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Spanish dancer

The Spanish dancer (Hexabranchus spp.), commonly referred to as H. sanguineus in older literature but now recognized as a species complex of at least five species following a 2023 taxonomic review,^[1] also known as the bloody sea goddess, is a large and vividly colored dorid nudibranch, a shell-less marine gastropod mollusk belonging to the family Hexabranchidae, renowned for its graceful swimming behavior that mimics the undulating movements of a flamenco dancer.^[2]^[3] Native to tropical and subtropical waters of the Indo-Pacific region, it inhabits coral reefs, rocky substrates, and sandy or rubble bottoms from the intertidal zone down to depths of at least 50 meters, often resting camouflaged during the day and becoming active at night.^[2]^[3] This taxon, first described as Doris sanguinea by Rüppell and Leuckart in 1830, can grow to impressive sizes, with adults reaching up to 40–60 cm in length, making it one of the largest known nudibranchs and among the world's biggest sea slugs.^[2]^[3] Its body features a broad, ruffled mantle with wide parapodia (lateral flaps) that display a mottled pattern of red, white, cream, yellow, and pink hues, while six bushy gills arranged in a rosette protrude from the dorsal surface—hence the genus name meaning "blood-colored six-gills."^[2] When threatened, the Spanish dancer propels itself through the water by rhythmically flapping its parapodia, a rare swimming ability among nudibranchs that allows relatively fast escape over short distances.^[3]^[2] Ecologically, Hexabranchus spp. are generalist feeders, primarily consuming a variety of sponges from at least 11 genera across five orders, though they occasionally ingest algae, coral mucus, and other organic matter; they sequester toxic chemicals from their sponge diet to deter predators, enhancing their own chemical defenses.^[3]^[2] As simultaneous hermaphrodites, individuals mate year-round, laying elongated pink or red egg ribbons that can contain millions of eggs and hatch into planktonic veliger larvae after 6–12 days, depending on water temperature.^[3] Found from East Africa through the Indian Ocean to the western Pacific (including Hawaii and French Polynesia), populations exhibit some color variation by region, but the complex faces no specific conservation threats and is not currently assessed by the IUCN.^[2]^[3]

Taxonomy

Classification

The Spanish dancer, Hexabranchus sanguineus, belongs to the kingdom Animalia, phylum Mollusca, class Gastropoda, subclass Heterobranchia, order Nudibranchia, suborder Doridina, superfamily Chromodoridoidea, family Hexabranchidae, genus Hexabranchus, and species H. sanguineus.^[4] The family Hexabranchidae, established by Bergh in 1891, is monotypic and encompasses only the single genus Hexabranchus.^[5] Recent integrative taxonomic studies, combining molecular and morphological data, have clarified the genus's diversity, recognizing at least six valid species, including the Indo-Pacific H. sanguineus and the Caribbean H. morsomus (previously placed in the synonymized genus Caribranchus), which is now confirmed as a distinct species based on radular differences and genetic divergence.^[6] Hexabranchus sanguineus was originally described as Doris sanguinea by Rüppell and Leuckart in 1830 from specimens collected in the Red Sea.^[4] Subsequent synonymy includes names such as Albania formosa Collingwood, 1881, and Doris imperialis Saville-Kent, 1897, reflecting early taxonomic confusion due to variable coloration and distribution.^[4] Hexabranchidae is distinguished from closely related nudibranch families like Chromodorididae—both within Chromodoridoidea—primarily by the unique arrangement of its six gills, which are individually inserted into the dorsal body wall rather than emerging from a shared vascular trunk as in chromodorids.^[2] This gill morphology, along with the branched, multi-pinnate structure of each gill, underscores the family's specialized evolutionary position among dorid nudibranchs.^[6]

Etymology

The common name "Spanish dancer" for Hexabranchus sanguineus originates from the nudibranch's distinctive frilly, red-and-white-edged mantle and its graceful, undulating swimming motion, which evoke the flowing dress and movements of a flamenco dancer.^[2] This descriptive moniker highlights the species' vibrant appearance and behavior, making it one of the few nudibranchs widely recognized by a common name in global marine biology contexts.^[7] The scientific name Hexabranchus sanguineus, established in its current form through taxonomic revisions, breaks down etymologically as follows: "Hexa-" derives from the Greek word for "six," referring to the six prominent branchial plumes or gills; "branchus" comes from the Greek "branchia," meaning gills; and "sanguineus" is Latin for "blood-red," describing the animal's typical deep crimson coloration.^[8] The genus Hexabranchus was originally erected by Ehrenberg in 1828, with the species first described as Doris sanguinea by Rüppell and Leuckart in 1830 from specimens collected in the Red Sea.^[8] This naming reflects key morphological features observed in the type locality, and the full binomial has been stabilized as a nomen protectum under the International Code of Zoological Nomenclature to preserve nomenclatural stability amid historical synonymy.^[8] The name's popularity in Indo-Pacific marine biology texts stems from the species' striking red hue and dramatic form, often likened to elements of Spanish cultural imagery such as a dancer's attire, contributing to its iconic status in popular and scientific accounts.^[2]

Description

Morphology

The Spanish dancer (Hexabranchus sanguineus) is a dorid nudibranch with a soft, flattened body and a broad, oval mantle that forms a skirt-like extension around the foot when extended. Unlike many gastropods, it lacks a protective shell and a traditional ctenidium, instead featuring typically six (occasionally varying to five or seven) large, feathery branchial plumes arranged in a rosette pattern around the anal opening; these plumes serve primarily for gas exchange.^[9] Adults typically measure 20–40 cm in length, though specimens up to 60 cm have been reported, with the mantle reaching widths of approximately 30 cm; juveniles are considerably smaller, often 5–10 cm long.^[9] Key external structures include modified oral tentacles that appear hand-like, consisting of a central lobe flanked by several finger-like digitate projections for manipulation, and a pair of retractile, club-shaped rhinophores positioned anteriorly for chemosensory detection of environmental cues. The dorsal mantle surface is generally smooth, lacking prominent warty projections (verrucae), which contributes to its streamlined form.^[9] Internally, H. sanguineus possesses a hermaphroditic reproductive system with distinct male and female openings on the right side of the body, allowing for reciprocal fertilization during mating. The digestive system is specialized for a spongivorous diet, featuring a protrusible proboscis and a broad, bilobed radula equipped with simple, hamate teeth that increase in length laterally; the central portion of the radular ribbon lacks teeth, facilitating rasping and ingestion of sponge tissue.^[9]

Coloration and patterns

The Spanish dancer, Hexabranchus sanguineus, displays a vivid primary coloration on its mantle that typically ranges from bright red to orange, creating a highly conspicuous appearance in Indo-Pacific coral reef environments. This coloration is often adorned with opaque white spots and lines, which form irregular patterns, including marginal bands and flecks concentrated on the posterior end and edges of the mantle. The branchial plumes, consisting of six multi-pinnate tufts, are frequently white-tipped or exhibit white with red lines, enhancing the overall striking visual profile.^[8]^[7] These color patterns serve an aposematic function, acting as a warning signal to visually oriented predators about the animal's unpalatability, with the pigments derived from defensive chemicals sequestered from its sponge diet, such as macrolides from species like Halichondria sp.^[10] Unlike some marine organisms, H. sanguineus lacks bioluminescence, relying solely on its static pigmentation for visual deterrence.^[8] Coloration exhibits notable variation across geographic ranges and developmental stages. Populations in the Red Sea tend toward deeper blood-red hues, while those in the Western Indian Ocean are more orange, French Polynesia features dark red with white rosettes, and Western Pacific specimens show red with clustered white flecks. Ontogenetically, juveniles are translucent in gray, pink, or yellow tones, gradually darkening and developing the full red pigmentation and white markings as they mature.^[8]

Distribution and habitat

Geographic range

The Spanish dancer (Hexabranchus sanguineus) has a primary geographic range spanning the tropical Indo-West Pacific, extending from the Red Sea and East Africa eastward to Hawaii, Japan, and Australia.^[11]^[3] This distribution encompasses latitudes from approximately 37°N to 36°S and longitudes from 30°E to 144°W, with individuals typically occurring at depths of 1–50 meters, usually 10–20 meters.^[12]^[11] Within this range, the species is commonly reported from key locations in the Indian Ocean, including the Maldives and Seychelles, as well as the Coral Triangle region encompassing Indonesia and the Philippines, and various Pacific islands such as those in French Polynesia.^[13]^[14]^[15] These areas represent biodiversity hotspots where sightings are frequent, particularly in association with coral reefs.^[3] Historically, H. sanguineus was first described from specimens collected in the Red Sea by Rüppell and Leuckart in 1830, establishing the western extent of its known distribution.^[16]^[17] The species is distinct from the Caribbean H. morsomus, which occupies the tropical western Atlantic and does not overlap with the Indo-Pacific range.^[18] Citizen science platforms like iNaturalist provide ongoing verification of range stability, with observations as recent as 2025 confirming consistent presence across the Indo-West Pacific without evidence of significant expansion or contraction.^[19]

Environmental preferences

The Spanish dancer, Hexabranchus sanguineus, inhabits shallow coral reefs, rocky subtidal zones, and areas of rubble or sand within tropical marine environments. These habitats provide structural complexity for shelter and proximity to food sources, with individuals often observed on fringing reef platforms.^[3]^[11] This species thrives in warm tropical waters with temperatures ranging from 24.7°C to 29.3°C, reflecting its adaptation to Indo-Pacific reef conditions. It exhibits nocturnal activity patterns, remaining hidden in reef crevices during daylight and becoming more active in low-light conditions at night to forage and reduce predation risk.^[11]^[20] Microhabitats favored by H. sanguineus include surfaces on or near encrusting sponges, which serve as prey and contribute to the nudibranch's chemical defenses. The species inhabits typical marine salinities consistent with its reef-associated lifestyle.^[3]^[11]^[10] As a benthic crawler, H. sanguineus navigates reef substrates but can transition to pelagic swimming when disturbed, aiding escape; most occurrences are between 10–20 m where conditions support sponge abundance.^[3]^[2]

Biology

Locomotion and behavior

The Spanish dancer, Hexabranchus sanguineus, primarily locomotes by crawling along substrates using its muscular foot, a typical mechanism for dorid nudibranchs that allows slow, deliberate movement over reef surfaces.^[2] When undisturbed, it typically folds its mantle over its back to facilitate this crawling while providing camouflage.^[2] For more rapid escape, it employs swimming via undulating waves of its expansive mantle, which flaps like wings to generate thrust through dorso-ventral angular deflections and propagating waves along the body.^[21] This swimming motion, often sustained for over an hour, propels the animal at terminal speeds of up to 1.33 body lengths per cycle, equivalent to approximately 0.67 body lengths per second assuming a 2-second cycle.^[21] The undulating display during swimming bears a resemblance to the flowing movements of a flamenco dancer, from which the common name derives.^[2] Behaviorally, H. sanguineus exhibits a nocturnal rhythm, remaining hidden during the day in crevices or under rocks and becoming active at night, as evidenced by frequent observations during night dives.^[2] It shows no territoriality, with individuals generally solitary except for occasional mating pairs, and aggregations are rare, though small groups may form in areas of high sponge density.^[2] In response to disturbance, the animal initiates swimming to flee, often displaying its bright red and white coloration in a deimatic manner.^[2] Sensory behaviors rely on specialized structures for navigation and environmental interaction. The retractable rhinophores serve as chemosensory organs, detecting chemical cues in the water column to guide movement.^[2] The branchial plumes, consisting of six exposed gills attached directly to the dorsal surface without a protective pouch, contribute to orientation during swimming by sensing water flow and aiding in postural stability amid the undulations.^[2]

Diet and feeding

The Spanish dancer, Hexabranchus sanguineus, is a generalist sponge feeder, consuming a variety of sponge species from at least 11 genera across five orders, including Halichondria spp.^[2]^[3] This diet provides essential nutrients while allowing the nudibranch to sequester defensive chemicals from toxic sponges, enhancing its own protection against predators.^[22] Preference for chemically defended prey ensures the incorporation of bioactive compounds like oxazole-containing macrolides into its tissues.^[2]^[23] Feeding occurs nocturnally, with the nudibranch protruding its buccal mass via a protractile proboscis to access sponge surfaces.^[24] It then employs its radula—a chitinous, toothed ribbon-like structure—to rasp and scrape sponge tissue, ingesting fragments that include skeletal spongin fibers and embedded toxins.^[25] Digestion takes place in the stomach, where enzymes break down the material to extract nutrients and defensive metabolites, such as macrolides derived from the sponge diet; these chemicals are stored for the nudibranch's defense and even transferred to egg ribbons.^[22]^[26] In its trophic role, H. sanguineus functions as a grazer on sessile sponges, contributing to reef ecosystem dynamics without evidence of cannibalistic behavior among conspecifics.^[2] This sponge-based intake supports substantial somatic growth, enabling individuals to reach lengths of up to 60 cm.^[27] Juveniles target smaller sponge fragments or encrusting forms suitable to their size, while adults consume larger patches.^[28]

Reproduction and development

The Spanish dancer, Hexabranchus sanguineus, is a simultaneous hermaphrodite, possessing both male and female reproductive organs within a single gonad located on the right side of the body.^[29] Internal fertilization occurs through reciprocal insemination, where mating pairs exchange spermatophores via their right-sided gonopores, allowing both individuals to function as donor and recipient simultaneously.^[29]^[2] No specific mate choice behaviors have been observed, and reproduction takes place year-round in tropical environments.^[2] Following mating, individuals deposit eggs in large, pink, ribbon-like masses attached to substrates such as coral or rocks, often near feeding areas; these consist of tightly coiled spirals forming rosette shapes typically 2–20 cm in diameter, containing numerous capsules each holding 8 to 44 eggs.^[29] Fecundity is high, with each mass containing an estimated 1.5 to 3.6 million eggs, reflecting the species' strategy to maximize offspring production in planktonic dispersal; the egg ribbons also incorporate defensive macrolides sequestered from the sponge diet.^[29]^[22] Development begins with trochophore larvae appearing by approximately day 6 after spawning (at 25–28°C), followed by hatching of planktonic veliger larvae after 8–9 days of incubation.^[29] These veligers, with shell lengths of 170–200 μm, remain in the pelagic phase for 20 to 30 days, feeding on phytoplankton before undergoing metamorphosis into juvenile forms that settle on the benthos.^[29] During this period, the larvae grow, developing characteristic features such as a translucent shell and digestive gland.^[29]

Ecology

Defenses and predators

The Spanish dancer (Hexabranchus sanguineus) employs chemical defenses derived from its sponge diet, sequestering oxazole-containing macrolides from species of Halichondria that render its tissues unpalatable to potential predators.^[30] These compounds, concentrated in the dorsal mantle and digestive gland at levels of 0.14–0.62% dry weight, impart a bitter taste that deters fish and invertebrates.^[30] The nudibranch modifies some dietary macrolides during digestion, enhancing their defensive efficacy while passing them to egg ribbons for protection of offspring.^[30] In addition to chemical protection, the Spanish dancer relies on physical evasion strategies, primarily rapid swimming using undulating movements of its mantle and parapodia when disturbed, allowing it to flee threats in open water.^[31] Its bright red coloration serves as an aposematic warning signal, advertising toxicity to visually oriented predators.^[9] Documented predators of the Spanish dancer are few, reflecting the effectiveness of its defenses; observed attackers include wrasses such as Thalassoma lunare (Labridae) and hermit crabs like Dardanus megistos.^[30] Some crabs and generalist reef fish may attempt predation, but encounters are rare due to the nudibranch's unpalatability and conspicuous warning signals.^[32] Field observations indicate high survival rates for the Spanish dancer in natural habitats, with low incidence of predation attributed to its multifaceted defenses. Laboratory assays demonstrate that its macrolides elicit rejection by predators at minimum effective concentrations of 0.01–0.02% dry weight in food pellets, resulting in near-complete avoidance after initial tasting.^[30]

Conservation status

The Spanish dancer (Hexabranchus sanguineus) has not been evaluated by the International Union for Conservation of Nature (IUCN) Red List as of 2025, rendering its global conservation status unknown and highlighting significant data deficiencies in population assessments.^[3]^[11]^[33] Primary threats to the species stem from anthropogenic pressures on its coral reef habitats, including coral bleaching driven by rising sea temperatures and ocean acidification, which degrade the structural complexity essential for sponge prey availability.^[34] Overfishing disrupts reef ecosystems by reducing herbivorous fish populations that maintain algal balance, indirectly affecting sponge habitats, while coastal pollution from agricultural runoff and wastewater introduces nutrients and toxins that diminish sponge diversity and health.^[35]^[34] Climate change exacerbates these issues by potentially shifting suitable temperature ranges, leading to range contractions in vulnerable tropical regions.^[36] Population trends are currently unknown, with no evidence of global declines or threats from human activities,^[3]^[37] though citizen science initiatives, including the Reef Life Survey, document infrequent but consistent occurrences across surveyed sites (3.9% of locations), aiding in monitoring but underscoring the need for expanded data collection to address knowledge gaps.^[33] Conservation measures focus on broader reef protection, with the species indirectly benefiting from marine reserves such as those in the Great Barrier Reef Marine Park, where restrictions on fishing and pollution help preserve sponge habitats. Recommendations emphasize sustainable diving practices to minimize disturbance and targeted efforts to restore sponge populations through pollution reduction and reef rehabilitation programs.^[25]^[19]

References

[1]
Hexabranchus sanguineus - The Sea Slug Forum
Hexabranchus sanguineus is a nudibranch found in the Tropical Indo-West Pacific, can swim, has unique gill structure, and can grow over 40cm long.
[2]
Spanish Dancers, Hexabranchus sanguineus
Spanish dancer nudibranchs, Hexabranchus sanguineus (Rüppell & Leuckart, 1830), are large and colourful nudibranchs (sea slugs with naked gills)
[3]
https://www.marinebio.org/species/spanish-dancers/hexabranchus-sanguineus/
[4]
https://www.marinespecies.org/aphia.php?p=taxdetails&id=220381
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[6]
Hexabranchus sanguineus - The Sea Slug Forum
Mar 26, 2010 · When they are small they are often mistaken for a species of Chromodoris. It's a bit hard to see but one way small Hexabranchus can be ...Missing: distinction | Show results with:distinction
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a review of the family Hexabranchidae Bergh, 1891 (Gastropoda ...
Jun 20, 2023 · The genus Caribranchus is considered a junior synonym of Hexabranchus Ehrenberg, 1828 and the ontogeny of color pattern is discussed. Similar ...Missing: roseus | Show results with:roseus
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Hexabranchus sanguineus - The Sea Slug Forum
Hexabranchus sanguineus is a nudibranch from the Tropical Indo-West Pacific, can swim, has unique gill structure, and can grow over 40cm long.
[9]
Spawn and Early Larval Development of Spanish Dancer ...
Hexabranchus sanguineus is best known as the Spanish dancer was found predominantly on the shallow fringing reef platforms all around Hurghada.Missing: literature | Show results with:literature
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Defensive chemicals of the Spanisch dancer nudibranch ...
In a reef environment dominated by visually oriented predators, the striking color pattern and behavioral responses of Hexabranchus may have arisen with a ...
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Hexabranchus sanguineus, Spanish dancer : fisheries - SeaLifeBase
The Spanish dancer, *Hexabranchus sanguineus*, is a benthic nudibranch found in the Indo-Pacific, 1-50m, with a max length of 25cm, and a unique swimming ...
[12]
Hexabranchus sanguineus - Spanish dancer - Snorkeling Report
Hexabranchus sanguineus. Spanish dancer. Distribution: from the Red Sea and South Africa to Southern Japan, Hawaii, and Australia.
[13]
Spanish Dancer? from the Maldives - The Sea Slug Forum
An animal that large, and red, is most probably a Spanish Dancer [Hexabranchus sanguineus]. If you look through the messages on the Hexabranchus page you will ...Missing: Seychelles | Show results with:Seychelles
[14]
Nudibranch from the Seychelles, Indian Ocean - The Sea Slug Forum
This is a form of Hexabranchus sanguineus. Have a look at the Fact Sheet and associated messages for further information.
[15]
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Dec 7, 2017 · The species diversity of marine heterobranch sea slugs found on field trips around Bunaken Island (North Sulawesi, Indonesia) and adjacent ...
[16]
Hexabranchus sanguineus from Red Sea - The Sea Slug Forum
This animal was first described from an animal collected in the Red Sea. The name 'sanguineus' means 'bloody' or 'blood red', which is very appropriate for ...
[17]
Description of Hexabranchus sanguineus : Reunion island (Indian ...
According Bill Rudman : " There is no easy way to distinguish it externally as a Hexabranchus at this small size because the characteristic oral lobes and ...Missing: distinction | Show results with:distinction
[18]
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marginata-grandifolia is considered a Lessepsian migrant and an alien species in the Mediterranean ... The nudibranch known as the Spanish dancer, Hexabranchus ...
[19]
A Spanish dancer? No! A troupe - iNaturalist Canada
Jun 22, 2023 · The nudibranch known as the Spanish dancer, Hexabranchus sanguineus, is a classic example with over 200 years of taxonomic confusion.
[20]
Spanish Dancer (Hexabranchus sanguineus) - iNaturalist
The Spanish dancer, *Hexabranchus sanguineus*, is a large, colorful dorid nudibranch, a marine gastropod mollusk, meaning 'blood-colored six-gills'.
[21]
Spanish Dancer (Hexabranchus sanguineus) - Red Sea Creatures
The Spanish Dancer is a vibrant sea slug, up to 25cm long, with a unique swimming style. It is nocturnal, feeds on sponges, and uses chemical defenses.
[22]
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[23]
Defensive chemicals of the Spanisch dancer nudibranch Hexabranchus sanguineus and its egg ribbons: macrolides derived from a sponge diet
### Summary of Coloration and Defensive Role in Hexabranchus sanguineus
[24]
[PDF] macrolides derived - UNCW
In a reef environment dominated by visually oriented predators, the striking color pattern and behavioral responses of Hexabranchus may have arisen with a ...
[25]
[PDF] The Book David A. Mullins 2023 – Diet, Feeding and Digestion
dorid Hexabranchus sanguineus shows us its buccal mass protruded and feeding upon a sponge colony. ... Their digestive systems in general follow a typical sea ...
[26]
Spanish Dancer | Life in Water Wiki | Fandom
The Spanish Dancer (Hexabranchus sanguineus) is a large and visually ... Reproduction: Spanish Dancers are hermaphroditic, possessing both male and female ...
[27]
Defensive chemicals of the Spanisch dancer nudibranch ...
Slices of the dorsal mantle tissue of the nudibranch were rejected in laboratory feeding assays employing two common sympatric predators: an Indo-Pacific reef ...Missing: verrucae projections<|control11|><|separator|>
[28]
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[29]
Feeding Hexabranchus sanguineus from Tofo, Mozambique
Hexabranchus is reported to feed on quite a few different species of sponges ... Juvenile Hexabranchus sanguineus from Christmas Id From: W.B. Rudman, May 10 ...Missing: seasonal | Show results with:seasonal
[30]
The effect of environmental factors on spatial-temporal variation of ...
Nov 9, 2022 · Furthermore, temperature and food sources may differ between seasons and depths, thereby influencing sea slug assemblages in this region.<|control11|><|separator|>
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Aug 9, 2025 · Edmunds (1968) suggested a difference in swimming behavior between H. lacer (as H. marginatus) and H. sanguineus but, as pointed out by Thompson ...Missing: locomotion | Show results with:locomotion
[33]
9 Spanish Dancer Sea Slug Facts - Fact Animal
In the tropical reefs around the world there's a branch of these slugs, many of whom are agile predators, feeding on sponges and attaching strange egg roses on ...
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Hexabranchus sanguineus - spanish dancer nudibranch
Information. Max Size: 60 cm. Sea Temperature Range: 19.1-28.3°C. Depth: N/A. Habitat Generalization Index: N/A. Also referred to as the SGI (Species ...
[35]
Threats to Coral Reefs | US EPA
Feb 9, 2025 · Overfishing can alter food-web structure and cause cascading effects, such as reducing the numbers of grazing fish that keep corals clean of ...Missing: Hexabranchus sanguineus
[36]
How does overfishing threaten coral reefs?
Jun 16, 2024 · The impacts from unsustainable fishing on coral reef areas can lead to the depletion of key reef species in many locations.Missing: Hexabranchus sanguineus
[37]
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Learn about coral bleaching, its causes, effects on the Great Barrier Reef, and how you can help combat this environmental threat.
[38]
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May 3, 2021 · Population trends of Spanish Dancers are unknown, but there is no evidence that human activities threaten the species. It is important ...