Examining annual catch trends and gear selectivity of bull sharks (Carcharhinus leucas): implications for Queensland’s coastal management
Riley W. Banaghan
A * , Alexis L. Levengood A and Bonnie J. Holmes A
A
Abstract
In coastal waters of eastern Australia, the bull shark (Carcharhinus leucas) is one of the most frequently caught shark species within the Queensland Shark Control Program (QSCP).
An analysis of 27 years of QSCP catch data (1996–2022) involving 2352 bull sharks from nine locations spanning 1700 km captured in gill-nets or on drumlines identified changes in catch composition and gear selectivity.
Linear regression tested trends in length and catchability, and selectivity and bias were evaluated through mean and distribution comparison tests.
Drumlines caught more sharks (80.1%), indicating stronger selectivity for C. leucas of all size classes. A decline in standardised catch per unit effort (CPUE) was noted in tropical regions for each gear type, yet there were increases of bull sharks caught within subtropical gill-nets (P < 0.001). The sex ratio was female biased on drumlines (1.85:1) and in gill-nets (1.53:1).
Our findings corroborated previous research, highlighting the significance of considering regional variations in CPUE, sex ratios and size compositions of sharks caught in shark control programs.
Catch rates provide insights into shark population trends, particularly where gear configurations have remained relatively consistent. Downward trends may indicate unsustainable harvest, or external influences.
Keywords: bather protection, bull shark, Carcharhinus leucas, conservation, drumlines, elasmobranchs, fisheries, gill-net.
Introduction
Elasmobranchs, including sharks, rays and skates, are globally distributed and play multifaceted roles in marine ecosystems (Myers et al. 2007; Pimiento et al. 2023). However, their life-history traits, such as slow growth, delayed reproduction, and low reproductive rates, make many species highly susceptible to anthropogenic exploitation (Hoenig 1990; Dulvy et al. 2017, 2021). Overfishing, habitat degradation and climate change have contributed to widespread declines, with recent global assessments estimating that nearly a third of all elasmobranch species face extinction risk (Dulvy et al. 2008, 2021; Pacoureau et al. 2021), making them one of the most vulnerable vertebrate classes (Ward-Paige et al. 2012). These declines have triggered significant ecological consequences, including trophic cascades and shifts in prey populations (Ferretti et al. 2010; Estes et al. 2011; Ripple et al. 2014; Hammerschlag et al. 2025). As apex predators, sharks exert a top-down influence on food webs, ecosystems and habitats (Heupel et al. 2014; Dulvy et al. 2017), underscoring the urgency of addressing ongoing population declines and the lack of recovery (Ferretti et al. 2010; Roff et al. 2018). Monitoring shark populations is often hindered by sharks’ elusive and wide-ranging nature (Heithaus et al. 2002; Werry et al. 2014) and the expansive geographic scale of marine ecosystems (Baum et al. 2003).
The bull shark (Carcharhinus leucas) belongs to the family Carcharhinidae, commonly known as whaler sharks in Australia (Last 2009), and requiem sharks globally. Among the species within this family, the genus Carcharhinus is the most diverse, comprising 35 of 57 species (Collareta et al. 2022). Being one of the most adaptable large-bodied shark species, bull sharks occupy marine, estuarine and freshwater environments across warm temperate, subtropical and tropical regions (Thorson 1972; Thomerson et al. 1977; Pillans et al. 2005; Smoothey et al. 2023). Currently listed as ‘vulnerable’ by the International Union for Conservation of Nature (IUCN) (Rigby et al. 2021), bull sharks exhibit natal philopatry, with females returning to specific estuarine and freshwater nurseries to give birth (Tillett et al. 2012; Heupel et al. 2015; Smoothey et al. 2019). In Australia, bull sharks are widely distributed along the northern, eastern and western coastlines, with genetic studies indicating high connectivity driven by large-scale male movements and female site fidelity (Devloo-Delva et al. 2023). Ontogenetic shifts in habitat use are well-documented, with juveniles primarily occupying low-salinity nursery areas, whereas subadults and adults undertake extensive seasonal migrations along the coast (Simpfendorfer et al. 2005; Heupel et al. 2010; Heupel and Simpfendorfer 2011).
Bull sharks frequently overlap with human activities, particularly in estuarine and nearshore environments. This overlap has led to increasing human–wildlife conflict, particularly in areas with high recreational and commercial fishing activity (Werry et al. 2012; Smoothey et al. 2023). For example, recent studies on shark depredation in Australia have indicated that bull sharks are contributing to the increasing depredation of fish catches (Mitchell et al. 2018; Vardon et al. 2021; Mitchell et al. 2023). Additionally, their presence in urban waterways, such as the Sydney Harbour and the Brisbane River, has led to concerns over public safety, with a small number of recorded shark attacks in these areas (Smoothey et al. 2019).
Across their range, bull sharks are susceptible to anthropogenic pressures relating to subsistence, recreational and commercial fishing activities, including those related to bather protection programs (Harry et al. 2011; Werry et al. 2012; Dulvy et al. 2021). Illegal targeting of large sharks by foreign fisheries throughout Australian waters (Field et al. 2009; Marinac 2022), along with unregulated, unmonitored exploitation, has played major roles in the over-exploitation of shark fisheries resources (Sumaila et al. 2006). Globally, it is estimated that ~1 million bull sharks are caught annually, equating to ~30,000 tonnes (Mg) (Clarke et al. 2006), and declines in bull shark lengths and population sizes have been reported in many locations (Cliff and Dudley 1991; O’Connell et al. 2007; Haig et al. 2018). Concomitantly, declines in several ‘whaler’ shark populations, which included bull sharks, have been reported in Queensland (Qld) (Dudley 1997; Roff et al. 2018; Henderson et al. 2024). For example, Haig et al. (2018) conducted a species-specific analysis on bull shark populations by using shark control data in Qld (1996–2012), reporting widespread and localised declines in size and abundance. However, despite these historical declines, recent reports suggest a potential increase in juvenile bull shark populations in some regions, possibly linked to warming temperatures and urbanisation (Mullins et al. 2024). Similar trends have been noted in Qld, where both recreational and commercial fishers report an increase in bull shark encounters in local waterways (Vardon et al. 2021). The recent rise in depredation rates in Australian fisheries further highlights the need to reassess bull shark populations and their interactions with human activities (Mitchell et al. 2018, 2023).
Given the scarcity of fisheries-independent monitoring for large sharks, shark control programs provide a valuable long-term dataset for assessing population trends (Sumpton et al. 2011; Haig et al. 2018; Lee et al. 2018, 2019; Niella et al. 2020, 2021; Lopes et al. 2024). Here, we analysed long-term temporal and spatial trends associated within bull shark catch across nine locations spanning 1700 km within the QSCP over a 27-year period (1996–2022). The objectives of this study were to (1) determine whether widespread declines in catch per unit effort (CPUE) within the QSCP are currently evident for bull sharks, (2) identify whether shifts in the average length of bull sharks are occurring within Qld waters, and (3) assess changes in relative catchability and relationships among gear type, location, sex and size of bull sharks. The outcomes of this study will enable a greater understanding of the stock status of bull sharks within Qld, and identify if shark numbers and size-shifts are occurring in coastal waters of Qld.
Methods
Queensland Shark Control Program (QSCP)
The QSCP extends along the eastern seaboard of Qld, from the Gold Coast to Cairns, using year-round deployments of a combination of gill-nets and drumlines along 85 beaches (Fig. 1). The drumlines are positioned ~500–1000 m from the shoreline and equipped with a single 14/0 J stainless steel hook (Sumpton et al. 2011), baited with ~2 kg of sea mullet (Mugil cephalus) or shark flesh. Gill-nets are deployed at the same distances from shore, running 186 m parallel to the shoreline, with a gauge of 50 cm and a 6-m drop (Holmes et al. 2012; Haig et al. 2018; Werry et al. 2018). All gill-nets used within this program are surface set to minimise entanglements with non-target benthic species (e.g. stingrays) (Sumpton et al. 2011).
The program comprises 10 contract areas, with Bribie Island being placed under the Sunshine Coast contract, with each region consisting of several beaches (Holmes et al. 2012). Independent shark contractors must service the gear every 1–3 days, with all gear being checked 15–20 times monthly (Holmes et al. 2012; Haig et al. 2018; Werry et al. 2018). Each time the gear is serviced, the contractor completes an activity report (Department of Agriculture and Fisheries 2024). When a shark is caught, contractors record key information, including species, sex (presence of claspers in males), total length (m TL), fate, location and gear type (Department of Agriculture and Fisheries 2024). The total length is measured from the tip of the nose to the end of the tail’s upper lobe in a natural position, measured on the dorsal side of the shark. During routine checks, any damaged gear and missing bait are replaced (Haig et al. 2018).
Size classes were identified and split into three life-stage categories. Adults were identified as larger than or equal to 2.2 m TL; this was chosen because the size at maturity for bull sharks was estimated between 2.04 and 2.25 m TL for females and between 1.90 and 2.20 m TL for males (Branstetter and Stiles 1987; Cruz-Martínez et al. 2005). Subadults were classified as those ≥1.75 m TL (but <2.2 m), because this is when bull sharks on the eastern coast of Australia are shown to make greater use of coastal habitats (Niella et al. 2022; Smoothey et al. 2023); juveniles subsequently were categorised as sharks <1.75 m TL, because they are often still site attached to riverine ecosystems (Branstetter and Stiles 1987; Werry et al. 2011).
Throughout a significant portion of its operating history, the species identification process within the QSCP has been considered unreliable, primarily owing to confusion among morphologically similar species commonly caught (Macbeth et al. 2018). For instance, the pig-eye shark (Carcharhinus amboinensis), which is also present in Queensland waters, closely resembles the bull shark and shares extensive habitat overlaps (Last 2009). Consequently, after a systematic review, the QSCP underwent significant changes in species identification protocols in 1996 (34 years after its establishment). Evaluations conducted in 1992 and 1996 by two Ministerial Committees highlighted substantial misidentifications, leading to recommendations for improved training in species-level identification (Gribble et al. 1998). Following these reviews, several specialised courses were introduced to enhance the accuracy of species identification within the program, particularly for whaler sharks (Family Carcharhinidae), to address long-standing classification challenges (Werry et al. 2018).
Given these issues, the data analysed herein were collected post-1995, when identification protocols were improved. However, this presents a significant limitation in reconstructing historical baselines, as the initial population structure and catch composition before these improvements remain uncertain. The lack of reliable species-level data from the program’s first three decades complicates efforts to assess long-term population trends and the full impact of the QSCP. If bull sharks were misidentified at a substantial rate during the early years of the program, the historical catch records may not accurately represent the species’ true abundance and size structure at the time. This uncertainty underscores the need for caution when interpreting long-term declines and highlights the importance of considering potential biases in historical data when assessing population trends.
Since its establishment, the QSCP has undergone significant variations in fishing configurations and effort. Initially, the program employed a large proportion of gill-nets throughout its operating beaches. However, because of unacceptable bycatch rates of non-target species, primarily turtles and dugongs (Sumpton et al. 2011), and a decrease in shark catch (Paterson 1990), they were changed to single-hook drumlines in many regions. To appropriately model CPUE metrics for each gear type, it was essential to adjust for these changes in effort (Fig. S1 of the Supplementary material). Effort data were acquired from the Queensland Department of Agriculture and Fisheries (QDAF), taking into consideration seasonal gear lifting and reconfigurations. To improve the accuracy of the analysis, data for several regions were also filtered and omitted. Specifically, data from the Capricorn Coast region were excluded post-2021 owing to the initiation of the Catch Alert Drumline (CAD) trial, which uses gear configurations different from those of the historical setups (Campbell and Scott-Holland 2023). Additionally, the period immediately preceding this trial (2019–2021) was omitted because of inconsistencies in gear configuration, deployment and servicing, as documented in the CAD trial report (Campbell and Scott-Holland 2023). Rainbow Beach was also excluded from the catch analyses owing to data quality concerns (Holmes et al. 2012), and therefore, the catch rate results here were not considered further. The gear type was recorded into the following three categories: ‘drumline’, ‘gill-net’ and ‘other’. However, the ‘other’ category was omitted because of uncertainty regarding the specific gear used when a shark catch was not properly assigned a gear type. Additionally, effort was not standardised across gear types, meaning that differences in the number of deployed drumlines and gill-nets (Fig. 1) were not accounted for in the analysis. This variation in effort should be considered when interpreting differences in catch rates among gear types.
Statistical analyses
To identify trends within the annual CPUE of bull sharks within the QSCP, the catch information (total sharks caught per year for each region) was fit into generalised linear models (GLMs) within R (ver. 4.3.2, R Foundation for Statistical Computing, Vienna, Austria, see https://www.r-project.org/). The significance level of all statistical tests was set at alpha = 0.05. Individual beaches were aggregated by region, allowing for the following nine study regions: Cairns, Townsville, Mackay, Capricorn Coast, Gladstone, Bundaberg, Sunshine Coast, North Stradbroke Island and Gold Coast. The QSCP regions of Bribie Island, Sunshine Coast North and Sunshine Coast South were aggregated to form the Sunshine Coast region. Each gear type (gill-net or drumline) and location were modelled independently. The dependent variable, the number of sharks caught per year, was assumed to follow a negative binomial distribution (Yates et al. 2015). This dependent variable was modelled against year for each location and gear type by using additive models to identify inter-annual trends and patterns within the catch, resulting in nine models assessed. Models included an offset term employed to represent the logarithm effort, quantified as sharks per gill-net per day or sharks per drumline per day. This offset enabled the response variable to be transformed into an adjusted CPUE pooled by year level. The annual adjusted CPUE for tropical and subtropical regions was investigated utilising the same methodology, although beaches were aggregated by latitude. The tropics were considered any location from Cairns to Bundaberg, whereas subtropics were classified as any area from Sunshine Coast to Gold Coast. Bundaberg was included within the tropical region because of the land bridge to the continental shelf, separating it from other subtropical regions; this is also consistent with Werry (2010) to enable direct comparisons.
Linear regression models were also utilised to analyse average length shifts for bull sharks within the QSCP. Sharks were pooled by region, with each gear type being modelled separately. Time-series means were employed to investigate shifts in the annual average length of sharks caught for each gear type for each location. To analyse relationships among gear type, sex, region and size, average lengths were compared across gear, sex and region by using independent-sample Student’s t-tests. Size distributions were grouped by gear type and were compared using a non-parametric Kolmogorov–Smirnov test (Chakravarti et al. 1967). The sex ratios of state-wide aggregated shark catches were tested using a Chi-Square test (χ2).
Results
Over the 27-year (1996–2022) study period, 2352 Carcharhinus leucas individuals were captured within the QSCP. Of these, 55.5% were female and 44.5% were male, with sex ratios significantly deviating from unity for both drumline (χ2 = 13.7, d.f. = 1, P < 0.001) and net catches (χ2 = 20.8, d.f. = 1, P < 0.001), displaying female to male ratios of 1.85:1 and 1.53:1 respectively (Table 1). The size of captured sharks across all gear types and locations ranged from 0.6 to 4.0 m TL, with a mean total length (TL) of 1.8 ± 0.5 m. The largest individual recorded was a 4.0 m TL female, caught on a drumline at the Sunshine Coast. Across age classes, the sex ratio varied, with 0.94:1 for juveniles (χ2 = 0.99, d.f. = 1, P = 0.32), 1.43:1 for subadults (χ2 = 21.37, d.f. = 1, P < 0.001) and 1.87:1 for adults (χ2 = 51.75, d.f. = 1, P < 0.001). Most drumline regions, except for the Capricorn Coast, Sunshine Coast and Townsville, captured significantly more females (Table 1). The capture of sharks on drumlines at North Stradbroke Island was predominantly female (85.1%), and similar trends were observed in the Gold Coast and Bundaberg regions. QSCP nets also showed a higher proportion of females across all regions, although no major skewing towards either sex was observed (i.e. <70% skew) (Table 1). In Townsville, where nets were removed in 2005, both sharks caught were female.
| Location | Drumline catch composition | Gill-net catch composition | ||||||
|---|---|---|---|---|---|---|---|---|
| ♂ (%) | ♀ (%) | Total (n) | ♂ (%) | ♀ (%) | Total (n) | |||
| Tropics | Cairns | 36.9 | 63.1 | 198 | 38.9 | 61.1 | 18 | |
| Townsville | 44.2 | 55.8 | 294 | 0 | 100 | 2 | ||
| Mackay | 43.3 | 56.7 | 261 | 42.2 | 57.8 | 204 | ||
| Capricorn Coast | 55.4 | 44.6 | 697 | |||||
| Gladstone | 44.4 | 55.6 | 196 | |||||
| Bundaberg | 27.1 | 72.9 | 85 | |||||
| Subtropical | Sunshine Coast | 47.4 | 52.6 | 79 | 40.9 | 59.1 | 149 | |
| Point Lookout | 14.9 | 85.1 | 47 | |||||
| Gold Coast | 24.2 | 75.8 | 33 | 32.3 | 67.7 | 96 | ||
| Queensland | 45.8 | 54.2 | 1883 | 39.45 | 60.55 | 469 | ||
Empty rows indicate regions with no gear deployed.
Gill-nets and drumlines exhibited significant differences in their catch composition. Approximately 19.9% (n = 469) of all sharks were caught within gill-nets, and 80.1% (n = 1883) were caught on drumlines (not correcting for effort among gear types). The total length distributions between the two gear types differed significantly (Z, P < 0.001). Mean TL was significantly larger for gill-net-captured sharks (mean TL = 2.04 ± 0.02 m) than for drumline-captured sharks (mean TL = 1.68 ± 0.01 m) (Fig. 2). Gill-nets primarily selected for adults (40.3%) and subadults (38%), whereas drumlines predominantly captured juveniles (53.5%) (Fig. 3). Regionally, the Capricorn Coast accounted for 49.28% (n = 515) of all juveniles caught during the study period (Fig. 3).
Size frequency distribution of C. leucas catch within drumline and gill-nets within the Queensland Sharks Control Program (QSCP), between 1996 and 2022. Mean length for each gear type is denoted by the dashed vertical line.
Catch per unit effort of C. leucas (CPUE; number of sharks caught per gear per day × 100) for each deployment area within the Queensland Shark Control Program (QSCP) and total number of sharks caught per each region between 1996 and 2022.
There was a significant interaction between total length and year for drumline and net catches, with notable differences among locations. Declines in annual mean length were observed at Capricorn Coast, Gladstone and Sunshine Coast (Table 2). Further investigation of gill-net catches at specific locations showed significant declines at Cairns, Sunshine Coast and Gold Coast, and a small but significant increase in mean length was detected in the gill-net catch from Mackay (Table 2).
| Area | Location | Gill-net | Drumline | Mean annual length | Mean annual length | Mean annual catch | Gear selectivity | |||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| r CPUE | r CPUE | r (net) | r (drumline) | ♂ n | ♀ n | Gill-net | Drumline | |||||
| Min. size (m TL) | Max. size (m TL) | Min. size (m TL) | Max. size (m TL) | |||||||||
| Tropics | Cairns | −0.089 | −0.065 | −0.049 | −0.003 | 3.14 | 5.59 | 0.90 | 2.90 | 0.70 | 3.20 | |
| Townsville | −0.270 | +0.024 | −0.050 | −0.002 | 4.58 | 5.72 | 2.15 | 2.20 | 0.70 | 3.15 | ||
| Mackay | −0.066 | −0.051 | +0.013 | +0.004 | 7.00 | 8.81 | 1.00 | 3.00 | 0.80 | 3.10 | ||
| Capricorn Coast | −0.001 | −0.027 | 14.96 | 12.52 | 0.50 | 3.20 | ||||||
| Gladstone | +0.033 | −0.021 | 3.50 | 3.72 | 0.60 | 2.70 | ||||||
| Bundaberg | +0.019 | −0.007 | 1.40 | 2.50 | 0.96 | 3.00 | ||||||
| Subtropics | Sunshine Coast | +0.032 | −0.001 | −0.026 | −0.047 | 3.52 | 4.00 | 1.00 | 3.10 | 1.00 | 4.00 | |
| Point Lookout | −0.062 | −0.002 | 1.17 | 1.85 | 1.30 | 3.30 | ||||||
| Gold Coast | −0.006 | −0.009 | −0.005 | −0.003 | 1.67 | 3.43 | 1.20 | 2.80 | 1.00 | 2.74 | ||
| Tropics | −0.029 | −0.013 | +0.010 | −0.015 | 27.24 | 29.15 | 0.90 | 3.00 | 0.50 | 3.20 | ||
| Subtropics | +0.017 | −0.019 | −0.015 | −0.021 | 14.96 | 8.84 | 1.00 | 2.80 | 1.00 | 4.00 | ||
| Queensland | −0.009 | −0.016 | −0.008 | −0.015 | 34.69 | 43.54 | 0.90 | 3.10 | 0.50 | 4.00 | ||
Min., mimimum; Max., maximum; TL, total length measured in metres; r, value of regression coefficient. Bold indicates significance (P < 0.05), + or − highlighting the direction of slope.
Analysis of annual trends in drumline CPUE indicated significant differences among areas (P < 0.001). When aggregated into subtropical and tropical zones, CPUE differed significantly between zones (P < 0.001) and years (P < 0.05). Significant declines in drumline CPUE were detected in Cairns, Mackay, North Stradbroke Island and the aggregated tropical areas, whereas increases were observed in Townsville and Gladstone (Fig. 4).
Trends in annual drumline CPUE for C. leucas (as the number of sharks caught per gear per day, with shaded area showing ±s.e.) within the Queensland Shark Control Program (QSCP), displaying regions that showed significant declines or increases in annual CPUE between 1996 and 2022.
Gill-net CPUE varied significantly among areas (P < 0.001). When aggregated into subtropical and tropical zones, there were significant regional differences (P < 0.001), but no significant trend was detected in state-wide gill-net catches (P = 0.307). However, Mackay experienced significant declines in CPUE over the study period (P < 0.001), whereas the Sunshine Coast showed a significant increase (P < 0.05) (Fig. 5).
Discussion
Shark control programs aim to reduce the risk of shark encounters in nearshore areas by culling or capturing potentially dangerous sharks by using methods such as drumlines and gill-nets. By lowering shark numbers near popular beaches, these programs seek to minimise the chance of negative interactions with humans. However, removing large apex predators can significantly affect ecosystem functioning and ecological balance (Estes et al. 2011; Ripple et al. 2014; Henderson et al. 2024; Hammerschlag et al. 2025). Despite the widespread distribution of sharks globally, comprehensive data on their long-term population trends and the extent of their declines remain limited (Clarke et al. 2006; Ferretti et al. 2010; Roff et al. 2016; Dulvy et al. 2024). This lack of knowledge makes it challenging to effectively monitor and understand the broader ecological consequences of their removal.
Analysing catch size and composition changes over time can provide insights into the exploitation or recovery of fish stocks (Jennings and Kaiser 1998; Guillemin et al. 2025). Additionally, assessing catch rates alongside biological characteristics offers a deeper understanding of a population trends (Maunder et al. 2006; Pacoureau et al. 2021), whereas variations in catch composition can indicate ecosystem status and community change (Greenstreet and Rogers 2006; Hammerschlag et al. 2019). Understanding the stock composition for important coastal species is critical given the burgeoning global shift in climate baselines (McHenry et al. 2019; Niella et al. 2022). By analysing 27 years of catch data from a state-wide shark control program, significant long-term trends in catch rates of bull sharks have been elucidated herein.
Gear-specific trends and catchability of bull shark catch
Gear selectivity significantly influences shark catch composition in fisheries and shark control programs (Dudley and Simpfendorfer 2006; Sumpton et al. 2011). A critical finding of this study is that the QSCP, despite targeting large, potentially dangerous sharks, predominantly captures juveniles and subadult bull sharks. This discrepancy between target (large adult sharks) and actual catch (primarily juveniles) raises important questions about the program’s effectiveness and ecological impact. In the QSCP, drumlines captured a higher proportion of juveniles (53.7%) and accounted for 80.1% of bull shark catches; however, effort was not standardised across gear types. Similar patterns were observed by Haig et al. (2018), where drumlines caught more bull sharks but selected for smaller individuals. By contrast, KwaZulu–Natal (KZN) South African’s shark control program found that gill-nets were far more effective for bull sharks than drumlines (Cliff and Dudley 2011), aligning with Sumpton et al. (2011), who identified bull sharks as the most susceptible target species in Queensland gill-nets. Standardised catch rates in south-eastern Queensland also support this trend (Lopes et al. 2024). Additionally, in the KZN shark control program, drumlines tended to catch smaller bull sharks than did gill-nets (Cliff and Dudley 2011), reinforcing global trends in gear efficiency and selectivity.
The higher number of smaller bull sharks caught on drumlines is likely to reflect a combination of species-specific behavioural traits, environmental factors and gear configurations. Bull sharks are highly opportunistic predators with strong olfactory sensitivity (Tricas and Sisneros 2004; Heupel and Simpfendorfer 2008), potentially making them susceptible to baited gear. Additionally, drumlines are often positioned near estuarine-influenced areas where juveniles are more common (Heupel and Simpfendorfer 2011). Seasonal movements, competition and environmental factors such as turbidity and water temperature may also drive catch selectivity (Lopes et al. 2024). Moreover, the larger number of beaches with drumlines than those with gill-nets may also help explain the higher catch rates.
This juvenile-biased selectivity has ecological implications, as sustained removals could contribute to recruitment overfishing (Diekert 2012) and alter population structures over time (Shin et al. 2005; Greenstreet and Rogers 2006; Svedäng and Hornborg 2017). The gear disparity in catch composition has key management implications. Drumlines appear more effective at targeting bull sharks, but if the goal is to remove larger, potentially more dangerous individuals, modifications such as adjusting hook size, bait type, or deployment depth may be necessary. Similar adjustments have been made in South Africa to gear deployments to help further refine selectivity (Cliff and Dudley 2011). Recent work by Lopes et al. (2024) on bull shark catch rates along eastern Australia further supports the need for strategic gear deployment based on seasonal habitat use patterns.
Length trends and composition across sites
Size trends in shark populations provide key insights into population dynamics and potential shifts in species composition over time (Field et al. 2012). In the QSCP, the predominance of juvenile bull sharks suggests that (1) gear type influences selectivity (e.g. hook size on drumlines), (2) estuarine habitats used by juveniles are near QSCP deployment sites, or (3) the recruiting stock is dominated by younger individuals. Similar declines in the mean total length (TL) of sharks have been documented in multiple regions, including in KZN program (Cliff and Dudley 1991), the QSCP up to 2012 (Haig et al. 2018), and the Gulf of Mexico, where bull shark length decreased in fishing tournaments over an 80-year period (Powers et al. 2013). These findings suggest a widespread trend of a declining shark size in response to fishing pressure and habitat changes.
Our study observed significant declines in the mean total length (TL) of bull sharks in Cairns, Sunshine Coast, Gladstone and Capricorn Coast, suggesting shifts in community composition towards smaller individuals and providing insights into the contemporary dynamics of these coastal shark populations. Annual length declines of 1.5 cm (cm) per year have been identified from statewide QSCP catch, with consistent patterns of decline shown throughout subtropical and tropical regions, as identified herein. This declining average TL aligns with global trends across several shark species in both commercial fisheries and shark control programs (blacknose shark, Carcharhinus acronotus; dusky shark, Carcharhinus obscurus; silky shark, Carcharhinus falciformis; sandbar shark, Carcharhinus plumbeus, Benavides et al. 2021; tiger sharks, Galeocerdo cuvier; whaler sharks, Roff et al. 2018). Similar reductions in the annual average length of bull sharks have been reported by Cliff and Dudley (1991) in the KZN program and by Haig et al. (2018) in the QSCP (up to 2012). Additionally, research in the Gulf of Mexico has indicated that bull shark length decreased during fishing tournaments over an 80-year period (Powers et al. 2013). Our findings indicated that the length trends Haig et al. (2018) observed have continued to decline over the past 10 years, indicating that there are ongoing shifts in the eastern Australian bull shark population that interact with the QSCP. Although the QSCP has been operational since 1962, these continued trends may reflect delayed or compounding effects of gear interaction on population structure. The consistent removal of larger and older individuals can truncate a stock’s structure, leading to fluctuations in abundances and making populations more volatile and less resilient (Anderson et al. 2008; Botsford et al. 2014; Secor et al. 2015). Interestingly, we identified significant increases in the average length of bull sharks in Mackay, a central Qld location known for high prey productivity and large number of estuaries (Haig et al. 2018). It may be that habitats such as these have been integral to maintaining the viability of the broader population, facilitating recovery of the stock from the individuals removed.
The observed declines in average length across the QSCP have significant ecological implications. Smaller average lengths indicate younger sharks, reducing the population’s overall reproductive output and resilience. This trend can lead to trophic downgrading, where the loss of large apex predators disrupts the balance of marine ecosystems, potentially causing an increase in the mesopredator population and subsequent declines in biodiversity (Hammerschlag et al. 2025). Shifts towards smaller sizes in shark populations have been linked to anthropogenic exploitation (Rago et al. 1998; Bradshaw et al. 2008; Roff et al. 2018), possibly influenced by the size-selectivity of fishing gear and improved technologies (e.g. side sonar, electric reels) (McLoughlin and Stevens 1994; Stevens et al. 2000). Our results are consistent with long-term declines in abundance and average length of coastal shark species in QSCP (Roff et al. 2018; Henderson et al. 2024) and highlight the need for adaptive management strategies. The observed changes within nearshore communities are likely to indicate anthropogenically induced trophic cascades (Henderson et al. 2024). Specifically, the shifts composition of juveniles may signal changes in population structure or habitat use, suggesting that these habitats serve as critical nursery regions. Localised shifts in catch composition necessitate targeted coastal management actions, such as refining spatial management zones, adjusting gear types or implementing additional protections in these important shark habitats.
Previous studies have established linkages between nearshore fisheries production and estuarine habitat structures (Lee 2004; Manson et al. 2005; Lefcheck et al. 2019), including relationships with the extent of intertidal habitat and mangrove cover (Lee 2004; Heithaus et al. 2009). Juvenile bull sharks rely on nearshore coastal environments as nursery habitats, providing protection and foraging opportunities (Simpfendorfer et al. 2005; Heupel and Simpfendorfer 2008; Heupel et al. 2010; Curtis et al. 2011; Bangley et al. 2018; Niella et al. 2022). Here, we found that catch of juvenile bull sharks on drumlines in waters off the Capricorn Coast waters was significantly higher than at all other QSCP locations. The Capricorn Coast region supports the greatest wetland area of all tropical sites and the second-highest area within all QSCP sites (Haig et al. 2018). The positive influence of wetlands on bull shark CPUE, as documented by Haig et al. (2018), indicates that the proximity of the QSCP gear to this area is likely to be driving the high juvenile catch rates.
Trends in CPUE
Bull sharks were caught in all QSCP locations, indicating a widespread distribution along the coastline. Regions within the tropics (north of Bundaberg <24.8°S) exhibited consistently greater CPUE of bull sharks than regions south of this point. A similar relationship can be seen in the northern hemisphere, where higher abundances were observed within the lower latitudinal waters off the Gulf of Mexico (~25°N) than in the higher latitudinal regions off Florida (~28°N) (Curtis et al. 2011). The nearshore tropical regions of eastern Australia are characterised by consistently warm temperatures, shallow waters (<50 m deep), and a distinct monsoon period. Various factors such as prey availability (Hammerschlag et al. 2012; Lubitz et al. 2023), water temperature (Smoothey et al. 2016; Niella et al. 2020; Lopes et al. 2024), rainfall (Werry et al. 2018) and habitat features (Haig et al. 2018) have been shown to influence the distribution of bull sharks. These sharks prefer warm waters at all life stages, with juveniles being observed to shift nursery areas in response to global climate change (Bangley et al. 2018; Mullins et al. 2024). Water temperature influences both the fine-scale habitat selections and large-scale migrations of bull sharks (Werry et al. 2018; Espinoza et al. 2021; Smoothey et al. 2023). Notwithstanding, changes in ocean conditions owing to climate-induced warming waters may affect these patterns. A study by Niella et al. (2020) off eastern Australia indicated a 3-month increase in the availability of favourable water temperatures for bull sharks at higher latitudes by 2030 under modelled future climate scenarios. These shifts in habitat use may lead to an increase within CPUE for higher latitudes within the QSCP and, consequently, possible increases in human–shark interactions within south-eastern Qld.
Local population fluctuations with significant inter-annual CPUE variations were observed across regions and gear types. Notably, over the study period, CPUE declines were seen in Cairns, Mackay and North Stradbroke Island, whereas Gladstone, Sunshine Coast and Townsville displayed significant CPUE increases. As noted in other studies, inter-annual bull shark catch rates are highly variable, influenced by environmental changes, prey availability and female philopatry (Dudley and Simpfendorfer 2006; Hammerschlag et al. 2012; Haig et al. 2018). In an unpublished study by Werry (2010), the reported QSCP bull shark catch rates from 1996 to 2006 had several key similarities to our findings herein. First, we identified CPUE increases within Townsville, as well as increases in catch at several tropical locations, which were attributed to several key environmental factors (Werry 2010). Werry (2010) suggested that unusually low rainfall between 1991 and 1995 (annual average 368 mm for the Ross River, Townsville) followed by substantial rainfall from 1996 to 2006 (annual average 995.4 mm for the Ross River) could account for the significant increases in juvenile catch. Werry (2010) also showed that peak catches for Cairns, Townsville, Mackay and Gladstone were attributed to peak rainfall years. Environmental drivers, such as rainfall, significantly influence bull shark occurrences, with salinity a known driver of fine-scale habitat use within juveniles (Schlaff et al. 2014). Both adult and juvenile bull sharks may utilise these lower salinity habitats to a higher extent after significant rain events to reduce inter-species competition, because they can tolerate low salinity for prolonged periods (Pillans et al. 2008). These relationships may contribute to our observed widespread inter-annual variations in catch rates, but our understanding of these drivers remains limited to very few empirical studies.
The observed fluctuations in localised CPUE are likely to be influenced, at least partly, by the highly complex and variable occurrence patterns of bull sharks, driven by several intrinsic and extrinsic factors (Werry et al. 2011; Heupel et al. 2015; Espinoza et al. 2016). These occurrence patterns involve large-scale migration (Heupel et al. 2015), residency (Heupel and Simpfendorfer 2008; Brunnschweiler and Barnett 2013), site fidelity (Brunnschweiler and Baensch 2011), regional philopatry (Tillett et al. 2012; Sandoval Laurrabaquio-A et al. 2019) and partial migrations (Espinoza et al. 2016). Intra-specific variations and the complex behaviours of bull sharks are likely to influence trends in CPUE within a fixed-effort fishery. Acoustic tracking studies have shown that bull sharks undergo partial migrations, meaning that some individuals will display residency whereas others undertake large migrations (Espinoza et al. 2016, 2021). This partial migration phenomenon is also seen in other large shark species along the eastern coast of Australia (e.g. tiger sharks, Galeocerdo cuvier; Holmes et al. 2014; Werry et al. 2014) and could explain some of the inter-annual variations displayed within nearshore catch rates of bull sharks. Additional data are needed to accurately elucidate the drivers behind these significant inter-annual trends within CPUE. Correlating tracking data with environmental information (e.g. rainfall, temperature) and fishing mortality is key to identifying potential drivers causing these trends. Although observed reductions in catch rates are consistent with localised stock depletions, cautious interpretations are needed, given the significant inter-annual variations in CPUE in the shark control data. Owing to the unknown levels of connectivity between coastal shark populations in Qld and accurate descriptions of the key drivers, interpretation of local catch data in isolation could result in very different conclusions than those made from data across wider regions.
Sex ratios
Sexual and size segregation have been identified in numerous oceanic and coastal shark species, such as bull, tigers, scalloped hammerheads (Sphyrna lewini), shortfin mako (Isurus oxyrinchus), blue shark (Prionace glauca) and blacktip reef sharks (Carcharhinus melanopterus) (Mucientes et al. 2009; Noriega et al. 2011; Werry et al. 2011; Werry and Clua 2013). Such intra-specific variability in behaviour and movement occurs for various reasons, including predator avoidance, reduced competition, differing nutritional requirements, or reproduction; however, these are highly context dependent (Lubitz et al. 2022). This study observed a significant sex bias in the state-wide program, with the catch of bull sharks predominately consisting of females across all life stages except juvenile. Female bias has been observed within whaler shark populations throughout the NSW Shark Meshing Program (Reid et al. 2011) and the KZN program (Cliff and Dudley 1991). One possible explanation for the observed sex bias is the unique reproductive strategy of bull sharks. Previous studies have shown that females return to rivers to pup bi-annually (Werry et al. 2011, 2012; Tillett et al. 2012), with gravid adult females using shallow waters (<5 m) for parturition (Lea et al. 2015). Espinoza et al. (2016) highlighted intra-specific differences, reporting that mature female bull sharks in the central Great Barrier Reef undertake wider migratory movements than do males. Our data indicated that sexual segregation occurs at the point nearing reproductive maturity, suggesting reproduction could be driving intra-specific variation in habitat use on the Australian eastern coast. Mating and courtship in sharks are often aggressive, with males inflicting significant bite wounds on females (Klimley 1987). This aggression may lead to sexual segregation in some species, because it can increase female fitness by reducing mating frequency (Mucientes et al. 2009). It is possible that bull shark harassment by males influences female sharks to display avoidance behaviours, which Mucientes et al. (2009) proposed to cause geographic-scale segregation within shortfin makos. Significant sex-based spatial segregation has also been observed in other shark species in eastern Australia (e.g. scalloped hammerheads, Noriega et al. 2011; tiger sharks, Holmes et al. 2012), and understanding the fine-scale dynamics of this phenomenon highlights the need for further research. Migrations and habitat use are complex and context-dependent, influenced by environmental variability and prey dynamics (Lubitz et al. 2023). Long-term tracking can help elucidate how factors such as size, sex and environmental context drive habitat choice and spatial segregation in sharks.
Management implications and future research
The observed declines in CPUE, average length and the significant sex bias in catch, highlight the need for further research to assess the complexities of assessing stock status of bull shark populations off the Australian eastern coast. Understanding gear selectivity and CPUE trends is essential for effective fisheries management (McClanahan and Mangi 2004; Sampson 2014; Maunder et al. 2020). Despite shark control programs being designed to catch and remove large, potentially dangerous, sharks from the population, the majority of bull sharks caught and killed in the QSCP during the study period were juveniles or immature subadults. This discrepency between target (large adults) and actual (primarily juveniles) catch suggests that current methods are not optimally addressing the program’s primary objective of reducing human–shark conflict.
On the basis of our findings, we recommend several management actions to improve the effectiveness of the program and reduce ecological impacts, including the following:
Modify gear specifications. Increasing hook sizes on drumlines could reduce juvenile captures while maintaining effectiveness for larger target sharks.
Alternative technologies. Integrate non-lethal alternatives such as expanding catch alert drumline (CAD) trial (which allow live release), electromagnetic deterrents and drone surveillance in high shark occurrence areas identified in this study.
Targeted research program. Implement a comprehensive tagging and tracking program for released individuals to better understand movement patterns and habitat use, which could inform more precise gear placement.
Regular review of effectiveness. Establish ongoing assessment of whether catch composition aligns with program objectives, with metrics to evaluate both safety outcomes and ecological impacts.
Local eastern coast commercial shark fishers also predominantly target juvenile bull sharks for flake and are fished for and landed within estuaries (Niella et al. 2020). Queensland’s shark fisheries operate under a gauntlet-style approach, in which only a subset of the population (small sharks) are subject to fisheries harvest (Kinney and Simpfendorfer 2009). This mangement style is strategised to conserve populations of long-lived, slow-to-reproduce, shark species (Simpfendorfer 1999). However, regional depletion in catch and long-term shifts in shark length indicate that the extraction of mature bull sharks is still frequent enough that these declines are being observed.
In the Gulf of Mexico, juvenile bull sharks have shown significant population increases in recent years, thriving in response to the region’s warming climate (Bangley et al. 2018; Mullins et al. 2024). Given that both QSCP and shark fisheries in the region are reporting large proportions of juveniles within catches, we must consider that these shifts may be driven by several successful pupping and recruitment pulses to the fishery over the past few years. Additionally, with south-eastern Queensland having been identified as a global climate change hotspot (Hughes and Steffen 2017), population shifts similar to those observed in the USA may also be occurring here. Anecdotally, local fishers are also reporting increases in shark abundance and depredation events in both recreational and commercial fishing pursuits. Future research should focus on identifying the species responsible for depredating catches, along with improved local population estimates and relatedness to gain insights into how bull sharks use nearshore regions of Qld. This information will be critical in determining whether the observed fluctuations within regional CPUE indicate population-level changes in abundance.
It is important to acknowledge that the reliance on catch data from control programs may introduce bias because of non-random sampling methods. Additionally, the analysis did not account for environmental variables, which influence shark distributions (Smoothey et al. 2023). The limitation of data starting from 1996, rather than the program’s inception in 1962, means we may be observing only part of the population response to long-term fishing pressure. Comparisons with the KZN shark control program, which documented significant catch declines in the first 5 years of implementation (Cliff and Dudley 1991), suggest that our analysis may underestimate the true extent of population impacts. Contemporary research efforts should incorporate these external factors to assess the drivers of bull shark CPUE shifts more accurately. Given the projected range shifts driven by water temperature for marine species, investigations into environmental drivers and ecological baselines for the species in Qld waters are advised, particularly considering the displayed variations in CPUE of the species temporally and spatially.
Species identification
A notable limitation of this study is the inherent difficulty in distiniguishing between bull sharks (C. leucas) and pig-eye sharks (C. amboinensis) at sea. Despite training provided to contractors, morphological similarities between these species, particularly in juvenile stages, create potential for misidentification. This challenge has been documented in shark control programs globally (Cliff and Dudley 2011; Reid et al. 2011) and may influence the accuracy of our species-specific catch data and subsequent trends.
To address this limitation in future research and strengthen confidence in species-specific trends, we recommend implementing validation protocols, including the following:
Genetic verification through tissue sampling of a subset of captured individuals, particularly in regions where both species co-occur.
Standardised photographic documentation of key diagnostic features to allow for expert verification.
Regular assessment of identification accuracy through blind tests with contractors to quantify and potentially correct for misidentification rates.
These validation measures would strengthen confidence in species-specific catch data and trends, providing more robust information for management decisions. The implementation of these methods would be particularly valuable given the different ecological roles and conservation statuses of these two species and the need for species-specific management approaches.
Data availability
The data that support this study will be shared upon reasonable request to the corresponding author.
Declaration of funding
This work was funded under the Sunshine Coast Bull Shark Program, led by the University of the Sunshine Coast, with partners the Queensland Department of Agriculture and Fisheries, Sunshine Coast Council, Noosa Biosphere Reserve Foundation and Sea Life Mooloolaba.
Acknowledgements
We acknowledge Elders past and present, who are the traditional custodians of the lands and waters, on which this work was conducted. We thank the QSCP contractors throughout Queensland for their efforts in collecting the data utilised within this study. We also thank Dr Tracey Scott-Holland and Dr Matthew Campbell from the Queensland Government: Department of Agriculture and Fisheries, for access to the historical QSCP data, historical effort information, and insights into the program’s operational history.
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