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Remora

Remoras, members of the family Echeneidae in the order Carangiformes, are a group of eight species of marine fishes distinguished by their specialized first dorsal fin, which is modified into an oval-shaped suction disc on the top of the head, enabling them to adhere firmly to larger hosts such as sharks, rays, sea turtles, cetaceans, and occasionally marine vessels.^[1]^[2] These hitchhiking fish exhibit a slender, elongated body with a flattened head, long-based dorsal and anal fins, and a protruding lower jaw, typically reaching lengths of 30 to 110 cm depending on the species.^[2]^[3] Distributed worldwide in tropical and warm temperate oceanic waters, often from coastal zones to depths exceeding 100 m, remoras lead an obligate commensal lifestyle, attaching to hosts for transportation, protection from predators, and opportunistic feeding on ectoparasites, food scraps, and plankton dislodged from the host's body.^[4]^[5]^[6] The suction disc, featuring transverse laminae with tooth-like structures, allows secure attachment with low energy cost, as the mechanism relies on hydrodynamic forces rather than constant muscular effort once engaged.^[5]^[6] This adaptation supports their pelagic existence, where juveniles and adults rarely swim independently and instead migrate across oceans via host associations.^[1]^[5] Ecologically, remoras play a role in marine symbioses that range from mutualistic—by removing parasites from hosts—to potentially parasitic under certain conditions, though net interactions are typically beneficial or neutral for the host.^[7] Host specificity varies among species; for instance, the common remora (Remora remora) prefers sharks and bony fishes, while the whalesucker (Remora australis) specializes in cetaceans.^[8]^[6] Their reproductive biology involves pelagic spawning, with eggs and larvae developing in open water before the disc forms and juveniles seek hosts, contributing to their wide dispersal in warm seas.^[9]

Taxonomy and evolution

Taxonomy

Remoras are classified in the family Echeneidae, which is included in the order Carangiformes according to phylogenetic analyses incorporating molecular data from studies after 2010.^[10]^[11] This placement reflects the family's close relation to carangoid fishes, with the suction disc representing a key derived trait.^[12] The term "remora" originates from the Latin remorari, meaning "to delay" or "hinder," alluding to historical accounts of these fish supposedly slowing ships by adhering to their hulls.^[13]^[14] Echeneidae encompasses three genera—Remora (five species), Echeneis (two species), and Phtheirichthys (one species)—for a total of eight extant species, all characterized by a modified dorsal fin forming an adhesive disc.^[15]^[16] The following table lists these species, including common names, maximum reported lengths, and brief distinguishing traits:

Genus	Species	Common Name	Max Length (TL unless noted)	Distinguishing Traits
Remora	R. remora	Common remora	86 cm	Uniformly dark gray to brown body; broad disc with 21–27 laminae; primarily attaches to sharks and turtles.^[14]^[17]
Remora	R. albescens	White suckerfish	35 cm	Short, wide disc (34–40% of SL); pale body; associates with mantas, sharks, and marlins.^[18]
Remora	R. australis	Whalesucker	76 cm	Elongate body; large disc with 25–28 laminae; specializes in attaching to whales and dolphins.^[19]
Remora	R. brachyptera	Spearfish remora	50 cm	Moderate disc with 18–22 laminae; slender form; adheres to billfishes like spearfish and swordfish.^[20]^[21]
Remora	R. osteochir	Marlin sucker	40 cm (SL)	Small disc with 15–19 laminae; pointed pectorals; attaches to marlins and sailfishes.^[22]
Echeneis	E. naucrates	Live sharksucker	110 cm	Slender body; long disc with 32–40 laminae; versatile host attachment including live sharks and rays.^[4]
Echeneis	E. naucratoides	Whitefin sharksucker	75 cm	Dark lateral band with white borders; disc with 25–33 laminae; broad host range including rays and turtles.^[23]
Phtheirichthys	P. lineatus	Slender suckerfish	76 cm	Bluish-black above, white below; three longitudinal stripes; rare, attaches to large fishes and turtles.^[24]^[25]

Evolutionary history

The fossil record of remoras (family Echeneidae) begins in the late Oligocene to early Miocene epochs, with the earliest known specimens dating to approximately 30 million years ago. The oldest described fossil remora is the stem-group taxon †Opisthomyzon, recovered from Rupelian (early Oligocene) deposits in the Engi Slate of the Canton of Glarus, Switzerland, which preserves an incomplete but diagnostic adhesion disc formed from modified dorsal-fin elements.^[26] Subsequent Miocene fossils, such as †Echeneis urupensis from the Maikop Formation in the North Caucasus region of Russia (dated to around 20 million years ago), indicate the presence of more crown-like echeneids with advanced disc structures, suggesting the family had diversified by the early Neogene.^[27] These fossils document a relatively sparse record, with no known remora remains from the Eocene or earlier, highlighting a relatively recent origin for the group compared to their percomorph relatives.^[26] Phylogenetically, remoras are nested within the order Carangiformes, specifically in the superfamily Echeneoidea, where Echeneidae forms a monophyletic clade sister to Rachycentridae and Coryphaenidae.^[10] Molecular phylogenomic analyses, including ultraconserved element (UCE) data from multiple loci, confirm this placement and support derivation from carangiform ancestors, with the family's specialized traits evolving as adaptations for ectoparasitic or commensal symbiosis on marine hosts.^[1] Morphological and genomic evidence indicates that the suction disc, a defining feature, arose through the transformation of the first dorsal fin into a multi-segmented adhesive organ, a specialization absent in outgroup carangiforms like carangids. A key evolutionary milestone was the modification of the dorsal fin into a functional sucker disc, evidenced in †Opisthomyzon around 30 million years ago, which likely facilitated attachment to mobile hosts such as sharks and rays during a period of marine ecosystem diversification in the Oligocene-Miocene transition. This innovation is linked to the radiation of potential host taxa, including elasmobranchs, whose lineages predate remoras but whose abundance increased in Neogene seas, enabling the symbiotic lifestyle that characterizes modern echeneids.^[10] The resulting evolutionary radiation has produced the eight extant species across three genera, reflecting adaptive specialization to diverse hosts.^[1]

Description and physiology

Physical characteristics

Remoras, belonging to the family Echeneidae, have an elongated, torpedo-shaped body with a flat ventral surface that supports their streamlined form. Their skin is covered in small, cycloid scales, contributing to a relatively soft texture. Body lengths typically range from 30 to 90 cm, with the maximum recorded up to 110 cm in species like the live sharksucker (Echeneis naucrates).^[28] These fish display dark coloration, varying from gray to black, which is nearly uniform across the dorsal and ventral surfaces. They possess a small terminal mouth, with the lower jaw extending beyond the upper jaw, and the head comprising about one-fifth of the total body length.^[28]^[4] Remoras feature a single dorsal fin, the anterior portion of which is modified into an oval-shaped suction disc, while the posterior portion remains as a soft-rayed fin. The anal fin mirrors the shape and structure of the dorsal fin, with 20-26 soft rays. The modified dorsal fin serves as the primary mechanism for attachment to host organisms.^[28]^[29] Morphological variations occur among species, particularly in the structure of the suction disc; for instance, spinules on the disc are shorter in the genus Echeneis compared to those in the genus Remora. These differences influence attachment efficacy across host types.^[1]

Physiological adaptations

Remoras possess a specialized suction disc formed from a modified first dorsal fin, featuring 18 to 28 transverse laminae—movable, plate-like structures—that enable attachment to hosts by creating a vacuum seal.^[30] These laminae rotate into position against the host surface while an outer fleshy lip seals the edges, and muscle contractions further enhance adhesion by generating negative pressure differentials averaging -92.7 kPa (approximately 0.92 atm).^[30] This mechanism allows remoras to maintain attachment across diverse host textures and speeds without expending significant energy once secured.^[30] The respiratory system of remoras is adapted for attachment to mobile hosts, primarily relying on ram ventilation where water flows over the gills due to the host's movement, minimizing the need for active pumping.^[31] In low-flow conditions, such as when a host is stationary or in shadowed areas with reduced water movement, remoras switch to active branchial ventilation, increasing oxygen consumption by 3.7% to 5.7% to sustain respiration.^[31] Their circulatory system includes unique cranial vein modifications, with enlarged veins positioned ventral to the disc that help maintain the suction seal by managing fluid dynamics and preventing pressure equalization from seepage.^[32] Sensory adaptations in remoras include a well-developed lateral line system, which detects water vibrations and movements from potential hosts, facilitating attachment and orientation during symbiotic interactions.^[33] Additionally, the adhesive disc contains embedded mechanoreceptor complexes in its soft lip that respond to tactile stimuli, allowing remoras to sense surface contact and adjust attachment precisely.^[34] Due to their pelagic lifestyle, remoras demonstrate tolerance to significant pressure variations, with records of individuals accompanying hosts to depths exceeding 1000 meters without physiological distress.^[35] This adaptability, combined with their oceanic distribution, enables them to endure salinity fluctuations typical of marine environments, from coastal to open-water conditions.^[28]

Habitat and ecology

Distribution and habitat

Remoras, belonging to the family Echeneidae, exhibit a cosmopolitan distribution across tropical and subtropical waters of the Atlantic, Indian, and Pacific Oceans, spanning latitudes from approximately 40°N to 40°S.^[15] Some species, such as the whalesucker (Remora australis), extend into temperate regions, with records from as far north as Vancouver Island, Canada, and southern California, USA.^[36] These fish primarily inhabit pelagic environments in the open ocean, rarely venturing into nearshore or coastal areas except occasionally around coral reefs or in shallow inshore waters.^[37] They occupy the epipelagic zone, from the surface down to depths of about 100 m, where they are often observed free-swimming singly or in small schools.^[14] Although capable of independent swimming, remoras preferentially attach to mobile hosts using their dorsal-fin-derived suction disc, a strategy that conserves energy in this dynamic habitat.^[1] Remoras thrive in marine conditions with water temperatures typically ranging from 20–30 °C and salinities of 30–35 ppt, aligning with their preference for warm, open-water ecosystems.^[14] These tolerances support their widespread occurrence in stable, oligotrophic oceanic waters but limit them to regions with suitable thermal profiles.^[6]

Symbiotic relationships

Remoras form primarily commensal symbiotic relationships with a variety of marine megafauna, gaining transportation across vast ocean distances, protection from predators, and incidental access to food scraps or ectoparasites without inflicting significant harm on their hosts.^[38]^[39] This phoretic association allows remoras to conserve energy by hitchhiking on faster-swimming or migrating hosts, such as sharks and whales, which can reach speeds exceeding 5 m/s.^[40] Hosts may derive secondary benefits from remoras removing skin parasites like copepods, though the extent of this service varies and does not typically impose costs on the host.^[41] Host preferences among remora species show notable specificity, influencing attachment choices based on host mobility, size, and ecology. The common remora (Remora remora) preferentially attaches to pelagic sharks, billfishes from the family Istiophoridae, and swordfish (Xiphias gladius), often targeting larger, open-ocean predators.^[1] In contrast, the slender sharksucker (Echeneis naucrates) exhibits broader preferences, commonly associating with sea turtles, teleost fishes, elasmobranchs like rays, and marine mammals, including dolphins and whales.^[42] Other species, such as the white suckerfish (Remora albescens), display strong fidelity to manta rays and similar filter-feeding elasmobranchs, while the family as a whole opportunistically attaches to boats or debris when natural hosts are unavailable.^[43] These preferences are shaped by evolutionary adaptations, with phylogenomic analyses revealing axes of host breadth from specialist (e.g., shark-focused) to generalist strategies across the eight remora species.^[44] Attachment occurs at strategic sites on the host's body, such as the dorsal fin, gills, underbelly, or smooth skin surfaces, where hydrodynamic drag is minimized and access to flowing water for respiration is optimized.^[45] For instance, on blue whales, remoras selectively adhere to low-drag regions like the ventral grooves, reducing their own energetic costs.^[46] These attachments can persist for extended durations, often lasting weeks to months, with remoras detaching temporarily to feed or during reproduction before reattaching. The specialized suction disc enables this reversible hold, allowing remoras to remain secure amid host movements while detaching as needed without causing injury.^[47] While the relationship is predominantly commensal, emerging evidence points to potential mutualistic elements, particularly through ectoparasite removal that enhances host hygiene and reduces infection risks, as seen in associations with whale sharks involving three-way symbioses with copepods.^[48] Recent observations as of November 2025, including video footage of remoras on whales removing sea lice and a study on sailfish-remora cleaning interactions, further support mutualistic elements through parasite removal.^[49]^[50] Historical views classify it as mutualistic due to reciprocal benefits like parasite control for the host and transport efficiency for the remora, though interactions can occasionally border on parasitism if attachments cause minor drag or irritation.^[39] Such dynamics underscore the eco-evolutionary spectrum of remora-host interactions, balancing costs and gains across diverse partnerships.

Life history

Reproduction

Remoras are oviparous fishes that reproduce through external fertilization, releasing pelagic eggs into the open ocean. Spawning likely occurs seasonally in some species, such as peaking in summer for Echeneis naucrates in the Gulf of Mexico, though direct observations of the behavior are scarce, with only limited records from aquarium settings describing batch spawning events.^[52]^[53]^[54] Female remoras exhibit high fecundity; for example, in Echeneis naucrates, mean batch fecundity is estimated at 1,710,000 ± 600,000 eggs, varying with body size and species, based on relative fecundity rates of approximately 40 hydrated oocytes per gram of ovary-free body mass. Little is known about specific spawning behaviors beyond occasional pairing of mature individuals on hosts.^[54]^[55] The eggs hatch into planktonic larvae that initially lack a suction disc and drift in the pelagic zone. The suction disc develops from modified dorsal fin elements during larval ontogeny, becoming functional around metamorphosis at 2–3 cm in length, at which point the juveniles begin associating with host organisms following settlement.^[56]^[26] Sexual maturity is attained at lengths of ~20–30 cm in some species (e.g., Echeneis naucrates), typically within 2–5 years, with lifespans of 4–10 years depending on species. No parental care is provided to eggs or larvae, relying instead on the dispersive nature of the pelagic habitat for survival and distribution.^[9]^[55]

Diet and feeding

Remoras are primarily carnivorous, exhibiting an opportunistic diet that relies heavily on their association with host organisms such as sharks, rays, and cetaceans. The bulk of their nutrition comes from ectoparasites, particularly copepods, which they remove from the host's skin and gills, supplemented by food scraps from the host's meals, sloughing epidermal tissue, and occasionally host feces.^[57] Zooplankton and smaller nektonic prey, such as baitfish, also contribute to their diet, especially when remoras are not attached to a host.^[39] There is no evidence of herbivory in remora feeding habits, confirming their exclusively carnivorous nature.^[57] Feeding occurs through a combination of passive and active strategies. While attached to a host using their specialized suction disc, remoras position themselves near the host's mouth or body to passively capture drifting food particles or actively nibble at parasites with their small, tooth-lined mouths.^[58] They periodically detach to pursue plankton or small fish independently, using bursts of swimming to chase prey in the water column, which allows them to exploit a broader range of resources beyond host proximity.^[59] This symbiotic hitchhiking enhances their access to concentrated food sources, reducing the energy costs of independent foraging.^[57] In marine food webs, remoras function as secondary consumers with a mean trophic level of 3.5, reflecting their position as predators of primary consumers like parasites and plankton while depending on higher-trophic-level hosts for optimal feeding efficiency. Their diet composition varies by species and life stage, but the reliance on host-derived resources underscores their ecological niche as cleaners and scavengers in pelagic environments.^[57]

Relationship with humans

Use in fishing

In traditional fishing practices of the Pacific Islands, particularly in the Torres Strait region of Melanesia, remoras (known locally as "gep" in the Meriam language) were employed as living tools to capture large marine animals, including turtles and occasionally large fish, prior to the early 20th century.^[60]^[61] Similar methods have been documented worldwide, including in East Africa for catching fish and turtles, in Cuba and Colombia for turtle fishing, and archaeologically in Southeast Arabia linked to coastal subsistence.^[61] Fishermen would capture a live remora, attach a strong line or cord to its tail or body—often securing it with a loop to avoid damaging the fish—and release it into the water near potential targets. The remora's specialized dorsal fin, modified into a powerful suction disc, would naturally adhere to the host animal, such as a hawksbill turtle or a large pelagic fish like tuna, allowing the fisherman to haul in the catch once attached.^[62]^[63] This technique exploited the remora's physiological adaptation for attachment, documented in 19th-century ethnographies by anthropologists like Alfred Cort Haddon during expeditions to Mabuiag Island in the Torres Strait, where locals described using the remora to target turtles by paddling canoes quietly and waiting for the attachment before pulling the line.^[61]^[63] Although similar methods were reported in broader Pacific contexts, such as Melanesian islands, there is limited specific documentation for Hawaii or core Polynesian societies like Samoa, where other line-and-hook techniques dominated for pelagic species like marlin. The practice's mechanism relied on the remora's innate behavior to seek hosts, with fishermen observing the line's movement to detect successful attachments, as noted in historical accounts from the late 1800s emphasizing its effectiveness in calm, shallow waters near reefs.^[62]^[64] Today, remora-assisted fishing is rare and largely obsolete across the Pacific, supplanted by synthetic hooks, lures, and motorized boats in both commercial and artisanal fisheries, with no evidence of large-scale modern application. Occasional anecdotal reports persist in small-scale, traditional contexts in remote island communities, but it remains confined to cultural preservation rather than practical use. Ethnographic records tie remora behavior to Polynesian and Melanesian fishing innovations, where the fish's attachment ability inspired lore about resourceful sea companions, as reflected in Torres Strait Islander stories linking the "gep" to ancestral hunting ingenuity and symbiotic marine relations.^[60]^[61]

Mythology and culture

In ancient Roman accounts, the remora, referred to as the echeneïs, was mythologized as a diminutive fish possessing the extraordinary power to halt the largest ships by adhering to their hulls with its suction disc, symbolizing obstruction and delay. Pliny the Elder, in his Natural History (Book XXXII, Chapter 1), detailed this belief, claiming that a remora attached itself to the flagship of Mark Antony during the Battle of Actium in 31 BCE, thereby impeding Cleopatra's fleet and contributing to their defeat by Octavian's forces.^[65]^[66] This ancient notion evolved into widespread maritime folklore across Europe, where remoras earned the moniker "ship-stayers" and were feared by sailors as harbingers of stalled voyages or ill fortune at sea. In Asian traditions, particularly Japanese folklore, remoras manifest as ayakashi—supernatural sea spirits—that latch onto vessels to immobilize them, often interpreted as omens of impending peril or divine intervention in navigation. In modern cultural references, remoras feature in 19th-century adventure literature, such as Jules Verne's Twenty Thousand Leagues Under the Sea (1870), where they illustrate opportunistic sea life amid explorations, and serve as metaphors for dependency or parasitic attachments in broader literary discourse on symbiosis and hindrance. Unlike figures in many mythologies, remoras lack prominent religious symbolism, appearing instead as emblems of tenacity or encumbrance in secular narratives.

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Remora australis (Bennett 1839), the Whalesucker ... The breeding behavior and the behavior of larvae and juveniles of the sharksucker, Echeneis naucrates.
[52]
Growth and reproduction of Echeneis naucrates from the eastern ...
This study describes growth and reproductive characteristics of a facultative elasmobranch symbiont, the sharksucker (remora) Echeneis naucrates, from the ...
[53]
Remora osteochir, Marlin sucker - FishBase
Preferred temperature (Ref. 123201): 17.3 - 28.2, mean 25.9 °C (based on 1457 cells). Phylogenetic diversity index (Ref. 82804): PD50 ...Missing: salinity | Show results with:salinity
[54]
Animal Morphology Journal | Wiley Online Library
Jul 25, 2012 · We studied the development of the sucking disc based on a series of larval, juvenile, and adult echeneids ranging from 9.3 mm to 175 mm standard ...
[55]
(PDF) Remoras - ResearchGate
This chapter examines the marine fish remoras of the family Echeneidae. Remora, suckerfish, diskfish, and sucker are some of the names describing eight species ...Missing: circulatory oxygen
[56]
Sea Wonder: Remora | National Marine Sanctuary Foundation
Apr 26, 2022 · Remoras are thin, elongated, and rather dark fishes that are often black or brown in color. Their heads are long and flat, with the lower jaw ...Missing: biology | Show results with:biology
[57]
International study uncovers secret surfing life of remoras hitchhiking ...
Oct 28, 2020 · The study shows the secrets behind the remora fish's success in hitchhiking aboard baleen whales more than 30 times their size to safely ...Missing: diet | Show results with:diet
[58]
Gep (Remora – Sucker Fish) - Canopy Art
Gep is Meuram language name for the remora, commonly known as the suckerfish. Torres Strait Islanders once used gep in traditional hunting methods.
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(PDF) Evidence for Fishing with Remora across the World and ...
People intentionally caught remora for use as a fishing aid, allowing them to haul in turtles, marine mammals, and other species to which remora attach ...
[60]
[PDF] On the Use of the Sucking-Fish for Catching Fish and Turtles
Fishing with the Remora. After Holder, 1899. FIG. 11. An Echeneis, twenty ... In Torres Strait and the lortllerln coast the hawksbill turtle . . . is.
[61]
Once More the Sucking-Fish - jstor
traveled eastward, to the Torres Strait and Melanesia, ... to America. 5. There are in America no corroborative stories of the remora fishing, except as derived ...
[62]
Studies in Echeneis or Remora, II
Fishing with the Remora. After Holder, 1899. FIG. 11. An Echeneis, twenty ... In Torres Strait and the lortllerln coast the hawksbill turtle . . . is.
[63]
the natural history of pliny. - Project Gutenberg
Some of our own authors have given this fish the Latin name of “mora.”12 It is a singular thing, but among the Greeks we find writers who state that, worn as an ...
[64]
The naval battle of Actium and the myth of the ship-holder - arXiv
May 27, 2019 · A fish, called echeneis or remora, is said to have the power to stop ships or to delay their motion by adhering to the hull.
[65]
What Does Ayakashi Mean in English?
Aug 21, 2013 · According to folk belief, if remoras fasten themselves to the underside of your boat, you will become stuck in the water and unable to move. In ...
[66]
The horrible natural history of remora - DIVE Magazine
Nov 9, 2021 · Remora, in Latin means delay, a reference to their supposed ability to slow down ships. The generic name Echeneis is derived from the Greek ...Missing: etymology | Show results with:etymology