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Devil fish

The devil fish (Mobula mobular), also known as the giant devil ray or spinetail devil ray, is a large species of pelagic ray in the family , characterized by a diamond-shaped formed by its pectoral fins, forward-projecting cephalic fins resembling horns, and a slender tail tipped with a venomous . Reaching a maximum width of approximately 3.5 meters, it is one of the larger members of its genus, though not as massive as some manta rays. Found in circumglobal tropical to warm temperate waters, often in coastal and epipelagic zones, the devil fish migrates seasonally and forms large aggregations, exhibiting acrobatic breaching that propels it several meters above the surface. As a , it primarily consumes such as euphausiids (), copepods, and small crustaceans, occasionally supplementing with small pelagic fishes, using specialized gill rakers to strain food from water passed through its mouth. Classified as Endangered on the since 2006, with populations declining due to intense in industrial fisheries and targeted exploitation for its gill plates used in traditional Asian , the faces additional pressures from slow reproductive rates—females mature late and produce few —and habitat . Conservation efforts include international trade regulations under Appendix II and regional fishing bans, though enforcement remains challenging in key areas like the and eastern Atlantic. Its distinctive appearance has historically contributed to negative perceptions and incidental harm, yet recent research underscores its ecological role in food webs.

Taxonomy and nomenclature

Classification and synonyms

The devil fish is classified in the domain Eukarya, kingdom Animalia, phylum Chordata, subphylum Vertebrata, class , subclass , infraclass Neoselachii, superorder , order , suborder Myliobatoidei, family , genus , and species Mobula mobular (Bonnaterre, 1788). This placement reflects its status as a cartilaginous adapted for pelagic life, distinguished from benthic rays by features such as reduced and cephalic fins. Synonyms for M. mobular include Aodon japonicus, Apterurus fabroni, Cephaloptera fabroniana, Cephaloptera giorna, Cephaloptera japanica, Cephaloptera mobular, Cephalopterus giorna, Ceratoptera dumerilii, Mobula auriculata, Mobula cuvieri, Mobula eregoodootenke, Mobula fabroniana, Mobula jayakari, Mobula kuhlii, and Mobula rochebrunei, among others documented in taxonomic revisions. Notably, Mobula japanica (Müller & Henle, 1841) was synonymized with M. mobular in 2016 based on morphological and genetic analyses confirming conspecificity, resolving prior debates over versus Atlantic distributions. These synonyms arise from historical misidentifications due to variable regional morphologies and limited specimens, with modern classifications prioritizing molecular data for delimitation within . Common names for M. mobular encompass "devil fish," "giant devil ray," "spinetail devil ray," and regionally specific terms such as "jamanta" or "manta mobula" in Spanish-speaking areas, reflecting its distinctive cephalic lobes and tail spine that evoked demonic imagery in early European accounts. The name "devil fish" historically extended to other mobulids but is now primarily associated with M. mobular to denote its large size and breaching behavior, distinguishing it from the shorter-tailed Mobula hypostoma (Atlantic devil ray).

Etymology and historical naming

The common name "devil fish" applied to Mobula mobular derives from the species' distinctive cephalic fins, which project forward from the head and resemble horns, instilling a sense of dread in early European sailors and fishermen who encountered the ray's imposing form in and . This nomenclature emerged in maritime during the Age of Sail, associating the creature's appearance with infernal imagery and contributing to historical superstitions that portrayed it as malevolent, despite its harmless filter-feeding nature. The scientific binomial Mobula mobular traces its origins to Pierre Joseph Bonnaterre's 1788 description under the name Raia mobular within the skate genus Raia. In 1810, erected the specifically for this species, adapting the specific epithet mobular. The of mobular is uncertain, with scholars proposing derivations from Latin mobilis (mobile), referencing the ray's pelagic migrations; Italian vernacular tavola cornuta ("horned table"), evoking the flattened disc and cephalic lobes; or a regional Azorean term for the animal. These interpretations underscore the name's reflection of observed and behavior rather than precise linguistic roots.

Physical characteristics

Morphology and anatomy

The devil fish, , features a dorso-ventrally flattened body with a rhomboidal disc formed by broad, triangular pectoral fins that extend anteriorly to form the leading edge of the disc. The head is equipped with two prominent cephalic fins that project forward from the sides, resembling horns, and function to channel water and prey toward the mouth during filter feeding. The is small, subterminal, and positioned ventrally, framed by the cephalic lobes, with five gill slits bearing specialized filter plates on the branchial arches that enable the separation of from ingested . A small originates posterior to the trailing margin of the pectoral , often preceded by one or more s or prickles; the pelvic fins are small and ventral. The tail is slender, whip-like, and longer than the disc width, terminating in a serrated caudal located near the base, with additional prickles present. Dorsal coloration is predominantly dark gray to blue-black, frequently featuring a whitish band encircling the head and a black crescent-shaped stripe across the upper shoulders, while the ventral surface is white; the dorsal fin bears a white tip. As an elasmobranch, M. mobular possesses a cartilaginous endoskeleton and spiracles for water intake, supporting its pelagic lifestyle.

Size, growth, and sexual dimorphism

The spinetail devil ray (Mobula mobular) attains a maximum disc width (DW) of approximately 350 cm in adults, though recorded averages range from 180 to 280 cm DW. Newborns measure around 150 cm DW at birth. Males reach sexual maturity at a DW of 200–220 cm, typically between 5 and 6 years of age, while females mature at 215–240 cm DW over a similar timeframe. Growth is characteristically slow for this species, consistent with its k-selected life history strategy. Age and growth studies utilizing vertebral band analysis have estimated a von Bertalanffy growth coefficient (k) of 0.05 year⁻¹, indicating protracted development and low somatic growth rates that contribute to extended generation times of about 9.2 years. These parameters underscore the species' vulnerability to overexploitation, as population recovery relies on minimal annual productivity. Sexual dimorphism is minimal in M. mobular, with no pronounced differences in coloration, body shape, or overall size between mature males and females beyond subtle variations in maturity thresholds. Males possess claspers for internal fertilization, a standard elasmobranch trait, but females do not exhibit markedly larger maximum sizes relative to males in verified records. This lack of strong dimorphism aligns with observations across mobulid rays, where reproductive roles drive minor size disparities at maturity rather than extreme sexual size differences.

Distribution and habitat

Global range

The spinetail devil ray (Mobula mobular) has a circumtropical distribution, occurring in tropical and warm temperate waters of the Atlantic, Indian, and Pacific Oceans, as well as adjacent seas including the Mediterranean and Red Seas. This species is the only member of the family Mobulidae regularly inhabiting the Mediterranean Sea, where it is particularly abundant. In the Atlantic Ocean, it ranges along the eastern seaboard from to and in the western Atlantic from , , to . Indo-Pacific records extend from the and eastward to the , with northern limits reaching and southern extents to . The species is typically epipelagic, inhabiting coastal and oceanic waters over continental shelves and near islands, often linked to productive zones such as those off , , and . Populations exhibit seasonal migrations influenced by water temperature preferences between 18–27°C and oceanographic features, with records from latitudes approximately 40°N to 40°S. Despite its broad range, the ' pelagic lifestyle results in sparse, patchily distributed populations vulnerable to localized threats.

Habitat preferences and environmental tolerances

The spinetail devil ray (Mobula mobular) primarily inhabits epipelagic and neritic zones in coastal and oceanic waters of tropical, subtropical, and warm temperate seas across the , , and Pacific Oceans, often occurring near the surface in productive areas associated with or oceanographic fronts that concentrate planktonic prey. It exhibits a preference for dynamic environments with elevated chlorophyll-a concentrations (indicating high primary productivity) and shallower depths, which facilitate access to dense patches essential for filter-feeding. Depth preferences center on the upper , with frequent sightings and records from 0 to 200 meters, though individuals can tolerate excursions to at least 700–1,000 meters during foraging dives or migrations, reflecting adaptations for exploiting vertically migrating prey. In the , summer aggregations are documented in shallow coastal waters (typically <50 meters), correlating with seasonal warming and prey availability. Temperature tolerances span 13–28°C, with peak abundances in regions where sea surface temperatures average 18–26°C, such as the eastern tropical Pacific and Atlantic, where cooler fronts (<26°C) relative to surrounding waters enhance presence by promoting prey aggregation. The species avoids prolonged exposure to temperatures below 13°C or above 28°C, migrating seasonally to track optimal thermal regimes, as evidenced by data showing southward movements in the Mediterranean during cooler periods. Salinity tolerances align with fully conditions (approximately 34–36 ), with no documented estuarine incursions or adaptations; occurrences are confined to open ocean , underscoring its pelagic lifestyle and sensitivity to freshwater influences that could disrupt prey distribution or . Limited data suggest broad tolerance within oceanic salinity gradients, but preferences favor stable, high- environments without significant that might limit vertical prey access.

Ecology and behavior

Diet and feeding mechanisms

The spinetail devil ray (Mobula mobular) is primarily planktivorous, with its diet dominated by small such as euphausiids, particularly Nyctiphanes simplex in regions like the . Stomach content analyses from specimens in the eastern tropical Pacific and Indonesian waters confirm euphausiids as the principal prey, comprising the majority of identifiable items by frequency of occurrence, alongside minor contributions from copepods, mysids, and occasionally small fish or cephalopods. places M. mobular at a of approximately 3.6, consistent with a specialized pelagic filter-feeding strategy focused on low-energy-density prey, though dietary niche breadth suggests some opportunistic intake of larger crustaceans or fish larvae during aggregations. Feeding occurs via ram filtration, where the ray swims forward with its mouth agape, directing plankton-laden seawater over specialized branchial filter plates using forward-projecting cephalic lobes that funnel and accelerate flow. These plates, composed of cartilaginous lobes forming primary and secondary pores (0.05–0.80 mm wide across mobulids), employ ricochet separation—a hydrodynamic mechanism involving captive vortices within pores to redirect and retain particles as small as one-fifth the pore size while expelling water, minimizing clogging compared to traditional sieving. Cephalic lobe morphology, with angles of 25–39° in mobulids, influences prey selectivity, enabling M. mobular to target euphausiids ≥0.5 mm effectively during sustained open-mouth cruising. Behavioral adaptations enhance efficiency in patchy distributions: individuals perform somersaults (360° backward loops) or chain formations to concentrate prey in water columns, often near the surface or bottom where cephalic fins splay centrally to corral particles. Aggregations may facilitate cooperative funneling during cyclone-like spirals, though M. mobular typically feeds solitarily or in loose groups in pelagic waters, contrasting with more structured in rays. This low-overlap trophic niche underscores vulnerability to declines from or , as the species' specialized limits dietary flexibility.

Reproduction and life history

Mobula mobular exhibits aplacental viviparous , characterized by matrotrophy, where embryos derive nourishment from uterine secretions after depletion. Females typically produce a single large pup per reproductive cycle, with litters of two observed rarely. lasts approximately 12 months, though estimates suggest it may extend to 20–25 months in some . Newborn pups measure 90–180 cm in disc width at birth. The interbirth interval spans 1–3 years, reflecting a low reproductive output that limits potential. involves males pursuing females in aggregations, with behaviors including abdominal pressing and insertion of claspers, though detailed observations remain limited for this . Sexual maturity occurs at a disc width of approximately 4 m, with age-at-maturity estimates ranging from 4.5–9.1 years depending on and priors. Age at first is estimated at 5.4–12.6 years. is slow, with vertebral band counts indicating median lifespan estimates around 11.5 years, though maximum may exceed this based on capture-recapture data. This K-selected life history emphasizes high maternal in few offspring, rendering populations vulnerable to .

Social behavior, migration, and aggregations

Mobula mobular typically exhibits solitary behavior but forms aggregations for feeding and reproductive activities, with schooling observed in productive waters. involves multiple males (up to nine) pursuing one or two females in trains, displaying endurance, evasion, and pre-copulatory positioning stages; these events occur seasonally from May to August, peaking in May along Mexico's . The species undertakes seasonal migrations driven by environmental factors such as , chlorophyll-a concentrations, and availability. In the , from 2016 to 2021 tracked individuals covering up to 14,802 km over 330 days, with mean daily distances of 30.2 km (range 21.2–44.9 km); rays shift from winter aggregations in the (e.g., off ) to summer distributions in the western and central basins, including the , Canyons, and . Evidence suggests north-to-south movements of at least 1,800 km at minimum speeds of 63 km/day, linked to prey productivity. Aggregations, often comprising hundreds of individuals, facilitate key functions including feeding on dense zooplankton patches and courtship, with summer events in Important Shark and Ray Areas (ISRAs) and winter concentrations in the Levantine Sea showing potential sexual segregation (over 90% males in Gaza landings from 2014–2016). These gatherings increase vulnerability to fisheries but highlight site-specific hotspots for monitoring, such as those tied to upwelling and oceanographic fronts.

Human interactions

Fisheries exploitation and trade

is primarily encountered as in industrial purse-seine fisheries targeting tropical tunas in the eastern and , where it is captured alongside species such as skipjack and . Targeted exploitation occurs in smaller-scale artisanal fisheries along West African coasts and in the , using gillnets and driftnets to capture individuals aggregating near the surface. Landings data for M. mobular are often aggregated under spp. in official reports, with the (FAO) recording global mobulid catches exceeding 7,000 metric tons in 2018, an increase from prior decades, though underreporting in small-scale fisheries likely underestimates true exploitation levels. Fishing mortality rates for devil rays in regions like the , where M. mobular sightings occur sporadically, exceed sustainable thresholds, with exploitation ratios often surpassing 0.77 based on demographic models. The primary products from M. mobular fisheries are for local consumption in communities and rakers ( plates) for in the . rakers, valued in for purported health benefits despite lacking empirical evidence of efficacy, are dried and shipped primarily to markets in and via established networks. Trade volumes are difficult to quantify precisely for M. mobular due to species misidentification and lumping in commercial records, but surveys indicate that plates constitute the majority (up to 96%) of mobulid products in surveyed Asian markets, with annual global mobulid plate implying the harvest of over 100,000 individuals pre-CITES regulations. All mobulid species, including M. mobular, were listed under Appendix II in 2014, mandating export permits and non-detriment findings for to ensure sustainability. Despite this, illegal, unreported, and unregulated (IUU) persists, particularly within protected areas and through misreporting as permitted species, sustaining a shadow trade estimated to evade detection in post-listing market surveys. Regional organizations, such as the International Commission for the Conservation of Atlantic Tunas (ICCAT), have implemented release requirements for live mobulids since 2018, yet enforcement gaps and demand-driven continue to drive exploitation. Population modeling indicates that current pressures exceed intrinsic population growth rates for M. mobular, rendering ongoing trade unsustainable without stricter controls.

Historical and cultural uses

The common name "devil fish" for Mobula mobular stems from the horn-like cephalic fins protruding from its head, evoking demonic imagery in maritime folklore across and Atlantic cultures, where the species was historically viewed as a harbinger of misfortune or a threat to vessels due to its breaching and size. This perception persisted in accounts into the early 20th century, despite the rays' harmless nature lacking defensive spines or aggressive tendencies toward humans. Prior to industrialized fisheries, M. mobular was exploited in Mediterranean and eastern Atlantic coastal communities for its , consumed fresh or dried as a low-cost protein source in local diets, particularly in regions like where targeted artisanal capture dates back decades. was dried for leather-like applications, and served as a filler in soups or traditional preparations akin to shark fin substitutes. In broader mobulid traditions, gill plates from species including M. mobular entered Asian medicinal practices by the , processed into tonics (known as pengyusai) claimed to address ailments from to purported cancer relief, though empirical validation remains absent and demand escalated post-1970s. Unlike manta rays revered as protective spirits in lore, M. mobular lacks documented spiritual roles in Mediterranean or cultures, with uses centered on utilitarian rather than symbolic .

Conservation and threats

The spinetail devil ray (Mobula mobular) is classified as on the following a reassessment in October 2025, reflecting severe ongoing declines driven primarily by fisheries and targeted exploitation. This upgraded status from its previous Endangered listing applies to the three oceanic devil ray species, including M. mobular, due to inferred population reductions exceeding 80% over the past three generations across major ocean basins. Global population trends indicate sharp declines, with suspected reductions of 50–79% or more over the last three generations (approximately 30–45 years, given generation lengths of 10–15 years) based on fishery-dependent data, landings records, and observed rates in industrial purse-seine and gillnet fisheries. In regions like the Mediterranean, eastern Atlantic, and , local populations have collapsed by over 90% in some hotspots due to high exploitation rates exceeding the species' low intrinsic rate (r ≈ 0.02–0.05 per year), characterized by slow maturation (8–10 years), low (one pup per cycle), and long lifespan (up to 25+ years). No evidence of recovery exists, as current mortality rates from unmanaged fisheries continue to outpace biological productivity.

Primary threats and causal factors

The primary threats to Mobula mobular stem from intense fishing pressure, encompassing both targeted fisheries and in nets, purse seines, and trawls across its range in tropical and temperate waters. Directed fisheries exploit the species for its plates (rakers), which are harvested for use in traditional Asian soups believed to confer benefits, alongside consumption and in regions like the Mediterranean, , and Atlantic. mortality is exacerbated by the ray's tendency to aggregate in surface waters, making it vulnerable during seasonal migrations, with low post-release survival rates due to stress and injury. Causal factors amplifying these threats include the species' K-selected life history traits, such as slow growth, late maturity (reached at approximately 2.5–3 meters disc width), and low —one pup per or longer period—which result in low intrinsic rates (r ≈ 0.04–0.06) and prolonged recovery times from exploitation. has driven inferred global population declines exceeding 50% over three generations (approximately 60 years), with regional hotspots like the Mediterranean showing sharp reductions due to unregulated artisanal fisheries landing thousands annually despite international protections. Demand-driven trade, particularly for rakers, has surged since the early 2000s, fueled by markets in and , outpacing regulatory enforcement in source countries. Secondary factors, such as habitat degradation from coastal and entanglement in , contribute marginally but are overshadowed by direct , as evidenced by stable environmental tolerances in unfished areas. Enforcement gaps post-CITES Appendix II listing in 2016 persist, with illegal, unreported, and unregulated (IUU) fishing sustaining pressure, particularly on immature individuals that comprise a disproportionate catch share.

Protection efforts and efficacy

All mobulid ray species, including Mobula mobular, were listed under Appendix II of the Convention on International Trade in Endangered Species (CITES) effective September 14, 2014, requiring non-detriment findings and export permits for international trade in specimens such as gill plates and meat. They were also included in Appendices I and II of the Convention on the Conservation of Migratory Species (CMS) starting in 2011 for manta rays and extended to devil rays by 2014, obligating signatory nations to prohibit capture and pursue cooperative conservation. The IUCN Species Survival Commission's Shark Specialist Group published a Global Devil and Manta Ray Conservation Strategy in 2016, outlining priorities such as reducing bycatch, enforcing trade regulations, and enhancing monitoring through research and citizen science. Regional fisheries management organizations (RFMOs) have implemented no-retention policies; for instance, the Tuna (IOTC) Resolution 19/03, adopted in 2019, prohibits retention of mobulid rays caught in association with tuna fisheries and mandates live release "as soon as possible" to minimize harm. Similar measures exist under the International Commission for the Conservation of Atlantic Tunas (ICCAT) and Inter-American Tropical Tuna (IATTC), including requirements for full release of live specimens and reporting of interactions. Nationally, protections include outright bans on landing and trade in the since 2011, the since 1995 (extended to mobulids), and countries like and . Best-practice guidelines for handling and releasing bycaught mobulids, promoted by organizations like the , emphasize minimizing air exposure and injury to boost post-release survival rates, with studies showing survival exceeding 90% under optimal conditions versus lower rates with improper methods. Efforts to enhance efficacy include fisher-developed sorting grids for purse seine vessels, tested in the Eastern Tropical Pacific, which allow live release of while retaining target species, potentially reducing mortality if widely adopted. Regional projects, such as the Conservation Project in the Eastern Tropical Pacific and Mediterranean-focused initiatives by groups like Save the Med, focus on mitigation, genetic monitoring, and mapping to inform targeted protections. Despite these measures, efficacy remains limited, as evidenced by ongoing population declines and a 2025 IUCN reassessment upgrading M. mobular and other oceanic devil rays to from Endangered, reflecting insufficient reversal of threats like in industrial fisheries. persistence in RFMO areas, with reports of over 90% declines in some regional s, stems from inconsistent enforcement, inadequate handling in high-seas fisheries, and illegal trade evading controls. No global abundance estimates exist, but diver surveys indicate declining sightings in nearly half of monitored regions over the past decade, underscoring the challenges of protecting highly migratory, low-productivity species amid transboundary exploitation. National measures often fail without complementary international action, as mobulids encountered in distant-water fleets bypass local bans.

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