Notes, observations and questions concerning Mediterranean moray eel, Muraena helena Linnaeus, 1758
A little over ten years ago, in September of 2014, I was snorkelling in the shallows off Salina when I saw my first Mediterranean moray eel (Figure 1).
Fig. 1: Mediterranean moray eel, Muraena helena (16th September 2014).
With its gaping mouth and defiant, languid posturing the signalling from one animal to another held an unclouded depth of clarity, that didn’t jump the species barrier so much as completely ignore it, lighting-up my amygdala and manifesting this thought within the primal depths of my brain, “Don’t go any closer”. I went closer. How can you not?
But equally, “how close is too close?”, is not a question I want an answer too. So, while awe and beauty pulled me in, they were tethered by caution (top-marks to the moray eel for that). Did I invade another creature’s space for a better shot. I did. Was my ‘rudeness’ limited? It was. Also, as I’m sure is true for many biologists, I am prone to navigate such intrusions to reduce disturbance and mitigate any likely distress. Afterall, surely, we lose nothing and probably see more, with a ‘light tread’? In this instance, I am happy to say that the meeting was navigated without physical injury. The memory of it still hits the glow-switch inside my head.
Here is a top predator with charisma and ecological importance that arguably, is understudied, misunderstood and largely ignored. Why? How can this big fish have been forgotten, dismissed?
Classification
The Family to which this species belongs is the Muraenidae. Above ‘Family’ we have the rank of ‘Order’ and below there is ‘Genus’, which in this case is ‘Muraena’, followed by ‘species’, ‘helena’.
According to FishBase the first fossil record of species in the Family Muraenidae was from the upper Tertiary Miocene (Berg, L.S. 1958). The Miocene lasted from 23.8–5.3 million years ago (Mya) and was the epoch within which Hominidae, or great apes also appeared. However, Reece and Mehta (2013) reference Benton (1993) and postulate that, "Moray eels probably appear in the fossil record between 34 and 54 Mya". That said, they also state that, "The oldest full-bodied fossil of a moray is dated to 5.3 Mya".
If Muraenidae did evolve in the Miocene this could suggest that the Family may prosper going forward. The Miocene was a period of warmer global temperatures than the preceding Oligocene epoch and the subsequent Pliocene epoch, with a climate and, atmospheric CO2 levels, similar to those projected for the future. Unsurprisingly, by studying and augmenting our understanding of the Miocene, scientists hope to understand more about the effect of climate change on living organisms.
Catch and consumption
So, how is the Mediterranean moray eel doing today? A 2011 paper stated that, in the decade up to 2009 it had, “become very frequent” with a “pronounced dominance” in, “longline catches of artisanal fishers” from the “outer islands in the middle and southern Adriatic” (Matić-Skoko et al. 2011).
Whether the same is true more broadly across the Mediterranean is not entirely clear but it has been noted that in the Ionian sea off Catania, Sicily, “during the last 5-10 years the abundance of M. helena seems increased” (Tiralongo et al. 2022). In correspondence with Simone Follone, a fisherman from Salina, I understood two things. Firstly, that Eolian Island people have always eaten Mediterranean moray eel where, for example, thinly sliced, unskinned fillets, might be fried or broiled with chestnuts; and, historically, local restaurants have not served moray eel, it being seldom asked for by their patrons. This, as touched on below, is in marked contrast to the high status and value of the species in the Roman era. Now, however, a Green Starred Michelin restaurant run by chef Martina Caruso, on Salina, is creating dishes using this locally caught fish (this Michelin award is given in recognition of excellence with respect to all aspects of environmental sustainability and ethical practice).
In terms of adult fish Simone Follone highlighted that the waters around the neighbouring island Filicudi, to the west, is full of them. An observation which itself sparks many questions! Why might moray eel catches be better around Filicudi? What supports such abundance there in particular?
It's worth mentioning Francis Adams here, a Scot from Banchory, a small town in Aberdeenshire. He was the son of a village labourer and according to Prof. Charles Singer, besides being an excellent and dedicated doctor, was “one of the greatest medical scholars of his age” (Nature, 1942). This side-hustle included his translation of ‘The seven books of Paulus Ægineta’ [more on this key text later] from which we learn the following:
“We need not say how much the murene was sought after by all the lovers of good eating in ancient Rome. Pliny, Martial, and Macrobius inform us that those from Sicily were in most esteem [especially if eaten before they had spawned]; and Brydone [a Scottish traveller and author] takes notice of the peculiar excellence of the Sicilian murenes at the time when he performed his tour through that island”.
This is reinforced by what is considered Europe’s oldest cookery book, the ‘De re coquinaria’ or, ‘The Art of Cookery’ (Vehling 1977):
“The ancients considered the murena one of the finest of fish; the best were brought from the straits of Sicily. Rich Romans kept them alive in their fish ponds, often large and elaborate marble basins called, piscina, fattened the fish, kept it ready for use”.
This collection of recipes, attributed to Apicius, includes a number dedicated to ‘Murena’; three for broiled fish and three for boiled fish. Although it has been argued that recipes in the ‘De re coquinaria’ for fish are more about the ingredient-rich sauces than the fish, with Vehling’s translation noting:
“Fish sauces tended to be particularly elaborate: boiled murena (likely eel), for example, called for pepper, lovage, dill, celery seed, coriander, dried mint, and rue, as well as pine nuts, honey, vinegar, wine, and oil (recipe 451)”.
At around the same time a Sicilian Greek called Archestratus, wrote what is described as a “mock epic poem on gastronomy” including this advice for the preparation of tender fish:
“sprinkle with a little salt only, and baste with oil. For it contains within itself alone the reward of joy” (Athenaeus, VII.321d).
This emphasis on simplicity is much more in tune with how people in the Eolian Islands and Sicily in general, cook today. Lastly, it is perhaps worth noting that Vehling’s translation notes the following:
“This part of Italy, and especially Sicily, because in close contact with Greece was for many years much farther advanced in art of cookery than the North”.
In the Eolian Islands, in more recent times, moray eels have not been a directly targeted species. Instead, they were landed as by-catch. Thus, if you set gill nets and catch smaller fish in this way, Moray eel may be coaxed from their hides to take advantage of this banquet and hence become entangled as well. Fixed gear in the form of bottom-set longlines and traps are two methods used by artisanal fisherman in the Mediterranean to catch M. helena (Matić-Skoko et al. 2011). Data from Global Fishing Watch (2024) indicates fixed gear was deployed off the south-east coast of Stromboli in depths of 200 - 500 metres in particular (but what species might have been landed from this activity was not seen).
Reflections
What can be seen in the image presented in Figure 2?
But equally, “how close is too close?”, is not a question I want an answer too. So, while awe and beauty pulled me in, they were tethered by caution (top-marks to the moray eel for that). Did I invade another creature’s space for a better shot. I did. Was my ‘rudeness’ limited? It was. Also, as I’m sure is true for many biologists, I am prone to navigate such intrusions to reduce disturbance and mitigate any likely distress. Afterall, surely, we lose nothing and probably see more, with a ‘light tread’? In this instance, I am happy to say that the meeting was navigated without physical injury. The memory of it still hits the glow-switch inside my head.
Here is a top predator with charisma and ecological importance that arguably, is understudied, misunderstood and largely ignored. Why? How can this big fish have been forgotten, dismissed?
Classification
The Family to which this species belongs is the Muraenidae. Above ‘Family’ we have the rank of ‘Order’ and below there is ‘Genus’, which in this case is ‘Muraena’, followed by ‘species’, ‘helena’.
According to FishBase the first fossil record of species in the Family Muraenidae was from the upper Tertiary Miocene (Berg, L.S. 1958). The Miocene lasted from 23.8–5.3 million years ago (Mya) and was the epoch within which Hominidae, or great apes also appeared. However, Reece and Mehta (2013) reference Benton (1993) and postulate that, "Moray eels probably appear in the fossil record between 34 and 54 Mya". That said, they also state that, "The oldest full-bodied fossil of a moray is dated to 5.3 Mya".
If Muraenidae did evolve in the Miocene this could suggest that the Family may prosper going forward. The Miocene was a period of warmer global temperatures than the preceding Oligocene epoch and the subsequent Pliocene epoch, with a climate and, atmospheric CO2 levels, similar to those projected for the future. Unsurprisingly, by studying and augmenting our understanding of the Miocene, scientists hope to understand more about the effect of climate change on living organisms.
Catch and consumption
So, how is the Mediterranean moray eel doing today? A 2011 paper stated that, in the decade up to 2009 it had, “become very frequent” with a “pronounced dominance” in, “longline catches of artisanal fishers” from the “outer islands in the middle and southern Adriatic” (Matić-Skoko et al. 2011).
Whether the same is true more broadly across the Mediterranean is not entirely clear but it has been noted that in the Ionian sea off Catania, Sicily, “during the last 5-10 years the abundance of M. helena seems increased” (Tiralongo et al. 2022). In correspondence with Simone Follone, a fisherman from Salina, I understood two things. Firstly, that Eolian Island people have always eaten Mediterranean moray eel where, for example, thinly sliced, unskinned fillets, might be fried or broiled with chestnuts; and, historically, local restaurants have not served moray eel, it being seldom asked for by their patrons. This, as touched on below, is in marked contrast to the high status and value of the species in the Roman era. Now, however, a Green Starred Michelin restaurant run by chef Martina Caruso, on Salina, is creating dishes using this locally caught fish (this Michelin award is given in recognition of excellence with respect to all aspects of environmental sustainability and ethical practice).
In terms of adult fish Simone Follone highlighted that the waters around the neighbouring island Filicudi, to the west, is full of them. An observation which itself sparks many questions! Why might moray eel catches be better around Filicudi? What supports such abundance there in particular?
It's worth mentioning Francis Adams here, a Scot from Banchory, a small town in Aberdeenshire. He was the son of a village labourer and according to Prof. Charles Singer, besides being an excellent and dedicated doctor, was “one of the greatest medical scholars of his age” (Nature, 1942). This side-hustle included his translation of ‘The seven books of Paulus Ægineta’ [more on this key text later] from which we learn the following:
“We need not say how much the murene was sought after by all the lovers of good eating in ancient Rome. Pliny, Martial, and Macrobius inform us that those from Sicily were in most esteem [especially if eaten before they had spawned]; and Brydone [a Scottish traveller and author] takes notice of the peculiar excellence of the Sicilian murenes at the time when he performed his tour through that island”.
This is reinforced by what is considered Europe’s oldest cookery book, the ‘De re coquinaria’ or, ‘The Art of Cookery’ (Vehling 1977):
“The ancients considered the murena one of the finest of fish; the best were brought from the straits of Sicily. Rich Romans kept them alive in their fish ponds, often large and elaborate marble basins called, piscina, fattened the fish, kept it ready for use”.
This collection of recipes, attributed to Apicius, includes a number dedicated to ‘Murena’; three for broiled fish and three for boiled fish. Although it has been argued that recipes in the ‘De re coquinaria’ for fish are more about the ingredient-rich sauces than the fish, with Vehling’s translation noting:
“Fish sauces tended to be particularly elaborate: boiled murena (likely eel), for example, called for pepper, lovage, dill, celery seed, coriander, dried mint, and rue, as well as pine nuts, honey, vinegar, wine, and oil (recipe 451)”.
At around the same time a Sicilian Greek called Archestratus, wrote what is described as a “mock epic poem on gastronomy” including this advice for the preparation of tender fish:
“sprinkle with a little salt only, and baste with oil. For it contains within itself alone the reward of joy” (Athenaeus, VII.321d).
This emphasis on simplicity is much more in tune with how people in the Eolian Islands and Sicily in general, cook today. Lastly, it is perhaps worth noting that Vehling’s translation notes the following:
“This part of Italy, and especially Sicily, because in close contact with Greece was for many years much farther advanced in art of cookery than the North”.
In the Eolian Islands, in more recent times, moray eels have not been a directly targeted species. Instead, they were landed as by-catch. Thus, if you set gill nets and catch smaller fish in this way, Moray eel may be coaxed from their hides to take advantage of this banquet and hence become entangled as well. Fixed gear in the form of bottom-set longlines and traps are two methods used by artisanal fisherman in the Mediterranean to catch M. helena (Matić-Skoko et al. 2011). Data from Global Fishing Watch (2024) indicates fixed gear was deployed off the south-east coast of Stromboli in depths of 200 - 500 metres in particular (but what species might have been landed from this activity was not seen).
Reflections
What can be seen in the image presented in Figure 2?
Fig. 2: Mediterranean moray eel, Muraena helena, in cobble habitat (17th June 2018).
Caution and curiosity? Fight or flight? The eels may not be known for their swimming skill but this one did not flee, perhaps understanding that the large creature lumbering above it was unlikely, or unable, to strike with any rapidity. It edges around serpentine elegance as it hugs the seabed, engaging the eye but without, I thought, the hypnotic tug inherent in its terrestrial equivalents. Instead, this broadside view emphasized its hefty flank, messaging bulk and strength. Another warning? If so, it back-fired for the moray, drawing me in, as it presented an unimpeded view with its skin, a living, smooth, aquatic parchment spotted and patterned with beguiling script-like scrawls. I was mesmerised, engaged, captured by this ancient weave, this dance on the cobble-carpet, this arcane visual spell-work, threading and stitching its way into my inner story.
Compared to my first encounter this was a much smaller eel. So, while the first may have been an adult, this one, with more certainty, was likely to be a juvenile. The remnant mermaid's cup, Acetabularia acetabulum, growths visible in the photograph give some indication of scale. Those shallow cups, or pale green parasols, at the tip of the short stems reach a maximum of just 1.2cm in diameter.
Labels commonly applied to the species include slimy, aggressive, predatory, cryptic, camouflaged. Inevitably, these labels carry some hefty, human baggage which, consciously or unconsciously, will colour our perception of the things to which they are applied. It can be argued that words such as these are reflective of what has been brought to the species, rather than what they say about the species. What adjectives would you use to describe Mediterranean moray eel?
The original description
Carl Linnaeus, who first described the species in his 10th Edition of ‘Systema Naturæ’, published in 1758, provided the following (Figure 3):
Compared to my first encounter this was a much smaller eel. So, while the first may have been an adult, this one, with more certainty, was likely to be a juvenile. The remnant mermaid's cup, Acetabularia acetabulum, growths visible in the photograph give some indication of scale. Those shallow cups, or pale green parasols, at the tip of the short stems reach a maximum of just 1.2cm in diameter.
Labels commonly applied to the species include slimy, aggressive, predatory, cryptic, camouflaged. Inevitably, these labels carry some hefty, human baggage which, consciously or unconsciously, will colour our perception of the things to which they are applied. It can be argued that words such as these are reflective of what has been brought to the species, rather than what they say about the species. What adjectives would you use to describe Mediterranean moray eel?
The original description
Carl Linnaeus, who first described the species in his 10th Edition of ‘Systema Naturæ’, published in 1758, provided the following (Figure 3):
Fig. 3: The original description from Linnaeus of Muræna helena in ‘Systema Naturæ’, 1758.
If your Latin, like mine, is a little rusty this translation may help (Table 1). The first is easy enough, ‘MURÆNA’ is the Genus, Muraena. Note ‘Æ’ or ‘æ’ is simply a ligature where two letters have been joined to form a single character and syllable, representing the Latin diphthong, or two sounds, in this case, ae so, your tongue moves to create the ‘gliding vowel’ sound. Here’s the rest (be aware that (Table 1, [1]) outlines features applicable to the Genus):
Table 1. A translation in [eight] sections of the original description for the species, Muraena helena by Linnaeus 1758.
Table 1. A translation in [eight] sections of the original description for the species, Muraena helena by Linnaeus 1758.
| 1 |
MURÆNA Caput læve. Nares tubulofæ. Membrana branchiostega radiis 10, ejusque Apertura ad pinnas pectorales. Corpus teretiufculum, lubricum. Pinna caudæ coadunata dorsali anique. Small head. Tubular nostrils. Branchial [gill] membrane with 10 rays, and its opening near the pectoral fins. The body is smooth, slimy. The tail fin is joined to the dorsal fin and the anal fin. |
| 2 |
Helena, 1. M. pinnis pectoralibus nullis. Muraena helena has no pectoral fins. [The ‘1’ is there because in this section Linnaeus describes 7 other species that he places within the Genus ‘Muraena’] |
| 3 |
Muræna pinnis pectoralibus carens. [This is prefaced with the source reference, ‘Art. gen. 25. syn. 41. Gron, muf I. n. 16. Muf. Ad. Fr. I. p. 319’ (this work is currently unknown to me)] A moray eel lacking pectoral fins. [Presumably, the cited reference states this and / or provides an illustration] |
| 4 |
Muræna maculata nigra, & viridis. [This is also prefaced with the source references, namely, ‘Seb. mus. 2. t. 69. f. 4, 5.’ (currently unknown to me); and, ‘Catesb. car. 2. t. 20, 21.’ (‘Catesb. car.’ is shorthand for Mark Catesby’s work titled, ‘The Natural History of Carolina, Florida and the Bahama Islands’. Volume 2 was published in 1743)] The moray eel spotted black, and green. |
| 5 |
Habitat in Europæ, Americæ Oceano. It lives in the ocean of Europe and America. [This appears to refer solely to the Atlantic]. |
| 6 |
Cænarum Helena, Rome in piscinis: natat ad magistrum delicata Muræna, Martial; The tender Moray eel swims towards the keeper, in tanks in Rome [raised for food consumption]; [Source, ‘Martial’, the Roman Poet, Marcus Valerius Martialis, who in Epigram X.30 wrote that at feeding time, “the tender moray eel swims towards its keeper” (“la delicata murena nuota al cenno dell'istruttore”)] |
| 7 |
Ved. Pollio damnata mancipia immergebat vivariis Murænarum, ut in visceribus suis aliquid de Servorum suorum corpore gustaret, Plin. IX: 23. Vedius Pollio had condemned slaves thrown into moray eel ponds so that he could taste something of the slaves' bodies in its viscera. [This is taken from ‘Plin. IX: 23’ or, more fully, Book 9, Chapter 23 / 24 of ‘The Natural History’, by Pliny the Elder (available in the Perseus Digital Library). Note that Pérez-Lloréns et al. (2021) indicate that it was a “common belief that human flesh greatly improved the fish quality” and that Pollio particularly, “enjoyed the liver of the fish that previously had eaten some body parts of his victim”] |
| 8 |
Morsu infesta. Its bite is dangerous. [Note that ‘Infesta’ on its own translates as ‘aggressive’, but together with ‘morsu’ it may translate more accurately as, ‘Fierce with its bite’, as used in Milton’s Comus, “Quos homines putas lupi sunt, &e Daunia, Getuliave monstra' morsu infesta” (Singleton, 1943)] |
That final note of caution from Linnaeus (Table 1, [8]) reminds me of the letter from Sirius Black to Harry Potter in the Goblet of Fire, “P.S. The bird bites”. It’s a safety announcement. So, you might interpret that Linnaeus is effectively signing off his description with, “Cave, hic piscis morsus”, be careful, this fish bites. In Harry’s situation Sirius’ post-script is a useful piece of information, delivered somewhat tardily for comic effect, because as we see, this owl quite simply, does bite. In terms of the Mediterranean moray eel there are, unfortunately, plenty of pages on various websites noting the bite, whilst neglecting the broader truth. Behavioural responses of the type ‘bite-first-and-ask-questions-later’ are not the norm in the animal kingdom. Strategies have, for example, an energetic cost and hence, do not come out of nowhere. There is always a ‘back-story’, both immediate and evolutionary. It could be argued, ‘morsu infesta’ does not deny this, especially if you make the link to Martialis (Table 1, [6]). Nevertheless, if we combine the ‘bond-villain’ horror of Vedius Pollio (Table 1, [7]), throwing condemned slaves to the jaws of this predator, with its fierce bite, the text, could be interpreted as perpetuating a dark mythology that the eel itself is inherently dangerous and prone to attack with aggression and fierceness. In any case, for Linnaeus, this information was characteristic, to the extent that he enshrined it in the description. Afterall, you would not want to be bitten by a moray eel as it would be dangerous. But in his day, who would commonly encounter live specimens and under what circumstances? I’ll take a wild guess that Roman fish-farming of the species is long gone, and that fisherman understand well-enough what to expect from a freshly landed fish. That said, I suspect ‘modern’ readers might benefit from greater clarity (I would be first in-line!). For example, “Hunc piscem mordere potest, si periculum senserit”, this fish can bite, if it feels threatened.
Why choose Pollio? It wasn’t as if Linnaeus would have been that short of alternative historical references (and he was known to be an avid reader). Instead of outlining Pollio’s predelictions he might have offered us something less sensationalist and more in keeping with the high regard wealthy Romans had for Mediterranean moray eels. For example, in Volume III, page 15 of the ‘Seven Books of the Saturnalia‘ by Macrobius (1400), we learn that the Roman orator Lucius Licinius Crassus wore black for the death of a moray eel that had been raised in the pool at his house, as if it were his daughter that had died (Macrobius, 1400; Adams, 1844; Pérez-Lloréns et al. 2021). Still interpretable as a bit mad, just not bloody. In the ‘Cookery and Dining in Imperial Rome’ Vehling notes the following:
“Pollio fattened murenas on human flesh, killing a slave on the slightest provocation and throwing the body into the fish pond; he would eat only the liver of such murenas. This is the only case of such cruelty on record, and it has often been cited and exaggerated”.
Still, perhaps I’m being a bit picky. Perhaps not. As Böhlke et al. (1989) note, “Large morays are much feared by divers and fishermen because of their malevolent appearance and occasional habit of provoked or unprovoked attacks upon hands thrust in their proximity. This is more a reflection of muraenid myopia than temperament (Randall, 1969)”. That last sentence is perfect and shows that concern with regard to how morays in general are portrayed or understood, is nothing new. It’s a shame then that Böhlke et al. chose the adjective “malevolent” to describe their appearance, given that the dictionary definition of that word, based on the Latin root is, “having or showing a wish to do evil to others”. What if they had said ‘scary appearance’ instead, wouldn’t that have captured a little more biological truth for both sides (with less hyperbole)? What sharp teeth you have Mr. Moray-Eel / All the better to bite you with!
What else do you notice as we work back through the original description provided by Linnaeus? Included is an indication of where each animal lives (Table 1, [5]). This is valuable information, if somewhat ambiguous, given, in part, that at the time there were only four known continents of the world: Europe, America, Asia and Africa. So, for M. helena we have, “Habitat in Europæ, Americæ Oceano”, with no direct mention of the Mediterranean. Odd, because in other species descriptions, for example Muraena myrus Linnaeus gives us, “Habitat in Mari Mediterraneo”; Blennius pholis – “Habitat in Oceano & Mari Mediterraneo”; Cottus scorpius – “Habitat in Oceano Europaeo” (for a species not recorded from the Mediterranean); Cottus cataphractus – “Habitat in M. Europaeo” (for a species not recorded from the Mediterranean); and, Pleuronectes solea – “Habitat in M. Europaeo” (for a species recorded from both the Atlantic and Mediterranean). This seems inconsistent from our perspective. But surely, we can put that to one side, afterall, the geographical references would have been of necessity, very ‘broadbrush’. It’s the binomial naming system, which is the key, if occasionally rather opaque, gift (who knew that Homo sapiens meant “Wise man”?!).
It seems reasonable to infer that Linnaeus would have been aware that the distribution of M. helena included the Mediterranean and therefore, ‘Habitat in Europæ, Americæ Oceano’ must, axiomatically, encompass it (unless it is simply an error and was meant to be explicitly recorded but was accidentally left out?). As a last thought, and leaving aside other potential inferences, it’s interesting to note that the Type Locality for Orcinus orca (Linnaeus, 1758) is detailed on the Global Biodiversity Information Facility (GBIF) as, ‘E North Atlantic ("Oceano Europaeo")’.
The 1758 description from Linnaeus also makes reference to colour, namely in the phrase, “Muræna maculata nigra, & viridis” (Table 1, [4]). This is lifted directly from ‘Catesb. car.’ Or Mark Catesby’s 1743 work, ‘The Natural History of Carolina, Florida and the Bahama Islands’ (Figure 4). But the citation particularly references ‘t 20, 21’ (Figure 5). These illustrations show the two forms of what Catesby calls a spotted moray (Muræna maculata), the Green (viridis) and the Black (nigra) (Figure 4; Figure 5). Notice too that Catesby also mentions, as evidenced by people in the Bahama’s, that this fish can bite (Figure 4).
Fig. 4: ‘Muræna maculata nigra, & viridis’ (Catesby, 1743).
Fig. 5: Catesby’s Green Moray T20 (above) and Black Moray T21 (below)
(taken from the digital collections of the University of South Carolina).
Why does Linnaeus direct us to Catesby’s ‘Muræna maculata’, given that it is clearly not M. helena (the colour and distribution being, for example, incorrect)? It must simply be to illustrate the type of fish he is describing, its more general features in comparison to other, similar spotted morays, already to be seen in the literature available at the time? I’m speculating because I don’t really know, but it seems reasonable. Also, there is no ‘M. maculata Catesby, 1743’ as the work is not describing species but detailing Natural History. Inevitably, there is a fish with the name Muraena maculata Lacepède, 1800 (the currently accepted name being, Haliophis guttatus (Forsskål, 1775)). However, this fish is not to be confused with Catesby’s as, among other things, it is only found off East Africa. See also Lacépède, & Buffon (1798) which refers to ‘la murene tachetée’ / ‘la murene ponctuée’ i.e. the spotted moray / the dotted moray, as well as Muræna guttata where ‘guttata’ comes from the Latin word ‘guttātus’, meaning "spotted" or "speckled"!
What about Catesby’s fish you ask? I mean, can we be sure there’s no more specific connection intended or implied? I think that we can be fairly confident that the species illustrated in Mark Catesby’s work and detailed by him is, Gymnothorax moringa (Cuvier, 1829). As you may well have guessed, its common name is, the spotted moray! Although it turns out, in this instance, to be unrelated to ‘moringa’, which derives from "moringue", a local name in the Caribbean for small moray eels (presumably with just a hint of irony, given the species can reach up to 3m in length!). Naturally, there is an unaccepted synonym for G. moringa, Muraena punctata Gronow, 1854 where ‘punctata’ does indeed mean, ‘spotted’ (but, not Muraena punctata Rafinesque, 1810, which was actually Muraena helena Linnaeus, 1758)! The connection to Catesby is clear both from Cuvier (1829) and from information provided for G. moringa on Fishbase which indicates, via Eschmeyer’s Catalog of Fishes, the following, “moringa, Muraena Cuvier [G.] (ex Catesby)”, with ‘ex’ meaning ‘from’.
Note too that on Fishbase, G. moringa is described as being, “unusually aggressive towards man”; and, that the distribution includes the “Western Atlantic: North Carolina, USA and Bermuda…”. All of which is perfect for Catesby.
So, with respect to colour, the original description provides no detail. Fear not because for that we have this, reported in Pliny the Elder’s ‘The Natural History’ as coming from the poem ‘Halieutica’ (concerning the natural history of fish), which was attributed by him to the Roman poet Ovid, "the muræna burning with its spots of gold". Whether you believe the Halieutica was written by Ovid or not, there is some debate on this point apparently, it's a nice line, capturing in eight words several key characteristics of the species and searing them into your brain. We know Linnaeus was familiar with this work from Pliny as it is the same source from which he took the anecdote about Vedius Pollio and yet he makes no reference to it. Curious.
The origin of the species name, ‘helena’
We turn again now to Francis Adams and his translation of the seven books of Paulus Ægineta (1844). Paulus Ægineta, or Paul of Aegina was a 7th-century Greek physician who compiled a medical encyclopedia commonly referred to as the ‘Medical Compendium in Seven Books’. Linnaeus was well acquainted with this encyclopedia. In ‘Species Plantarum’, ‘The Species of Plants’, published in 1753 he created the Genus ‘Aeginetia’ in honour of Paul of Aegina.
In his commentary on Section XC (90), ‘On Fishes’ of the seven books, Adams says this:
“According to Icesius, it is as nutritious as the eel. (Athen. vii.) He mentions that the murene was called “the Helen of suppers”.
From which Adams then goes on to postulate, “and hence, no doubt, Linnæus has named the fish muræna Helena”.
We need to unpick this a little as it turns out this is not the full quote. So, here we go. Adams cites 'Athen. vii.'. This is shorthand for, Athenaeus, Book 7. The book in question is The Deipnosophists (or, ‘Banquet of the learned of Athenaeus’) written in the 3rd Century AD. Athenaeus is the author. The term ‘deipnosophist’ can be translated as meaning, “A master of the art of dining, particularly learned conversation in the manner of Athenaeus's characters” (Wiktionary, 2024)). The full quote, from a translation by Yonge (1854) is:
“But a certain Epicurean, who was one of our party, when an eel was served up, said, -
Here is the Helen of the feast; I therefore will be the Paris!”
This confirms that, within the context of the quote, the reference is to Helen of Troy, who was taken by Paris, an action that precipitated the Trojan War. In this case if you read on you see that the unnamed Epicurean devours the eel. An action which, according to Jacob (2013), wins him the prize for gluttony. If Adams is correct in his assertion that this reference is the source of the species name, ‘helena’, selected by Linnaeus, then perhaps we are all Paris, as we gaze upon the Helen of the eels. Not only that but its application turns out to be a banner, trying to tell us something we've forgotten, that murene are good to eat.
Shallow water cobble habitat in the Eolian Islands
If you’re not looking, you might think the stony seabed just north of Santa Marina’s busy little ferry port a barren place, harbouring little but rocks and the occasional crab, echinoderm, bearded fireworm or snail. There are no showy corals advertising their presence with bold forms and bright colours. It would be easy to miss the life that is here because mostly, it doesn’t pay to be seen, particularly if you are an ambush predator like the Mediterranean moray eel (Figure 6).
What about Catesby’s fish you ask? I mean, can we be sure there’s no more specific connection intended or implied? I think that we can be fairly confident that the species illustrated in Mark Catesby’s work and detailed by him is, Gymnothorax moringa (Cuvier, 1829). As you may well have guessed, its common name is, the spotted moray! Although it turns out, in this instance, to be unrelated to ‘moringa’, which derives from "moringue", a local name in the Caribbean for small moray eels (presumably with just a hint of irony, given the species can reach up to 3m in length!). Naturally, there is an unaccepted synonym for G. moringa, Muraena punctata Gronow, 1854 where ‘punctata’ does indeed mean, ‘spotted’ (but, not Muraena punctata Rafinesque, 1810, which was actually Muraena helena Linnaeus, 1758)! The connection to Catesby is clear both from Cuvier (1829) and from information provided for G. moringa on Fishbase which indicates, via Eschmeyer’s Catalog of Fishes, the following, “moringa, Muraena Cuvier [G.] (ex Catesby)”, with ‘ex’ meaning ‘from’.
Note too that on Fishbase, G. moringa is described as being, “unusually aggressive towards man”; and, that the distribution includes the “Western Atlantic: North Carolina, USA and Bermuda…”. All of which is perfect for Catesby.
So, with respect to colour, the original description provides no detail. Fear not because for that we have this, reported in Pliny the Elder’s ‘The Natural History’ as coming from the poem ‘Halieutica’ (concerning the natural history of fish), which was attributed by him to the Roman poet Ovid, "the muræna burning with its spots of gold". Whether you believe the Halieutica was written by Ovid or not, there is some debate on this point apparently, it's a nice line, capturing in eight words several key characteristics of the species and searing them into your brain. We know Linnaeus was familiar with this work from Pliny as it is the same source from which he took the anecdote about Vedius Pollio and yet he makes no reference to it. Curious.
The origin of the species name, ‘helena’
We turn again now to Francis Adams and his translation of the seven books of Paulus Ægineta (1844). Paulus Ægineta, or Paul of Aegina was a 7th-century Greek physician who compiled a medical encyclopedia commonly referred to as the ‘Medical Compendium in Seven Books’. Linnaeus was well acquainted with this encyclopedia. In ‘Species Plantarum’, ‘The Species of Plants’, published in 1753 he created the Genus ‘Aeginetia’ in honour of Paul of Aegina.
In his commentary on Section XC (90), ‘On Fishes’ of the seven books, Adams says this:
“According to Icesius, it is as nutritious as the eel. (Athen. vii.) He mentions that the murene was called “the Helen of suppers”.
From which Adams then goes on to postulate, “and hence, no doubt, Linnæus has named the fish muræna Helena”.
We need to unpick this a little as it turns out this is not the full quote. So, here we go. Adams cites 'Athen. vii.'. This is shorthand for, Athenaeus, Book 7. The book in question is The Deipnosophists (or, ‘Banquet of the learned of Athenaeus’) written in the 3rd Century AD. Athenaeus is the author. The term ‘deipnosophist’ can be translated as meaning, “A master of the art of dining, particularly learned conversation in the manner of Athenaeus's characters” (Wiktionary, 2024)). The full quote, from a translation by Yonge (1854) is:
“But a certain Epicurean, who was one of our party, when an eel was served up, said, -
Here is the Helen of the feast; I therefore will be the Paris!”
This confirms that, within the context of the quote, the reference is to Helen of Troy, who was taken by Paris, an action that precipitated the Trojan War. In this case if you read on you see that the unnamed Epicurean devours the eel. An action which, according to Jacob (2013), wins him the prize for gluttony. If Adams is correct in his assertion that this reference is the source of the species name, ‘helena’, selected by Linnaeus, then perhaps we are all Paris, as we gaze upon the Helen of the eels. Not only that but its application turns out to be a banner, trying to tell us something we've forgotten, that murene are good to eat.
Shallow water cobble habitat in the Eolian Islands
If you’re not looking, you might think the stony seabed just north of Santa Marina’s busy little ferry port a barren place, harbouring little but rocks and the occasional crab, echinoderm, bearded fireworm or snail. There are no showy corals advertising their presence with bold forms and bright colours. It would be easy to miss the life that is here because mostly, it doesn’t pay to be seen, particularly if you are an ambush predator like the Mediterranean moray eel (Figure 6).
Fig. 6: Cobble habitat and juvenile Mediterranean moray eels, Salina, Le Isole Eolie.
What you learn as you snorkel between the baking sun above and the salted sea below, with the slow-soundscape crackling all around, is patience, and the treasure-box of rewards it gifts. Because, observing juvenile M. helena in the shallow water cobble habitat off Salina, whilst not easy, is inevitable. From depths of less than a metre down to about 5m, encountering these young fish is just a matter of time, particularly if you’re taking one last quick snorkel before the sun sets. They are there, and it occurs to me that if I’m seeing one or two in a relatively small area, how many remain hidden, how many do I miss while finning over this mosaic in failing light? A study from the Adriatic suggests that immature individuals, ranging “from 31.5 to 78 cm, with an average TL [total length] of 62.55 cm”, in conjunction with males, dominate in coastal shallow waters (Matić-Skoko et al. 2011). Results from Matić-Skoko et al. (2011) also show that immature individuals were distributed across all three sampling depths in that study (5-15m, 15-30m and 30-80m). The limited observations presented here, though only providing anecdotal evidence are, in terms of the juveniles, consistent with this data. These observations also suggest that the depth range of 0-5m is utilized by the species. If nothing else, I would argue that this is a good reminder of the value of cobble habitat.
This value is already well understood of course because the location where these images were taken, is within the Natura 2000 site, ‘Fondali dell'Arcipelago delle Isole Eolie’ (Figure 7).
This value is already well understood of course because the location where these images were taken, is within the Natura 2000 site, ‘Fondali dell'Arcipelago delle Isole Eolie’ (Figure 7).
Fig. 7: The Special Area of Conservation Fondali dell'Arcipelago delle Isole Eolie, designated under the European Habitats Directive.
As the Standard Data Form (SDF) for the site indicates, those Annex I habitat types presence in the Fondali dell'Arcipelago delle Isole Eolie are, Posidonia beds (Posidonion oceanicae), Site Code 1120; and, Reefs, Site Code 1170. Both are recorded as category ‘A’, the highest assessment level, with respect to ‘Representativity’ and ‘Conservation’.
The Interpretation Manual of European Union Habitats EUR 28 (European Commission 2013) details, “Reefs can be either biogenic concretions or of geogenic origin. They are hard compact substrata on solid and soft bottoms, which arise from the sea floor in the sublittoral and littoral zone”.
The manual goes on to clarify that:
“Geogenic origin” means: reefs formed by non biogenic substrata;
“Hard compact substrata” are: rocks (including soft rock, e.g. chalk), boulders and cobbles (generally >64 mm in diameter); and,
“Arise from the sea floor" means: the reef is topographically distinct from the surrounding seafloor.
It is clear from this that the cobble reef habitat seen in the images here and swam and snorkelled over by thousands of people each year, are a protected habitat. But the humble moray eel itself? Unprotected and, in the context of SAC’s, without a mention in the optional section, ‘Other important species of flora and fauna’. But why would you want to mention the species there, given their noted importance in the literature as piscovores that, “may have an important role in structuring local reef-fish assemblages” (Matić-Skoko et al. 2011, 2014; Hixon et al. 2012; Zubak et al. 2017); which coupled with the high site fidelity attributed to the species could mean that protecting moray eel explicitly, enhances stability and any associated resilience in these wider fish communities (Hixon et al. 2012)? Following on from this to what extent if any might the presence of M helena be considered characteristic of shallow water cobble reef habitat in the Mediterranean; or, more broadly, could it be a candidate indicator of community health? What might be the knock-on positive effects for artisanal fisheries if applied management strategies included tangible actions based on this and what evidence could be supplied to legitimate such a stance?
Can we identify nursery habitat for M. helena in the Eolian Islands?
There is another inference which might reasonably be made based on the currently available information for immature eels presented here. This inference could be of added value to the recognition of the importance of shallow water cobble habitat. Let’s start by asking this, where might the nursery areas for M. helena be and can we map them based on current evidence (Figure 8)? Note that the depth contours and, MC35 and MC252 marine habitat classifications from the European Nature Information System (EUNIS), were accessed via EMODnet (European Marine Observation and Data Network).
The Interpretation Manual of European Union Habitats EUR 28 (European Commission 2013) details, “Reefs can be either biogenic concretions or of geogenic origin. They are hard compact substrata on solid and soft bottoms, which arise from the sea floor in the sublittoral and littoral zone”.
The manual goes on to clarify that:
“Geogenic origin” means: reefs formed by non biogenic substrata;
“Hard compact substrata” are: rocks (including soft rock, e.g. chalk), boulders and cobbles (generally >64 mm in diameter); and,
“Arise from the sea floor" means: the reef is topographically distinct from the surrounding seafloor.
It is clear from this that the cobble reef habitat seen in the images here and swam and snorkelled over by thousands of people each year, are a protected habitat. But the humble moray eel itself? Unprotected and, in the context of SAC’s, without a mention in the optional section, ‘Other important species of flora and fauna’. But why would you want to mention the species there, given their noted importance in the literature as piscovores that, “may have an important role in structuring local reef-fish assemblages” (Matić-Skoko et al. 2011, 2014; Hixon et al. 2012; Zubak et al. 2017); which coupled with the high site fidelity attributed to the species could mean that protecting moray eel explicitly, enhances stability and any associated resilience in these wider fish communities (Hixon et al. 2012)? Following on from this to what extent if any might the presence of M helena be considered characteristic of shallow water cobble reef habitat in the Mediterranean; or, more broadly, could it be a candidate indicator of community health? What might be the knock-on positive effects for artisanal fisheries if applied management strategies included tangible actions based on this and what evidence could be supplied to legitimate such a stance?
Can we identify nursery habitat for M. helena in the Eolian Islands?
There is another inference which might reasonably be made based on the currently available information for immature eels presented here. This inference could be of added value to the recognition of the importance of shallow water cobble habitat. Let’s start by asking this, where might the nursery areas for M. helena be and can we map them based on current evidence (Figure 8)? Note that the depth contours and, MC35 and MC252 marine habitat classifications from the European Nature Information System (EUNIS), were accessed via EMODnet (European Marine Observation and Data Network).
Fig. 8: An initial attempt to identify suitable areas of potential importance as nursery habitat for Muraena helena, Salina.
Figure 8 is an initial attempt to map potential nursery habitat for M. helena. The requirement for rocky seabed habitat within which the eels can find suitable holes and hideaways is clearly critical (D'Ancona, 1929; Randall & Golani 1995; Jiménez et al. 2007; Matić-Skoko et al. 2011). The only region where I have tangible evidence for this habitat is in the shallow water areas which hug the coast of Salina. The evidence is a mix of limited ground-truthed observation from snorkelling and, in conjunction with this, subsequent visual analysis of the aerial view provided by the Google Satellite layer within a QGIS environment.
The EUNIS habitat layer available via EMODnet suggests that Salina is ringed by largely, soft sediment habitat. Extensive areas of Posidonia oceanica seagrass beds are also predicted (Figure 8). It is important not to undervalue that word, ‘predicted’ and its coupling with ‘suggests’ in the previous sentence. These are necessary to keep in mind because they remind us of the inherent uncertainty in what has been mapped. This ‘conscious awareness of ignorance’ keeps us on our mental toes, slows us down and, with any luck, leads to better subsequent questions and actions. I have attempted to map Posidonia off Santa Marina, Salina previously.
In deeper water there is a ring of circalittoral coarse sediment (CCS) (Figure 8). Crucially, EUNIS indicates that CCS has no identified linkage to reef habitat. So, if we trust our model, it is perhaps more likely than not that hard substrate in this context, if present, will at most be patchy and discontinuous. I have found no published evidence to indicate where, or to what extent if any, reef habitat might actually be present, beyond the narrow band circling the coast (Figure 8, orange polygon). However, it may be that the ring of CCS could be a band within which, for example, slope currents expose some examples of hard substrate.
The area highlighted in a darker blue has a lower boundary that follows the 100m contour around the island. This is an acknowledgement of the evidence from published work that indicates immature M. helena could be found down to at least 80m given the presence of suitable habitat (Matić-Skoko et al. 2011). But this blue layer is simply an ‘area of interest’ for possible future search. I am not suggesting that it can currently be considered as containing potential nursery habitat for the eel, only that it might.
The possible nursery area indicated in Figure 8 is, I trust, more interesting and to a degree, less speculative. Similar rings might be drawn around most of the other islands in the Eolian archipelago. The orange polygon in question here was simply created as a buffer feature extending out 50m from the coastline mapped in QGIS (this distance covers most of the hard substrate visible beneath the sea from the satellite imagery). It was then modified slightly to capture some local detail, such as the small rocky island off the north-west coast of Salina (Plate 8). Not all seabed areas within the area mapped in orange will comprise rocky / cobble habitat. I have observed patches of bare soft sediment as well as seagrass beds within this zone but much of the area is rock habitat of the type which, arguably, could be utilized by moray eels. But a further caveat is of course that we cannot assume all such habitat to be equally suitable, or optimal to an eel searching for a refuge from which to hunt. Note too that the polygon clearly edges around four locations visible in Plate 8 which are man-made structure (concrete piers / harbours) and these are less likely to present suitable habitat.
The area covered by the orange polygon is just under 1km2 (0.982 km2 to be more precise). How much of that area could be occupied nursery habitat is not known so I would suggest it is talked about in terms of being less than 1km2. From my limited snorkel activity, I have observed a maximum of three M. helena in an area of cobble habitat roughly 0.005km2 in extent. I suspect this is an under-estimate of the true number. Perhaps it could be taken as potentially representing a conservative estimate, bearing in mind the concerns / limiting factors mentioned above, in terms of the number of eels the overall area might support. For what it’s worth, the ‘back-of-the-envelope’ calculation based on these numbers suggests that the entire orange envelope might support in the region of 589 juveniles. That may not sound like a lot but equally within the context of the local / regional population, it might well be of structural significance. Given the scant knowledge currently available I would say that at the very least it reinforces the need to ensure loss of, or disturbance to, this protected coastal habitat is avoided. The arguments often rolled out in environmental impact assessment work are that if loss or damage is minimal in terms of the overall extent / magnitude then the result can be dismissed. Unfortunately, such fallacious and unsupported arguments are all too common (Wright et al. 2013).
The presence of Posidonia meadows in close proximity to cobble habitat in the coastal environment here is notable. M. helena are piscivores and well known nocturnal, stalk-and-attack predators within these habitats (Zubak et al. 2017). In fact, it has been shown that piscivores have a strong, “association with the prey community” and as such are likely to be one of the key structuring forces for that community (Zubak et al. 2017). Therefore, one of the reasons supporting / enhancing the value of the identified potential nursery area around Salina may be its close association to such foraging areas.
What are the chances?
In the Spring of 2023, I was lucky enough to find a small, recently washed-up, Anguilliform eel on Salina (Figure 9). The species remains largely, a mystery. The developmental stage as indicated by the presence of red blood cells, clearly visible in the inset image, suggest that it is perhaps either a late glass eel or early elver (Miller 2009). My money is on the latter, given the translucent rather than glassy appearance. Or maybe it is metamorphosing and hence somewhere in-between? I would be very happy to be corrected and to know more, if anyone reading this can shed some light! It’s intriguing because it’s obviously tempting to speculate that as an elver it’s possible it was starting to seek out potential juvenile habitat within which to continue its growth to adulthood.
The EUNIS habitat layer available via EMODnet suggests that Salina is ringed by largely, soft sediment habitat. Extensive areas of Posidonia oceanica seagrass beds are also predicted (Figure 8). It is important not to undervalue that word, ‘predicted’ and its coupling with ‘suggests’ in the previous sentence. These are necessary to keep in mind because they remind us of the inherent uncertainty in what has been mapped. This ‘conscious awareness of ignorance’ keeps us on our mental toes, slows us down and, with any luck, leads to better subsequent questions and actions. I have attempted to map Posidonia off Santa Marina, Salina previously.
In deeper water there is a ring of circalittoral coarse sediment (CCS) (Figure 8). Crucially, EUNIS indicates that CCS has no identified linkage to reef habitat. So, if we trust our model, it is perhaps more likely than not that hard substrate in this context, if present, will at most be patchy and discontinuous. I have found no published evidence to indicate where, or to what extent if any, reef habitat might actually be present, beyond the narrow band circling the coast (Figure 8, orange polygon). However, it may be that the ring of CCS could be a band within which, for example, slope currents expose some examples of hard substrate.
The area highlighted in a darker blue has a lower boundary that follows the 100m contour around the island. This is an acknowledgement of the evidence from published work that indicates immature M. helena could be found down to at least 80m given the presence of suitable habitat (Matić-Skoko et al. 2011). But this blue layer is simply an ‘area of interest’ for possible future search. I am not suggesting that it can currently be considered as containing potential nursery habitat for the eel, only that it might.
The possible nursery area indicated in Figure 8 is, I trust, more interesting and to a degree, less speculative. Similar rings might be drawn around most of the other islands in the Eolian archipelago. The orange polygon in question here was simply created as a buffer feature extending out 50m from the coastline mapped in QGIS (this distance covers most of the hard substrate visible beneath the sea from the satellite imagery). It was then modified slightly to capture some local detail, such as the small rocky island off the north-west coast of Salina (Plate 8). Not all seabed areas within the area mapped in orange will comprise rocky / cobble habitat. I have observed patches of bare soft sediment as well as seagrass beds within this zone but much of the area is rock habitat of the type which, arguably, could be utilized by moray eels. But a further caveat is of course that we cannot assume all such habitat to be equally suitable, or optimal to an eel searching for a refuge from which to hunt. Note too that the polygon clearly edges around four locations visible in Plate 8 which are man-made structure (concrete piers / harbours) and these are less likely to present suitable habitat.
The area covered by the orange polygon is just under 1km2 (0.982 km2 to be more precise). How much of that area could be occupied nursery habitat is not known so I would suggest it is talked about in terms of being less than 1km2. From my limited snorkel activity, I have observed a maximum of three M. helena in an area of cobble habitat roughly 0.005km2 in extent. I suspect this is an under-estimate of the true number. Perhaps it could be taken as potentially representing a conservative estimate, bearing in mind the concerns / limiting factors mentioned above, in terms of the number of eels the overall area might support. For what it’s worth, the ‘back-of-the-envelope’ calculation based on these numbers suggests that the entire orange envelope might support in the region of 589 juveniles. That may not sound like a lot but equally within the context of the local / regional population, it might well be of structural significance. Given the scant knowledge currently available I would say that at the very least it reinforces the need to ensure loss of, or disturbance to, this protected coastal habitat is avoided. The arguments often rolled out in environmental impact assessment work are that if loss or damage is minimal in terms of the overall extent / magnitude then the result can be dismissed. Unfortunately, such fallacious and unsupported arguments are all too common (Wright et al. 2013).
The presence of Posidonia meadows in close proximity to cobble habitat in the coastal environment here is notable. M. helena are piscivores and well known nocturnal, stalk-and-attack predators within these habitats (Zubak et al. 2017). In fact, it has been shown that piscivores have a strong, “association with the prey community” and as such are likely to be one of the key structuring forces for that community (Zubak et al. 2017). Therefore, one of the reasons supporting / enhancing the value of the identified potential nursery area around Salina may be its close association to such foraging areas.
What are the chances?
In the Spring of 2023, I was lucky enough to find a small, recently washed-up, Anguilliform eel on Salina (Figure 9). The species remains largely, a mystery. The developmental stage as indicated by the presence of red blood cells, clearly visible in the inset image, suggest that it is perhaps either a late glass eel or early elver (Miller 2009). My money is on the latter, given the translucent rather than glassy appearance. Or maybe it is metamorphosing and hence somewhere in-between? I would be very happy to be corrected and to know more, if anyone reading this can shed some light! It’s intriguing because it’s obviously tempting to speculate that as an elver it’s possible it was starting to seek out potential juvenile habitat within which to continue its growth to adulthood.
Fig. 9: Anguilliform eel, note the presence of red blood cells visible in the inset image,
Salina, April 9th 2023.
If my inference is correct and this specimen is at the elver stage of development, this could potentially discount one species from the short list of possibilities. Namely, Anguilla anguilla, the common eel because in that species, “Glass eels develop into elvers during upstream migration” and this individual was clearly, a long way from either a suitable estuarine or freshwater environment (Miller, 2009).
In the preceding, pelagic larval stage, this eel would have looked very different with a body that was laterally compressed and transparent (Figure 10) (Miller 2009).
In the preceding, pelagic larval stage, this eel would have looked very different with a body that was laterally compressed and transparent (Figure 10) (Miller 2009).
Fig. 10: The leptocephalus larvae of M. helena as provided by D’ Ancona (1929)
M. helena are oviparous and spawn in the late summer / autumn (Ré & Meneses 2008; Matić-Skoko et al. 2011). The eggs are buoyant and planktonic; spherical and smooth; about 5mm in diameter; and, unpigmented with no oil globules (D'Ancona, 1931-1933; Castle, 1984; Ré & Meneses 2008; Crec’hriou 2015). Post-hatching, eel larvae, or leptocephali, play an essential role in the distribution of the species (Miller 2009; Arai, 2020). Matić-Skoko et al. (2012) found that larvae collected in the south and middle Adriatic Sea using a small pelagic trawl, “ranged in length from 7.7 to 13.1 cm, and were most abundant along the off-shore islands”. Could it be that when such offshore islands are present in the Mediterranean, they represent an important feature in the maintenance of local / regional populations?
The pelagic larval duration (PLD) of up to 2 years is very protracted compared to other reef fishes (Jiménez, 2007). Though the playing field in this respect is not entirely level as, “Leptocephali develop larger sizes and undergo a longer larval period than other fish larvae” (Arai, 2020). Species, with long dispersal times are understood to, “maintain greater geographic-genetic homogeneity among populations than do species having very short PLDs” (Reece et al. 2011). This may explain why M. helena show, “little genetic divergence among specimens from different localities”, as they are not, “sufficiently genetically isolated for endemics to develop” (Jiménez, 2007).
It is perhaps important to note here that Reece et al. (2011) cite Weersing & Toonen (2009) in respect to the above quoted statement. But the 2009 paper makes it clear that in the main, “PLD is, at best, a poor predictor of connectivity” and that, “the significance of the relationship between PLD and genetic structure is driven largely by the presence or absence of a pelagic stage rather than the length of pelagic duration per se”. However, Weersing & Toonen (2009) are studying fish larvae in general, without identifying the type of larvae in question. This is clearly of some significance given, not least, what is known about leptocephali. A point acknowledged indirectly in their closing argument where they underline that accurate estimates of dispersal and population connectivity need, “to consider the interplay of important biophysical, behavioral, and developmental factors through time” (Weersing & Toonen, 2009).
What have moray eels ever done for us?
Lastly, I would like to mention something that I came across whilst writing this article. In 2022, Tiralongo et al. published the first evidence to show that the non-native species Percnon gibbesi, the Sally-lightfoot crab, was being consumed by M. helena. Photographs taken on 24 February 2014 showed a regurgitated Sally Lightfoot in the mouth of a small Mediterranean moray eel. The eel had been taken by a speargun fisherman in the Ionian Sea, just north-east of the Porto di San Giovanni li Cuti, Catania. As already indicated these authors state that M. helena abundance has increased in the last 5-10 years in the recorded area. Interestingly, they then go on to speculate that “the massive presence of the alien crab” may be, in part, one of the drivers behind this reported increase (i.e. through an increase in prey availability). Evidence of the importance of crustacea in the diet of Mediterranean moray eels in shallow waters more generally, has been published (Matić-Skoko et al. 2014). Furthermore, Matić-Skoko et al. (2014) consider M. helena to be an opportunistic species in terms of feeding, with diet reflecting prey availability locally.
In 2007 whilst snorkelling off Salina I noted the presence of large numbers of the Sally-lightfoot crab. Subsequently, over the years, my sense has been that this abundance has not been maintained. It is not unreasonable to infer that this may be in part down to predatory control. Could it be that the presence of P. gibbesi in the shallow sublittoral is now one of the factors supporting favourable conditions for immature moray eels? Was the habitat already fairly optimal or did the availability of this non-native species provide a ‘boost’ to some level of optimality? We cannot know, but perhaps such questions may help frame future research.
And finally…
It is winter as I write this article and my snorkel is dry and unsalted, and I am far from the sea where the Helen of the eels’ swims. But putting these words down has been a useful distraction, sorting my thoughts a little, raising my sight towards the year ahead when I hope to be, at least for a short time, floating head down over cobble habitat, camera at the ready.
Is the fish forgotten? Well maybe or, maybe not.
This article is dedicated to my father, who understood the importance of observation and taking time to enjoy, and to wonder at the beauty, to be found in the world, "What is this life, if full of care, we have no time to stand and stare?" (W.H. Davies, 1911).
References
Adams, F. 1844. The seven books of Paulus Ægineta (Vol. 1). Sydenham Society, London
Arai, T. 2020. Ecology and evolution of migration in the freshwater eels of the genus Anguilla Schrank, 1798. Heliyon, 6(10), e05176. https://doi.org/10.1016/j.heliyon.2020.e05176
Athenaeus. The Deipnosophists. Or Banquet Of The Learned Of Athenaeus. London. Henry G. Bohn, York Street, Covent Garden. 1854.
https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:2013.01.0003
Berg, L.S., 1958. System der rezenten und fossilen Fischartigen und Fische. VEB Verlag der Wissenschaften, Berlin.
Böhlke, E. B., McCosker, J. E., & Böhlke, J. E. 1989. Family Muraenidae. Morays. In Orders Anguilliformes and Saccopharyngiformes: Part 9, Volume 1 (pp. 104–206). Yale University Press. https://doi.org/10.12987/9781933789323-010
Castle, P.H.J., 1984. Notacanthiformes and Anguilliformes: development. p. 62-102. In American Society of Ichthyologists and Herpetologists. Ontogeny and systematics of fishes, based on an international symposium dedicated to the memory of E.H. Ahlstrom, 15-18 August 1983, La Jolla, California. Spec. Publ. Am. Soc. Ichthyol. Herpetol., 1, 1-760.
Catesby, M. & Royal Society of Great Britain 1743. The Natural History of Carolina, Florida and the Bahama Islands, Vol. 2, p. 20. Printed at the expence of the author, and sold by W. Innys and R. Manby, at the West End of St. Paul’s, by Mr. Hauksbee, at the Royal Society House, and by the author, at Mr. Bacon’s in Hoxton. https://www.biodiversitylibrary.org/page/40680119
Crec’hriou, R., Marinaro, J. Y., & Planes, S. 2015. Advance in identification of pelagic eggs of Mediterranean teleostean fish: Development of a new identification key. Vie Milieu.
Cuvier, G. 1829. Le règne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. (Vol. 2, p. 350). https://www.biodiversitylibrary.org/page/31771657
D'Ancona, Umberto 1929. Muraena helena (Linnaeus 1758). Faune ichthyologique de l'Atlantique Nord (1928–1938). Report. https://doi.org/10.17895/ices.pub.24211029.v1
D'Ancona, U. 1931-1933. Apodes (Muraenoidei). Uova, larve e stadi Giovanili di Teleostei: monografia elaborate con l’uso del materiale raccolto e sweriato da Salvatore Lo Bianco. Fauna e Flora Golfo Napoli Monogr. 38, 84 – 146.
European Commission 2013. Interpretation Manual of European Union Habitats EUR 28. DG ENVIRONMENT, Nature ENV B.3, 146pp.
Jacob, C. 2013. The Web of Athenaeus. Hellenic Studies Series 61. Washington, DC: Center for Hellenic Studies. http://nrs.harvard.edu/urn-3:hul.ebook:CHS_JacobC.The_Web_of_Athenaeus.2013.
Jiménez, S., Schönhuth, S., Lozano, I. J., González, J. A., Sevilla, R. G., Diez, A., & Bautista, J. M. 2007. Morphological, Ecological, and Molecular Analyses Separate Muraena augusti from Muraena helena as a Valid Species. Copeia, 1, 101–113. https://doi.org/10.1643/0045-8511(2007)7[101:MEAMAS]2.0.CO;2
Lacépède, & Buffon, Georges Louis Leclerc 1798. Histoire naturelle des poissons 33, 265). Plassan. https://www.biodiversitylibrary.org/page/41229247
Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata [10th revised edition], vol. 1: 824 pp. Laurentius Salvius: Holmiae., available online at https://biodiversitylibrary.org/page/726886
Macrobius, A. A. T. A., Seven Books of the Saturnalia 1400. Volume III, pXV. Publisher not identified.
Matić-Skoko, S., Tutman, P., Petrić, M., Skaramuca, D., Đikić, D., Lisičić, D., & Skaramuca, B. 2011. Mediterranean moray eel Muraena helena (Pisces: Muraenidae): biological indices for life history. Aquatic Biology, 13(3), 275–284. https://doi.org/10.3354/ab00373
Matić-Skoko, S., Vilibić, I., Tutman, P., & Pallaoro, A. 2012. Importance of the South Adriatic Pit as the spawning area. Retention and dispersion of Mediterranean moray eel, Muraena helena leptocephali from deep open waters to coastal area. 138–138. https://www.croris.hr/crosbi/publikacija/prilog-skup/588224
Matić-Skoko, S., Tutman, P., Bojanić Varezić, D., Skaramuca, D., Đikić, D., Lisičić, D., & Skaramuca, B. 2014. Food preferences of the Mediterranean moray eel, Muraena helena (Pisces: Muraenidae), in the southern Adriatic Sea. Marine Biology Research, 10(8), 807–815. https://doi.org/10.1080/17451000.2013.863351
Miller, M. 2009. Ecology of Anguilliform Leptocephali: Remarkable Transparent Fish Larvae of the Ocean Surface Layer. Aqua-BioScience Monographs, 2(4). https://doi.org/10.5047/absm.2009.00204.0001
Nature 1942. Francis Adams (1796-1861). 150, 286–287. Citing an article by Prof. Charles Singer published in the June edition of the Bulletin of the History of Medicine https://doi.org/10.1038/150286d0
Pérez-Lloréns, J. L., Acosta, Y., & Brun, F. G. 2021. Seafood in Mediterranean countries: A culinary journey through history. International Journal of Gastronomy and Food Science, 26, 100437. https://doi.org/10.1016/j.ijgfs.2021.100437
Pliny the Elder, The Natural History, ed. John Bostock and H. T. Riley, in the Perseus Digital Library, http://data.perseus.org/citations/urn:cts:latinLit:phi0978.phi001.perseus-eng1:1.dedication.
Randall, J. E. 1969. How Dangerous is the Moray Eel? Australian Natural History, 16(6), 177–182.
Randall, J. E., & Golani, D. 1995. Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56(3), 849–880.
Ré, P., & Meneses, I. 2008. Early stages of marine fishes occurring in the Iberian Peninsula. Early stages of marine fishes occurring in the Iberian Peninsula. https://comum.rcaap.pt/handle/10400.26/48140
Reece, J. S., Bowen, B. W., Smith, D. G., & Larson, A. 2011. Comparative phylogeography of four Indo-Pacific moray eel species (Muraenidae) reveals comparable ocean-wide genetic connectivity despite five-fold differences in available adult habitat. Marine Ecology Progress Series, 437, 269–277. https://doi.org/10.3354/meps09248
Reece, J. S., & Mehta, R. S. 2013. Evolutionary history of elongation and maximum body length in moray eels (Anguilliformes: Muraenidae): Evolution of Body Elongation and Size in Moray Eels. Biological Journal of the Linnean Society, 109(4), 861–875. https://doi.org/10.1111/bij.12098
Singleton, R. H. 1943. LI Milton’s Comus and the Comus of Erycius Puteanus. PMLA/Publications of the Modern Language Association of America, 58(4_1), 949–957. https://doi.org/10.2307/458920
Tiralongo, F., Akyol, O., Al Mabruk, S. A. A., Battaglia, P., Beton, D., Bitlis, B., Borg, J. A., Bouchoucha, M., Çinar, M. E., Crocetta, F., Dragičević, B., Jdulčić, J. D., Evangelopoulos, A., Jevans, J., Fortič, A., Gauff, R. P. M., Georgiadis, C. G., Gökoğlu, M., Daniele Grech, D., … Velasquez, X. 2022. New Alien Mediterranean Biodiversity Records (August 2022). Mediterranean Marine Science, 23(3), 725–747. https://doi.org/10.12681/mms.31228
Vehling, J. D. 1977. Cookery and Dining in Imperial Rome: A Bibliography, Critical Review, and Translation of the Ancient Book Known as Apicius de Re Coquinaria: Now for the First Time Rendered Into English. United Kingdom: Dover Publications.
Weersing, K., & Toonen, R. 2009. Population genetics, larval dispersal, and connectivity in marine systems. Marine Ecology Progress Series, 393, 1–12. https://doi.org/10.3354/meps08287
Wright, A. J., Dolman, S. J., Jasny, M., Parsons, E. C. M., Schiedek, D., & Young, S. B. 2013. Myth and Momentum: A Critique of Environmental Impact Assessments. Journal of Environmental Protection, 04(08), 72–77. https://doi.org/10.4236/jep.2013.48A2009
Zubak, I., Kruschel, C., & Schultz, S. 2017. Predators structure fish communities in Posidonia oceanica meadows: Meta-analysis of available data across the Mediterranean basin. Marine Ecology Progress Series, 566, 145–157. https://doi.org/10.3354/meps12038
Websites:
Global Fishing Watch
https://globalfishingwatch.org/
Natura 2000 Viewer, Fondali dell'Arcipelago delle Isole Eolie, Italy (ITA030041)
https://natura2000.eea.europa.eu/?sitecode=ITA030041&views=Sites_View
Wiktionary contributors, "deipnosophist," Wiktionary, https://en.wiktionary.org/w/index.php?title=deipnosophist&oldid=82036239
(accessed November 28, 2024).
With grateful thanks to Simone Follone for sharing his local knowledge; and, Giuseppina Crisafi for translating the Latin text and her guidance and help in this respect.
The pelagic larval duration (PLD) of up to 2 years is very protracted compared to other reef fishes (Jiménez, 2007). Though the playing field in this respect is not entirely level as, “Leptocephali develop larger sizes and undergo a longer larval period than other fish larvae” (Arai, 2020). Species, with long dispersal times are understood to, “maintain greater geographic-genetic homogeneity among populations than do species having very short PLDs” (Reece et al. 2011). This may explain why M. helena show, “little genetic divergence among specimens from different localities”, as they are not, “sufficiently genetically isolated for endemics to develop” (Jiménez, 2007).
It is perhaps important to note here that Reece et al. (2011) cite Weersing & Toonen (2009) in respect to the above quoted statement. But the 2009 paper makes it clear that in the main, “PLD is, at best, a poor predictor of connectivity” and that, “the significance of the relationship between PLD and genetic structure is driven largely by the presence or absence of a pelagic stage rather than the length of pelagic duration per se”. However, Weersing & Toonen (2009) are studying fish larvae in general, without identifying the type of larvae in question. This is clearly of some significance given, not least, what is known about leptocephali. A point acknowledged indirectly in their closing argument where they underline that accurate estimates of dispersal and population connectivity need, “to consider the interplay of important biophysical, behavioral, and developmental factors through time” (Weersing & Toonen, 2009).
What have moray eels ever done for us?
Lastly, I would like to mention something that I came across whilst writing this article. In 2022, Tiralongo et al. published the first evidence to show that the non-native species Percnon gibbesi, the Sally-lightfoot crab, was being consumed by M. helena. Photographs taken on 24 February 2014 showed a regurgitated Sally Lightfoot in the mouth of a small Mediterranean moray eel. The eel had been taken by a speargun fisherman in the Ionian Sea, just north-east of the Porto di San Giovanni li Cuti, Catania. As already indicated these authors state that M. helena abundance has increased in the last 5-10 years in the recorded area. Interestingly, they then go on to speculate that “the massive presence of the alien crab” may be, in part, one of the drivers behind this reported increase (i.e. through an increase in prey availability). Evidence of the importance of crustacea in the diet of Mediterranean moray eels in shallow waters more generally, has been published (Matić-Skoko et al. 2014). Furthermore, Matić-Skoko et al. (2014) consider M. helena to be an opportunistic species in terms of feeding, with diet reflecting prey availability locally.
In 2007 whilst snorkelling off Salina I noted the presence of large numbers of the Sally-lightfoot crab. Subsequently, over the years, my sense has been that this abundance has not been maintained. It is not unreasonable to infer that this may be in part down to predatory control. Could it be that the presence of P. gibbesi in the shallow sublittoral is now one of the factors supporting favourable conditions for immature moray eels? Was the habitat already fairly optimal or did the availability of this non-native species provide a ‘boost’ to some level of optimality? We cannot know, but perhaps such questions may help frame future research.
And finally…
It is winter as I write this article and my snorkel is dry and unsalted, and I am far from the sea where the Helen of the eels’ swims. But putting these words down has been a useful distraction, sorting my thoughts a little, raising my sight towards the year ahead when I hope to be, at least for a short time, floating head down over cobble habitat, camera at the ready.
Is the fish forgotten? Well maybe or, maybe not.
This article is dedicated to my father, who understood the importance of observation and taking time to enjoy, and to wonder at the beauty, to be found in the world, "What is this life, if full of care, we have no time to stand and stare?" (W.H. Davies, 1911).
References
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Arai, T. 2020. Ecology and evolution of migration in the freshwater eels of the genus Anguilla Schrank, 1798. Heliyon, 6(10), e05176. https://doi.org/10.1016/j.heliyon.2020.e05176
Athenaeus. The Deipnosophists. Or Banquet Of The Learned Of Athenaeus. London. Henry G. Bohn, York Street, Covent Garden. 1854.
https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:2013.01.0003
Berg, L.S., 1958. System der rezenten und fossilen Fischartigen und Fische. VEB Verlag der Wissenschaften, Berlin.
Böhlke, E. B., McCosker, J. E., & Böhlke, J. E. 1989. Family Muraenidae. Morays. In Orders Anguilliformes and Saccopharyngiformes: Part 9, Volume 1 (pp. 104–206). Yale University Press. https://doi.org/10.12987/9781933789323-010
Castle, P.H.J., 1984. Notacanthiformes and Anguilliformes: development. p. 62-102. In American Society of Ichthyologists and Herpetologists. Ontogeny and systematics of fishes, based on an international symposium dedicated to the memory of E.H. Ahlstrom, 15-18 August 1983, La Jolla, California. Spec. Publ. Am. Soc. Ichthyol. Herpetol., 1, 1-760.
Catesby, M. & Royal Society of Great Britain 1743. The Natural History of Carolina, Florida and the Bahama Islands, Vol. 2, p. 20. Printed at the expence of the author, and sold by W. Innys and R. Manby, at the West End of St. Paul’s, by Mr. Hauksbee, at the Royal Society House, and by the author, at Mr. Bacon’s in Hoxton. https://www.biodiversitylibrary.org/page/40680119
Crec’hriou, R., Marinaro, J. Y., & Planes, S. 2015. Advance in identification of pelagic eggs of Mediterranean teleostean fish: Development of a new identification key. Vie Milieu.
Cuvier, G. 1829. Le règne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. (Vol. 2, p. 350). https://www.biodiversitylibrary.org/page/31771657
D'Ancona, Umberto 1929. Muraena helena (Linnaeus 1758). Faune ichthyologique de l'Atlantique Nord (1928–1938). Report. https://doi.org/10.17895/ices.pub.24211029.v1
D'Ancona, U. 1931-1933. Apodes (Muraenoidei). Uova, larve e stadi Giovanili di Teleostei: monografia elaborate con l’uso del materiale raccolto e sweriato da Salvatore Lo Bianco. Fauna e Flora Golfo Napoli Monogr. 38, 84 – 146.
European Commission 2013. Interpretation Manual of European Union Habitats EUR 28. DG ENVIRONMENT, Nature ENV B.3, 146pp.
Jacob, C. 2013. The Web of Athenaeus. Hellenic Studies Series 61. Washington, DC: Center for Hellenic Studies. http://nrs.harvard.edu/urn-3:hul.ebook:CHS_JacobC.The_Web_of_Athenaeus.2013.
Jiménez, S., Schönhuth, S., Lozano, I. J., González, J. A., Sevilla, R. G., Diez, A., & Bautista, J. M. 2007. Morphological, Ecological, and Molecular Analyses Separate Muraena augusti from Muraena helena as a Valid Species. Copeia, 1, 101–113. https://doi.org/10.1643/0045-8511(2007)7[101:MEAMAS]2.0.CO;2
Lacépède, & Buffon, Georges Louis Leclerc 1798. Histoire naturelle des poissons 33, 265). Plassan. https://www.biodiversitylibrary.org/page/41229247
Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata [10th revised edition], vol. 1: 824 pp. Laurentius Salvius: Holmiae., available online at https://biodiversitylibrary.org/page/726886
Macrobius, A. A. T. A., Seven Books of the Saturnalia 1400. Volume III, pXV. Publisher not identified.
Matić-Skoko, S., Tutman, P., Petrić, M., Skaramuca, D., Đikić, D., Lisičić, D., & Skaramuca, B. 2011. Mediterranean moray eel Muraena helena (Pisces: Muraenidae): biological indices for life history. Aquatic Biology, 13(3), 275–284. https://doi.org/10.3354/ab00373
Matić-Skoko, S., Vilibić, I., Tutman, P., & Pallaoro, A. 2012. Importance of the South Adriatic Pit as the spawning area. Retention and dispersion of Mediterranean moray eel, Muraena helena leptocephali from deep open waters to coastal area. 138–138. https://www.croris.hr/crosbi/publikacija/prilog-skup/588224
Matić-Skoko, S., Tutman, P., Bojanić Varezić, D., Skaramuca, D., Đikić, D., Lisičić, D., & Skaramuca, B. 2014. Food preferences of the Mediterranean moray eel, Muraena helena (Pisces: Muraenidae), in the southern Adriatic Sea. Marine Biology Research, 10(8), 807–815. https://doi.org/10.1080/17451000.2013.863351
Miller, M. 2009. Ecology of Anguilliform Leptocephali: Remarkable Transparent Fish Larvae of the Ocean Surface Layer. Aqua-BioScience Monographs, 2(4). https://doi.org/10.5047/absm.2009.00204.0001
Nature 1942. Francis Adams (1796-1861). 150, 286–287. Citing an article by Prof. Charles Singer published in the June edition of the Bulletin of the History of Medicine https://doi.org/10.1038/150286d0
Pérez-Lloréns, J. L., Acosta, Y., & Brun, F. G. 2021. Seafood in Mediterranean countries: A culinary journey through history. International Journal of Gastronomy and Food Science, 26, 100437. https://doi.org/10.1016/j.ijgfs.2021.100437
Pliny the Elder, The Natural History, ed. John Bostock and H. T. Riley, in the Perseus Digital Library, http://data.perseus.org/citations/urn:cts:latinLit:phi0978.phi001.perseus-eng1:1.dedication.
Randall, J. E. 1969. How Dangerous is the Moray Eel? Australian Natural History, 16(6), 177–182.
Randall, J. E., & Golani, D. 1995. Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56(3), 849–880.
Ré, P., & Meneses, I. 2008. Early stages of marine fishes occurring in the Iberian Peninsula. Early stages of marine fishes occurring in the Iberian Peninsula. https://comum.rcaap.pt/handle/10400.26/48140
Reece, J. S., Bowen, B. W., Smith, D. G., & Larson, A. 2011. Comparative phylogeography of four Indo-Pacific moray eel species (Muraenidae) reveals comparable ocean-wide genetic connectivity despite five-fold differences in available adult habitat. Marine Ecology Progress Series, 437, 269–277. https://doi.org/10.3354/meps09248
Reece, J. S., & Mehta, R. S. 2013. Evolutionary history of elongation and maximum body length in moray eels (Anguilliformes: Muraenidae): Evolution of Body Elongation and Size in Moray Eels. Biological Journal of the Linnean Society, 109(4), 861–875. https://doi.org/10.1111/bij.12098
Singleton, R. H. 1943. LI Milton’s Comus and the Comus of Erycius Puteanus. PMLA/Publications of the Modern Language Association of America, 58(4_1), 949–957. https://doi.org/10.2307/458920
Tiralongo, F., Akyol, O., Al Mabruk, S. A. A., Battaglia, P., Beton, D., Bitlis, B., Borg, J. A., Bouchoucha, M., Çinar, M. E., Crocetta, F., Dragičević, B., Jdulčić, J. D., Evangelopoulos, A., Jevans, J., Fortič, A., Gauff, R. P. M., Georgiadis, C. G., Gökoğlu, M., Daniele Grech, D., … Velasquez, X. 2022. New Alien Mediterranean Biodiversity Records (August 2022). Mediterranean Marine Science, 23(3), 725–747. https://doi.org/10.12681/mms.31228
Vehling, J. D. 1977. Cookery and Dining in Imperial Rome: A Bibliography, Critical Review, and Translation of the Ancient Book Known as Apicius de Re Coquinaria: Now for the First Time Rendered Into English. United Kingdom: Dover Publications.
Weersing, K., & Toonen, R. 2009. Population genetics, larval dispersal, and connectivity in marine systems. Marine Ecology Progress Series, 393, 1–12. https://doi.org/10.3354/meps08287
Wright, A. J., Dolman, S. J., Jasny, M., Parsons, E. C. M., Schiedek, D., & Young, S. B. 2013. Myth and Momentum: A Critique of Environmental Impact Assessments. Journal of Environmental Protection, 04(08), 72–77. https://doi.org/10.4236/jep.2013.48A2009
Zubak, I., Kruschel, C., & Schultz, S. 2017. Predators structure fish communities in Posidonia oceanica meadows: Meta-analysis of available data across the Mediterranean basin. Marine Ecology Progress Series, 566, 145–157. https://doi.org/10.3354/meps12038
Websites:
Global Fishing Watch
https://globalfishingwatch.org/
Natura 2000 Viewer, Fondali dell'Arcipelago delle Isole Eolie, Italy (ITA030041)
https://natura2000.eea.europa.eu/?sitecode=ITA030041&views=Sites_View
Wiktionary contributors, "deipnosophist," Wiktionary, https://en.wiktionary.org/w/index.php?title=deipnosophist&oldid=82036239
(accessed November 28, 2024).
With grateful thanks to Simone Follone for sharing his local knowledge; and, Giuseppina Crisafi for translating the Latin text and her guidance and help in this respect.
Citation: This article was originally published in the Autumn 2025 edition of the Bulletin of the Porcupine Marine Natural History Society and can be cited as follows:
Barfield, P. D. (2025). Notes, observations and questions concerning Mediterranean moray eel, Muraena helena Linnaeus, 1758. Bulletin of the Porcupine Marine Natural History Society, No. 24, 59-73. ISSN 2054-7137.
Note: Some small edits and additions have been made to this online version of the article.
© Sea-nature Studies, 2025. All rights reserved in all media.