1. Introduction
Marine fishes host diverse parasite assemblages that can affect host physiology, survival, and ecosystem functioning [
1,
2,
3,
4]. These impacts may be intensified under climate variability as parasite prevalence and diversity often respond to seasonal and interannual changes [
5,
6,
7,
8]. In artisanal fisheries, parasitic infections also compromise product quality and marketability, with implications for food security and human health [
9,
10]. Beyond pathology, parasites are increasingly recognized as integral components of marine food webs and trophic networks [
11,
12], offering insights into host diet, resource use, and ecological interactions. Comparative studies in tropical carangids have demonstrated their utility as biological tags for stock discrimination [
13,
14,
15], while broader ecological syntheses highlight their role as sentinels of ecosystem change [
4,
16,
17,
18]. Together, these findings establish a conceptual framework in which parasites serve as bioindicators of both fisheries stock structure and ecosystem resilience, situating local artisanal fisheries within global debates on sustainability and climate-driven variability.
Current research increasingly examines parasite biology and ecology to illuminate host dynamics [
19,
20,
21,
22], analyze trophic networks [
9,
11,
23,
24,
25], investigate phylogenetic relationships [
26,
27], evaluate environmental quality [
16,
17], predict population trends [
13], and project species distributions under climate change scenarios [
4,
18,
28,
29,
30,
31]. These approaches underscore the value of parasites as bioindicators, not only for local stock discrimination, but also for understanding broader ecosystem responses to global environmental change.
Caranx caballus Günther, 1868 (Carangiformes; Carangidae) is distributed along the northern Colombian Pacific coast and occurs year-round; it is also an important target of artisanal fisheries [
32,
33]. Given its relevance to coastal fisheries in the central Eastern Tropical Pacific (ETP), recent studies have characterized the
Caranx caballus fishery in the Las Palmas Archipelago, Panama [
34], its feeding habits off Manzanillo, Colima, Mexico [
35], and, more recently, its parasite fauna across three Mexican Pacific localities, where select parasite species were proposed as biological markers for stock discrimination [
13]. Comparative studies across the ETP therefore provide an opportunity to situate Colombian findings within a regional framework, contributing to a broader understanding of host–parasite interactions and their implications for fisheries management under climate variability.
In Colombia, studies on fish parasite ecology are sporadic and generally limited to a few commercially important Caribbean species. In contrast, the Pacific region—where the largest national fisheries operate—remains understudied. To address these knowledge gaps, this study provides the first systematic characterization of the metazoan parasite community associated with Caranx caballus (Green Jack) in the northern Colombian Pacific, within the Eastern Tropical Pacific. We hypothesized that parasite prevalence and mean intensity would vary seasonally in relation to environmental conditions, that endoparasites community composition would be linked to host diet—particularly crustacean prey such as brachyuran megalopa—and that host relative condition (Kn) would remain unaffected under low infection intensities. Our objectives were: (1) to identify the parasite species associated with C. caballus captured during artisanal fishing operations; (2) to describe parasite community structure, prevalence, and infection levels across seasons and localities; (3) to evaluate whether parasitism influences host condition under current infection intensities; and (4) to explore the role of parasite assemblages as ecological indicators of trophic interactions, fisheries stock structure, and climate-driven variability. By testing seasonal variation, diet linkages, and host condition alongside the bioindicator role of parasite assemblages, this study presents results from the first systematic investigation of the parasite community associated with an artisanal fish species of importance in the northern Colombian Pacific and contributes to a broader conceptual framework in which parasites inform host ecology, fisheries stock structure, and climate-driven ecosystem variability, thereby linking local artisanal fisheries in the northern Colombian Pacific to global debates on food security and marine resilience.
2. Materials and Methods
The study was conducted in the northern Colombian Pacific, specifically in the Gulf of Cupica, between Bahía Solano (6°13′38.15″ N; 77°24′15.60″ W) and Punta Piñas (6°39′24.50″ N; 77°31′25.90″ W). This area lies within the Eastern Tropical Pacific, is influenced by the Panama and Colombia Currents, and has been designated as an Exclusive Artisanal Fishing Zone (ZEPA), extending 2.5 nautical miles from Punta Solano to Punta Ardita along the border with Panamá. Three sampling sites were selected within artisanal fishing grounds: Bahía Solano (Site 1), La Tebada (Site 2), and Punta Piñas (Site 3) (
Figure 1).
Routine artisanal fishing operations targeting
C. caballus (Green Jack) were conducted during two contrasting seasonal periods: the dry season (January) and the wet season (April), following regional climatic classifications [
36]. Specimens were captured using standardized nocturnal handline fishing (6 h per site). For each fish, standard length (cm) and weight (g) were recorded. External surfaces (skin and fins) were inspected in situ, muscle samples were examined by compression between glass slides, a widely applied technique in fish parasitology [
37,
38,
39,
40,
41] that is comparable in sensitivity to candling under field conditions, and internal organs were fixed in hot 4% formalin. Gills and fins were stored separately for laboratory analysis. No histopathological evaluation was performed as the scope of this study was limited to parasitological identification, quantification, and epidemiological assessment of infection patterns in
C. caballus.
Oceanographic variables (temperature and salinity) were measured prior to fishing effort using a YSI Professional Plus multiparameter probe (Xylem Inc., Yellow Springs, OH, USA) following protocols by Giraldo et al. [
42], and vertical profile from surface to 80 m depth was registered using a CTD Castaway probe Sontek
® (Xylem Inc., Yellow Springs, OH, USA). These parameters were included to explore potential associations between parasite infection metrics and environmental conditions.
Parasite identification was conducted under stereomicroscopy using morphological criteria. Monogeneans and digeneans were stained and dehydrated following the method of Vidal-Martínez et al. [
37], while copepods were cleared in glycerin–ethanol solutions [
38,
43]. Species determination relied on standard taxonomic keys and references [
39,
40,
44,
45], and voucher specimens were deposited in institutional collections. Molecular confirmation was not possible, and this limitation has been acknowledged. Epidemiological indices metrics followed those of Bush et al. [
41], including prevalence (% infected hosts), mean abundance (parasites per host, including uninfected fish), and mean intensity (parasites per infected host). Parasite communities were analyzed at infracommunity (within individual hosts) and component community (across host subsets and environmental contexts) levels. Community structure was characterized by species richness, dominance (Berger–Parker index), and diversity (Brillouin and Shannon–Wiener indices). Qualitative similarity between assemblages was assessed using the Jaccard index (J), calculated as the proportion of shared species across seasons, and its ecological importance was quantified using the Specific Importance Index (IE = Prevalence + [Mean Abundance × 100]; [
46,
47]), with parasites classified as primary (>65% prevalence), secondary (40–65%), or satellite (<40%) species [
48].
Fish condition was assessed using Fulton’s condition factor, corrected for relative condition [
49,
50]. Regression parameters were derived from non-parasitized individuals and compared to parasitized fish using
t-tests in Minitab v16. Stomach contents of
C. caballus were analyzed to assess dietary preferences and their relationship to parasite load. Prey items were identified to the lowest taxonomic resolution, grouped into categories, and quantified using the vacuity index (VI), frequency of occurrence (FA), and numerical frequency (FN) [
51,
52]. The Index of Relative Importance (IRI; [
53]) was calculated to classify prey groups as low (0–9.9%), secondary (10–40%), or high (40–100%) trophic relevance.
Comparisons of prevalence and abundance among sites, seasons, and host sex were performed using
t-tests, chi-square (χ
2), Fisher’s exact test, and Mann–Whitney (MW) U, implemented in Quantitative Parasitology QP v.3.0 [
54] and PAST v3.06. Relationships between prevalence and mean intensity were tested using Spearman’s rank correlation (rs). Aggregation patterns were evaluated using dispersion indices (variance-to-mean ratio K, Green’s IG, Morisita’s Im, and Lloyd’s mean crowding IMC) in PASSaGE v2 [
48,
55]. Diet–parasite linkages were explored by correlating prey categories with infection metrics to evaluate trophic transmission pathways. Statistical significance was set at α = 0.05.
4. Discussion
This study represents the first systematic characterization of the metazoan parasite community associated with
Caranx caballus in the northern Colombian Pacific, a region of high artisanal fishing activity but limited parasitological research. A total of 46 individuals were collected (18 during the dry season and 28 during the wet season), and we acknowledge that this seasonal imbalance constrains the robustness of statistical comparisons. Moreover, although our sampling effort achieved 74% representativity, the absence of a formal power analysis, due to logistical constraints inherent to artisanal fisheries, further limits statistical inference. Accordingly, our results should be interpreted as baseline information that provides a foundation for future expanded sampling. Despite these constraints, the dataset contributes novel baseline knowledge on parasite assemblages in
C. caballus and situates local artisanal fisheries within the broader Eastern Tropical Pacific (ETP) framework, where even modest datasets have proven valuable for understanding parasite diversity, stock discrimination, and ecosystem monitoring under climate variability [
13,
19,
21,
56,
57,
58].
The 74% representativity indicates that the effort was sufficient to capture most of the parasite diversity present in
C. caballus, although additional sampling would likely reveal further species and increase richness estimates, as reported in other tropical marine fish studies [
7,
13,
59]. Overall, the parasite assemblage exhibited low richness and moderate prevalence, dominated by copepods—particularly
Caligus sp.1—which emerged as the most abundant and prevalent species across seasons (
Table 2 and
Table 3). The detection of
Allopyragraphorus caballeroi with aggregated distribution patterns (
Table 4) underscores the heterogeneity of host–parasite interactions within this fishery. Importantly, these findings contribute novel baseline data for the northern Colombian Pacific and situate local artisanal fisheries within the broader ETP framework, where parasites are increasingly recognized as bioindicators of stock structure, ecosystem variability, and resilience under climate change [
13,
16,
57,
58,
60].
Despite significant differences in host diet composition and environmental conditions between sampling periods (
Table 5,
Table 6 and
Table 7), parasite communities exhibited seasonal stability in richness and prevalence. The strong reliance of
C. caballus on crustacean prey, especially brachyuran megalopa, suggests trophic transmission pathways that may explain the dominance of copepod parasites. Infection intensities were generally low, and host condition factors did not differ between parasitized and non-parasitized individuals (
Figure 2), indicating resilience of
C. caballus under current parasitic pressures [
58,
59,
60,
61].
These findings underscore the dual role of parasites as ecological indicators. Locally, they provide insights into host feeding ecology, stock discrimination, and transmission dynamics. Globally, they serve as sentinels of ecosystem change under climate variability. By linking parasite assemblages to diet, host condition, and oceanographic drivers (
Figure 4), this study situates artisanal fisheries of the Colombian Pacific within broader debates on sustainability, food security, and resilience in tropical marine ecosystems.
Of the 30 parasite species previously reported for C. caballus, we identified three (10%) in our samples. Notably, three copepods (Caligus sclerotinosus, Caligus sp., Caligus sp.1), two monogeneans (Microcotylidae sp. and Pseudomazocraes sp.), and one digenean (Stephanostomum sp.) constitute new records. All species reported here are the first for Colombia, thereby extending their known distribution within the Eastern Tropical Pacific (ETP).
Host condition was evaluated using Fulton’s condition factor, derived from length–weight relationships [
49,
50], and compared between parasitized and non-parasitized individuals. Although no significant differences were detected, this standardized approach provides a transparent baseline for assessing parasite influence on host health. Contrary to expectations from previous studies, no significant relationships were found between host size, sex, and parasite prevalence. Similar results were reported by Violante-González et al. [
13] in Mexico, where total length was unrelated to prevalence in two of three localities. The absence of such associations in our study may reflect the predominance of immature individuals, whose habitat use and resource exploitation differ from adults [
62,
63,
64]. Juvenile habitat partitioning can influence parasite exposure, as infracommunities and component communities are shaped by host resource use and trophic position [
12]. Given that prevalence and mean intensity were not linked to size, we suggest that fish across sampled sites and seasons did not exhibit significant differences in resource use, consistent with the >50% similarity in parasite composition between seasons.
Parasite communities at both infracommunity and component levels exhibited low richness, low diversity, and dominance by a single species. The categorization of low richness in the parasitofauna of
C. caballus from Colombia is relative and based on comparisons with published studies of carangid populations in the Mexican Pacific [
17,
56], which reported higher species counts and prevalence under similar sampling conditions. These benchmarks justify our classification of the assemblage as low richness, reflecting both the limited number of species detected and the dominance of a single taxonomic group. Component community richness did not vary significantly between seasons, differing from records for other carangids [
5,
14]. The addition of new species in season 2 likely reflects increased sample size rather than structural change, consistent with nested subset patterns described by Guegán & Hugueny [
65]. Such nestedness may arise from environmental influences on free-living infective stages or host distribution, leading to parasite gains or losses [
5,
66].
Comparisons with Mexican populations of
C. caballus [
13] revealed lower prevalence and mean intensity in Colombian fish, despite shared species (
A. caballeroi,
C. robustus). Parasite associations are rarely stable across host populations as communities are dynamic assemblages shaped by climate and latitude [
6]. Interestingly, our results contrast with [
7], who reported increased ectoparasite prevalence at lower latitudes, suggesting that local environmental conditions and host ecology may override latitudinal trends.
Copepods were the most represented group (56%), consistent with findings in Mexico [
13]. As one of the most diverse groups of marine fish ectoparasites, copepods, together with monogeneans, dominate parasite assemblages globally [
8]. Their prevalence in our study aligns with host diet, which was dominated by crustaceans, particularly brachyuran megalopa. This result is consistent with Saucedo-Lozano et al. [
35], although their January samples were dominated by cnidarians and mollusks, highlighting regional dietary variability. The predominance of crustaceans in both seasons suggests a relatively homogeneous prey supply, corroborated by Espinal-García et al. [
67], who reported year-round availability of brachyuran larvae in Colombian Pacific waters. Such resource homogeneity may limit exposure to intermediate hosts of endoparasites, explaining the absence of nematodes, acanthocephalans, and other taxa commonly reported in Mexican populations where diets included penaeid shrimps and adult fishes. The consumption of larval fishes in our samples may further explain the absence of nematodes, which typically infect intermediate hosts at later developmental stages [
68].
The strong reliance of
C. caballus on crustacean prey, especially brachyuran megalopa, suggests trophic transmission pathways that may explain the dominance of copepod parasites. Copepods typically have direct life cycles, attaching directly to hosts without intermediate hosts [
69,
70]. However, frequent ingestion of free-living crustaceans such as mysids and megalopa may increase exposure opportunities through shared habitats and trophic overlap, reinforcing the ecological link between host diet and parasite prevalence [
57,
59,
71]. This mechanism highlights how host feeding behavior directly shapes parasite community structure, consistent with reports in other tropical marine fishes where crustacean-rich diets are associated with higher copepod infection rates [
58,
61,
72]. Such findings align with broader evidence that caligid copepods exploit trophic pathways opportunistically, reflecting their ecological versatility and capacity to dominate parasite assemblages in crustacean-dependent fish population [
56,
60].
Spatial distribution analyses revealed contrasting patterns:
A. caballeroi exhibited aggregated distribution, which is consistent with monogeneans that remain on hosts and rapidly form dense populations [
73]. In contrast,
Caligus sp.1 displayed random distribution, likely reflecting its mobility and colonization strategy. Caligid copepods possess swimming appendages that enable dispersal across host populations, reducing the need for aggregation and minimizing intraspecific competition. Their presence in plankton samples supports this interpretation, highlighting their capacity to exploit hosts opportunistically across heterogeneous environments [
15].
Host-genus-specialist parasites often dominate communities, while most parasites behave as generalists [
74]. In this study, the dominant species was a copepod, a group generally recognized as taxonomic generalists capable of parasitizing diverse fish families. For example,
Caligus robustus has been reported in scombrids, lutjanids, and haemulids [
75]. The absence of central specialist species in our samples limits the predictability of parasite community structure in
C. caballus, a conclusion supported by the lack of correlation between prevalence and mean intensity. Beyond these structural patterns, parasites fulfill a dual role: they function as biological tags for stock discrimination [
13,
14,
15,
21,
56] and act as sentinels of ecosystem change under climate variability, with shifts in prevalence and assemblage composition reflecting broader environmental and trophic alterations [
4,
5,
6,
7,
8,
16,
17,
18,
66].
Environmental variation, particularly in temperature and salinity between seasons, may have influenced parasite diversity, even though overall community structure appeared temporally stable (
Table 7). These shifts in oceanographic structure are ecologically relevant. Warmer, saltier, and more stratified waters in the dry season may favor parasite transmission by enhancing host aggregation near the surface, whereas cooler, fresher, and more mixed conditions in the wet season could reduce parasite encounter rates and alter host feeding opportunities. Among monogeneans,
A. caballeroi has been reported from Peru in
Caranx hippos (likely misidentified, as this species does not occur in the Pacific, suggesting
C. caninus instead; FishBase, accessed 2 February 2025). This indicates a broad distribution across the ETP. Similarly, the genus
Pseudomazocraes (e.g.,
P. selene) has been reported in the Atlantic parasitizing
Selene vomer. These records challenge the expectation that monogeneans are strictly host-genus specialists [
76,
77]. In our study, the unidentified microcotylid and
Pseudomazocraes sp. may represent host-specific parasites of
C. caballus, but further taxonomic resolution is required.
The only endoparasite recorded was the digenean
Stephanostomum sp., typically found in the digestive tract, especially the intestine [
78,
79]. In this case, individuals were located in the esophagus, in a nearly external position. Their presence provides insights into the trophic role of
C. caballus, as digeneans require intermediate hosts to complete their life cycle. However, species-level identification and life-cycle studies are necessary to determine whether
C. caballus functions as a secondary intermediate host or as a definitive host in the region.
Taken together, the patterns observed in parasite prevalence, diversity, and aggregation, along with the associations between host condition and diet composition, underscore the multifaceted role of parasites in artisanal fisheries. These findings highlight how parasite assemblages not only reflect local ecological processes—such as trophic transmission and seasonal variability—but also provide signals of broader ecosystem dynamics influenced by climate drivers.