low for five of the six species examined. The exceptional species—N. tetricus—exhibited a 70% rise in fish sighted by diver following algal clearance despite trapping data indicating no increase in fish numbers.Diver estimates of fish length were on average 7% greater than measured lengths. Divers possessed a clear tendency to make increasingly inaccurate size estimates as fish length deviated in either direction from 300 mm. Lengths of 175-mm animals were underestimated by ≈20% and 400-mm fishes were overestimated by ≈10%. These changes in diver bias with fish size were largely independent of fish species, site and diver. The precision of size estimates, as indicated by the standard deviation of bias, also varied with fish size, with values varying from ≈13% at 200 mm fish length to ≈8% at 400 mm length. The decline in precision at small body size largely reflected size intervals used by divers to bin data. Divers appear capable of making more precise size estimates than the 25-mm interval used at small fish sizes.     

Improvement of fish length estimates for underwater visual census of reef fish biomass

Accuracy and precision are of great importance in the assessment of reef fish biomass when conducting an underwater visual census (UVC). Quantification and subsequent correction of the bias is required in order to standardize the estimates and correct for underwater distortion. To optimize the UVC, the observer should receive length‐measurement training in order to obtain in situ‐measurements that are as accurate and precise as possible. The objective of this study was to quantify the bias of fish length measurements made by divers with and without training in order to enhance reef fish biomass estimates. Adaptation of the diver to estimate fish lengths was analysed as a part of reef fish biomass monitoring in the Karimunjawa National Park, a national marine sanctuary in the Java Sea, Indonesia. Two divers practiced estimating a variety of fish in a natural environment by using styrofoam models attached to strings and sinkers. Analyses showed that by training the diver, his/her accuracy and precision improved substantially. Proving its reliability, an underwater visual census (UVC) becomes a useful and reliable method to assess the biomass of reef fishes.     

Sustainable dive ecotourism on a South African coral reef

SCUBA ecotourism provides economic support for local human communities and for conservation. Such support should be sustainable and as ecologically non-destructive as possible to its coral reef foundation. In recent years, the major increase in SCUBA activity has become a significant consideration at some locations. Among these are the South African coral reefs at Sodwana Bay, which receive 80 000 dives per year. Between May 1998 and January 1999, assessments using line-intercept and belt transects showed that 5.9% of hard corals and 4.0% of soft corals were damaged from various causes, while only 0.02% of the hard corals were dead. Of the 222 diver contacts observed, only 1.63% produced instant and visually-clear damage, with no measurable decline in overall coral reef health. Direct observations on 89 recreational divers showed that photographers made damaging contacts on nine out of ten dives, while novice divers made one such contact per six dives, moderately experienced divers about once every 14 dives, and very experienced divers about once every 123 dives. Increased time lapsed since the last dive increased diver damage by novices. We calculated that the 80 000 dives were equivalent to 1 m² of reef receiving 2.3 diver contacts per year. Acropora austera was by far the most significantly damaged coral, with one out of four branches broken per year. But as this species accounted for only 0.2% of reef benthos cover, this impact was not representative of overall reef health. This heavy species-specific impact makes A. austera a good bio-monitor.     

Variability in Abundance of Temperate Reef Fishes Estimated by Visual Census

Identifying sources of sampling variation and quantifying their magnitude is critical to the interpretation of ecological field data. Yet, most monitoring programs of reef fish populations based on underwater visual censuses (UVC) consider only a few of the factors that may influence fish counts, such as the diver or census methodology. Recent studies, however, have drawn attention to a broader range of processes that introduce variability at different temporal scales. This study analyzes the magnitude of different sources of variation in UVCs of temperate reef fishes off Patagonia (Argentina). The variability associated with time-of-day, tidal state, and time elapsed between censuses (minutes, days, weeks and months) was quantified for censuses conducted on the five most conspicuous and common species: Pinguipes brasilianus, Pseudopercis semifasciata, Sebastes oculatus, Acanthistius patachonicus and Nemadactylus bergi. Variance components corresponding to spatial heterogeneity and to the different temporal scales were estimated using nested random models. The levels of variability estimated for the different species were related to their life history attributes and behavior. Neither time-of-day nor tidal state had a significant effect on counts, except for the influence of tide on P. brasilianus. Spatial heterogeneity was the dominant source of variance in all but one species. Among the temporal scales, the intra-annual variation was the highest component for most species due to marked seasonal fluctuations in abundance, followed by the weekly and the instantaneous variation; the daily component was not significant. The variability between censuses conducted at different tidal levels and time-of-day was similar in magnitude to the instantaneous variation, reinforcing the conclusion that stochastic variation at very short time scales is non-negligible and should be taken into account in the design of monitoring programs and experiments. The present study provides baseline information to design and interpret results from visual census programs in temperate reefs.     

Habitat complexity and fish size affect the detection of Indo-Pacific lionfish on invaded coral reefs

A standard approach to improving the accuracy of reef fish population estimates derived from underwater visual censuses (UVCs) is the application of species-specific correction factors, which assumes that a species’ detectability is constant under all conditions. To test this assumption, we quantified detection rates for invasive Indo-Pacific lionfish (Pterois volitans and P. miles), which are now a primary threat to coral reef conservation throughout the Caribbean. Estimates of lionfish population density and distribution, which are essential for managing the invasion, are currently obtained through standard UVCs. Using two conventional UVC methods, the belt transect and stationary visual census (SVC), we assessed how lionfish detection rates vary with lionfish body size and habitat complexity (measured as rugosity) on invaded continuous and patch reefs off Cape Eleuthera, the Bahamas. Belt transect and SVC surveys performed equally poorly, with both methods failing to detect the presence of lionfish in >50 % of surveys where thorough, lionfish-focussed searches yielded one or more individuals. Conventional methods underestimated lionfish biomass by ~200 %. Crucially, detection rate varied significantly with both lionfish size and reef rugosity, indicating that the application of a single correction factor across habitats and stages of invasion is unlikely to accurately characterize local populations. Applying variable correction factors that account for site-specific lionfish size and rugosity to conventional survey data increased estimates of lionfish biomass, but these remained significantly lower than actual biomass. To increase the accuracy and reliability of estimates of lionfish density and distribution, monitoring programs should use detailed area searches rather than standard visual survey methods. Our study highlights the importance of accounting for sources of spatial and temporal variation in detection to increase the accuracy of survey data from coral reef systems.     

Do Reef Fish Habituate to Diver Presence? Evidence from Two Reef Sites with Contrasting Historical Levels of SCUBA Intensity in the Bay Islands,Honduras

Contact between humans and the marine environment is increasing, but the capacity of communities to adapt to human presence remains largely unknown. The popularization of SCUBA diving has added a new dimension to human impacts in aquatic systems and, although individual-level impacts have been identified, cumulative effects on ecosystem function and community-wide responses are unclear. In principle, habituation may mitigate the consequences of human presence on the biology of an individual and allow the quick resumption of its ecological roles, but this has not been documented in aquatic systems. Here, we investigate the short-term impact of human presence and the long-term habituation potential of reef-fish communities to recreational SCUBA divers by studying symbiotic cleaning interactions on coral reefs with differing levels of historical contact with divers. We show that incidences of human contact result in a smaller decline in ecosystem function and more rapid resumption of baseline services on a reef in Utila, Honduras that has heavy historical levels of SCUBA diver presence, compared to an un-dived reef site in the Cayos Cochinos Marine Protected Area (CCMPA). Nonetheless, despite the generally smaller change in ecosystem function and decades of regular contact with divers, cleaning behavior is suppressed by >50% at Utila when divers are present. We hypothesize that community-wide habituation of reef fish is not fully achievable and may be biologically restricted to only partial habituation. Differential responses to human presence impacts the interpretation and execution of behavioral research where SCUBA is the predominant means of data collection, and provides an important rationale for future research investigating the interplay between human presence, ecosystem function, and community structure on coral reefs.     

Behaviour of temperate and sub-tropical reef fishes towards a stationary SCUBA diver

Using remote underwater stereo-video systems we examined fish behaviour towards a stationary SCUBA diver at temperate (Rottnest Island) and sub-tropical (Houtman Abrolhos Islands) reefs in Western Australia. Changes in species richness, relative abundance, fish length, and the mean distance of individual fish from stereo-video cameras, in the presence and absence of a SCUBA diver, were assessed to infer changes in behaviour. Results show that a stationary SCUBA diver may obtain accurate measures of species richness and of the composition of fish assemblages in an area. However, the usefulness of these measures to reflect changes in fish behaviour appears limited as responses of fish towards the stationary SCUBA diver were highly species specific. Several species differed in their mean relative abundance (Heterodontus portusjacksoni, Coris auricularis, Thalassoma lunare), length (Ophthalmolepsis lineolatus, C. auricularis), and the distance to which they would approach the stereo-video systems (Kyphosus sydneyanus, Scarus schlegeli) when a SCUBA diver was present. Here, species-specific changes in the behaviour of several common and abundant fish species towards a stationary SCUBA diver advises caution to avoid biases when interpreting results obtained by SCUBA divers.     

Behaviour of temperate and sub-tropical reef fishes towards a stationary SCUBA diver

Using remote underwater stereo–video systems we examined fish behaviour towards a stationary SCUBA diver at temperate (Rottnest Island) and sub-tropical (Houtman Abrolhos Islands) reefs in Western Australia. Changes in species richness, relative abundance, fish length, and the mean distance of individual fish from stereo–video cameras, in the presence and absence of a SCUBA diver, were assessed to infer changes in behaviour. Results show that a stationary SCUBA diver may obtain accurate measures of species richness and of the composition of fish assemblages in an area. However, the usefulness of these measures to reflect changes in fish behaviour appears limited as responses of fish towards the stationary SCUBA diver were highly species specific. Several species differed in their mean relative abundance (Heterodontus portusjacksoni, Coris auricularis, Thalassoma lunare), length (Ophthalmolepsis lineolatus, C. auricularis), and the distance to which they would approach the stereo–video systems (Kyphosus sydneyanus, Scarus schlegeli) when a SCUBA diver was present. Here, species-specific changes in the behaviour of several common and abundant fish species towards a stationary SCUBA diver advises caution to avoid biases when interpreting results obtained by SCUBA divers.     

Human,Oceanographic and Habitat Drivers of Central and Western Pacific Coral Reef Fish Assemblages

Coral reefs around US- and US-affiliated Pacific islands and atolls span wide oceanographic gradients and levels of human impact. Here we examine the relative influence of these factors on coral reef fish biomass, using data from a consistent large-scale ecosystem monitoring program conducted by scientific divers over the course of >2,000 hours of underwater observation at 1,934 sites, across ~40 islands and atolls. Consistent with previous smaller-scale studies, our results show sharp declines in reef fish biomass at relatively low human population density, followed by more gradual declines as human population density increased further. Adjusting for other factors, the highest levels of oceanic productivity among our study locations were associated with more than double the biomass of reef fishes (including ~4 times the biomass of planktivores and piscivores) compared to islands with lowest oceanic productivity. Our results emphasize that coral reef areas do not all have equal ability to sustain large reef fish stocks, and that what is natural varies significantly amongst locations. Comparisons of biomass estimates derived from visual surveys with predicted biomass in the absence of humans indicated that total reef fish biomass was depleted by 61% to 69% at populated islands in the Mariana Archipelago; by 20% to 78% in the Main Hawaiian islands; and by 21% to 56% in American Samoa.     

Estimating length frequency distributions of large reef fish underwater

We describe the training of divers to recognise and remove bias in estimating lengths of fish underwater. Divers were asked to allocate objects, from a population (N=50) with a known length frequency distribution, to ten 100 mm size classes. Observed and expected distributions were then compared and the divers informed of their errors. Training continued until divers consistently produced length frequency distributions that were not significantly different from the expected distribution (=0.8) by the one sample Kolmogorov-Smirnov (K-S) test. Divers were trained in five trials, but after six months they had lost all their ability and had to be retrained. Three trained divers observing the same population of the large reef fish Plectropomus leopardus (Serranidae) produced length frequency distributions that were not significantly different (P>0.1) on 67% of occasions. Data collected by divers can be used to detect small but real differences in length frequency distributions of populations when analysed using the two sample K-S test. We suggest a means of determining within site variation in length frequency relative to between site variation.     

Use of sonar transects to improve efficiency and reduce potential bias in visual surveys of reef fishes

A sonar transect method has been found to be a useful alternative to traditional tape-transect methods in quantification of the species richness and abundance of reef fish. No significant difference was found between species richness measured by the methods, and high Jaccard’s similarity coefficients were calculated for each site using the two methods, indicating a similar response to fish of both the sonar and tape transects. Abundance estimates of one species only (Chaetodon tricinctus) were greater when the transect tape method was used, probably because fish were being attracted to the divers who deployed the transect tape. Substantial time saving was also noted for the sonar transects, indicating overall that estimating distance using sonar rather than transect tape may enable increased efficiency and accuracy in␣underwater visual surveys for both ecological␣and management-related research on reef fishes.     

Evaluating the use of roving diver and transect surveys to assess the coral reef fish assemblage off southeastern Hispaniola

The relatively little-studied fish fauna off southeastern Hispaniola was rapidly assessed using a combination of visual survey techniques including transects and roving diver surveys. It was found that when combined, both methods provided a more complete overall species assessment than either method was able to provide in isolation. Being able to conduct rapid species assessments is becoming increasingly more important as a conservation tool. Data on species composition, sighting frequency, and abundance of all fishes were collected using both methods. Abundance was recorded in four logarithmic-based categories (roving diver method) while the number of fishes were counted within 40-m² transects (transect method). Both methods were similar in recording the most abundant species, while a greater number of rare species (especially fishery-targeted species) were recorded with the roving diver method. The most abundant groupers were Cephalopholis cruentata and C. fulva. The most abundant parrotfishes were Scarus taeniopterus, Sparisoma aurofrenatum, and Scarus iserti. The most conspicuous differences between fishes off southeastern Hispaniola and elsewhere in the tropical western Atlantic were the low abundance and smaller size of harvested species such as groupers, snappers (Fam. Lutjanidae), and grunts (Fam. Haemulidae). With the roving diver method, more time could be spent surveying (instead of placing transect lines), resulting in a greater number of species being recorded. Additionally, well-trained volunteers can adopt the roving diver method as part of their regular diving program. Transect surveys were able to provide information such as length (biomass) and actual density measures that were not recorded in roving diver surveys. Thus, these methods were complementary and should be used in conjunction when conducting rapid assessments of fish assemblages, especially to detect the effects of overfishing.     

Comparison of efficiency of direct observations by scuba diver and indirect observations via video camera for measuring reef-fish behaviour

The present study investigates how the humbug damselfish Dascyllus aruanus, subject of a large number of ecological, evolutionary and behavioural studies, responds to the presence of human observers (effect of scuba diver presence-absence) and how the method of data collection (directly by a scuba diver v. indirectly via video camera) may affect the quality of behavioural data. Scuba diver presence had only subtle effects on fish behaviour. The efficiency of the method of scoring fish behaviour depended on the behaviour under consideration: those behaviours that occur in close proximity to the corals were scored more effectively directly by a scuba diver while those that are performed in a more rapid or repetitive fashion were scored more effectively indirectly via video camera. These results provide a foundation for future behavioural research on D. aruanus and other fishes where scuba divers or video cameras are the prevalent means of data collection.     

An analysis of fish community responses to coral mining in the Maldives

Synopsis Coral mining takes place on shallow reef flats at a number of localities in the Maldives, but not on the adjacent deeper reef slopes. A semi-quantitative census method for fish species abundance and biomass is described. Fish community structure is compared on mined and non-mined reef flats and their adjacent slopes using a variety of univariate, graphical/distributional and multivariate statistical techniques. In general, univariate and graphical distributional methods do not indicate significant differences between mined and non-mined localities with respect to the relative abundances and biomasses of species. Multivariate methods (both classification and ordination), however, indicate very clear-cut effects of mining on the reef flats, and also significant effects on reef slopes adjacent to mined flats. The effect was equally clear using non-quantitative (presence/absence) data. The fish species mainly responsible for the differences between mined and non-mined localities are identified, and the differences are explained in terms of the feeding biology of these species.     

A preliminary examination of depth associated spatial variation in fish assemblages on small artificial reefs

Small concrete artificial reef modules (hemisphere-shaped, approximately 1.3 m diameter, 1 m high) were placed at two sites, eight modules per site. The sites were in 7 m and 21 m of water, 1.6 km apart, off the coast of southeast Florida, USA. The reefs were censused monthly for fish over a 19-month period. Species, number of fish, and estimated total length of each individual were recorded by divers using SCUBA. After the monthly census, all fish were removed from the reefs with a piscicide. A total of 88 species were recorded in the study, with significantly greater diversity on the deep reefs (monthly mean of 7.6 versus 3.0 shallow, P < 0.001, anova ). There were also significantly more biomass (calculated from length), and more large fish (>5 cm) on the reefs at 21 m than at 7 m (P < 0.001). There were more small fish at the shallow site (P < 0.05). Although it is not clear what variable(s) associated with the two depths is responsible for the differences, these results highlight the potential differences in artificial reef and ambient environment interactions within a localized area.     

Acoustic Telemetry Validates a Citizen Science Approach for Monitoring Sharks on Coral Reefs

Citizen science is promoted as a simple and cost-effective alternative to traditional approaches for the monitoring of populations of marine megafauna. However, the reliability of datasets collected by these initiatives often remains poorly quantified. We compared datasets of shark counts collected by professional dive guides with acoustic telemetry data from tagged sharks collected at the same coral reef sites over a period of five years. There was a strong correlation between the number of grey reef sharks (Carcharhinus amblyrhynchos) observed by dive guides and the telemetry data at both daily and monthly intervals, suggesting that variation in relative abundance of sharks was detectable in datasets collected by dive guides in a similar manner to data derived from telemetry at these time scales. There was no correlation between the number or mean depth of sharks recorded by telemetry and the presence of tourist divers, suggesting that the behaviour of sharks was not affected by the presence of divers during our study. Data recorded by dive guides showed that current strength and temperature were important drivers of the relative abundance of sharks at monitored sites. Our study validates the use of datasets of shark abundance collected by professional dive guides in frequently-visited dive sites in Palau, and supports the participation of experienced recreational divers as contributors to long-term monitoring programs of shark populations.     

Comparisons among survey methodologies to test for abundance and size of a highly targeted fish species

Three sampling methods for estimating abundance and size of blue cod Parapercis colias were compared inside and outside Kapiti Marine Reserve, New Zealand (40° 49′ 31·77^′′ S; 174° 55′ 02·87^′′ E). Two baited methods, baited underwater video (BUV) and experimental angling (EA), were more efficient and had lower levels of estimate variation than diver‐based underwater visual census (UVC). The BUV and EA recorded more fish and of greater size ranges than UVC, and also had fewer zero count replicates. The BUV and EA methodologies revealed highly significant differences in abundance and size of fish between sites (reserve v. non‐reserve), whereas UVC revealed no such differences. These results indicate that BUV is likely to be the most accurate, cost‐effective and easy to use methodology for the surveying of carnivorous temperate reef fishes for future monitoring. It is noted, however, that new data acquired using the BUV methodology may need to be compared over a calibration period to data acquired using the UVC methodology to ensure that historical data sets derived from UVC still have validity and application for future monitoring activity.     

Geography and island geomorphology shape fish assemblage structure on isolated coral reef systems

We quantify the relative importance of multi‐scale drivers of reef fish assemblage structure on isolated coral reefs at the intersection of the Indian and Indo‐Pacific biogeographical provinces. Large (>30 cm), functionally‐important and commonly targeted species of fish, were surveyed on the outer reef crest/front at 38 coral reef sites spread across three oceanic coral reef systems (i.e. Christmas Island, Cocos (Keeling) Islands and the Rowley Shoals), in the tropical Indian Ocean (c. 1.126 x 106 km²). The effects of coral cover, exposure, fishing pressure, lagoon size and geographical context, on observed patterns of fish assemblage structure were modelled using Multivariate Regression Trees. Reef fish assemblages were clearly separated in space with geographical location explaining ~53 % of the observed variation. Lagoon size, within each isolated reef system was an equally effective proxy for explaining fish assemblage structure. Among local‐scale variables, ‘distance from port’, a proxy for the influence of fishing, explained 5.2% of total variation and separated the four most isolated reefs from Cocos (Keeling) Island, from reefs with closer boating access. Other factors were not significant. Major divisions in assemblage structure were driven by sister taxa that displayed little geographical overlap between reef systems and low abundances of several species on Christmas Island corresponding to small lagoon habitats. Exclusion of geographical context from the analysis resulted in local processes explaining 47.3% of the variation, highlighting the importance of controlling for spatial correlation to understand the drivers of fish assemblage structure. Our results suggest reef fish assemblage structure on remote coral reef systems in the tropical eastern Indian Ocean reflects a biogeographical legacy of isolation between Indian and Pacific fish faunas and geomorphological variation within the region, more than local fishing pressure or reef condition. Our findings re‐emphasise the importance that historical processes play in structuring contemporary biotic communities.     

Drivers of reef fish assemblages in an upwelling region from the Eastern Tropical Pacific Ocean

Reef fish assemblages are exposed to a wide range of anthropogenic threats as well as chronic natural disturbances. In upwelling regions, for example, there is a seasonal influx of cool nutrient-rich waters that may shape the structure and composition of reef fish assemblages. Given that climate change may disrupt the natural oceanographic processes by altering the frequency and strength of natural disturbances, understanding how fish assemblages respond to upwelling events is essential to effectively manage reef ecosystems under changing ocean conditions. This study used the baited remote underwater video stations (BRUVS) and the traditional underwater visual census (UVC) to investigate the spatiotemporal patterns of reef fish assemblages in an upwelling region in the North Pacific of Costa Rica. A total of 183 reef fish species from 60 families were recorded, of which 166 species were detected using BRUVS and 122 using UVC. Only 66% of all species were detected using both methods. This study showed that the upwelling had an important role in shaping reef fish assemblages in this region, but there was also a significant interaction between upwelling and location. In addition, other drivers such as habitat complexity and habitat composition had an effect on reef fish abundances and species. To authors’ knowledge, this is the first study in the Eastern Tropical Pacific that combines BRUVS and UVC to monitor reef fish assemblages in an upwelling region, which provides more detailed information to assess the state of reef ecosystems in response to multiple threats and changing ocean conditions.