8
Aristotle’s observations of the foraging interactions of the red mullet (Mullidae: Mullus spp) and sea bream (Sparidae: Diplodus spp) JASON A. TIPTON St John’s College, Box 2800, Annapolis, Maryland 21404, USA (email: [email protected]). ABSTRACT: A brief but detailed observation which Aristotle made regarding the foraging behavior of the red mullet (Mullus surmuletus) is examined closely in order to better understand the substance of the discussion. Aristotle’s description of the heterospecific red mullet/sea bream foraging behavior is evaluated in light of contemporary observations and life history characteristics. Possible methods, including underwater observations with the use of diving equipment and observations in fish ponds, employed by Aristotle in making these observations are discussed. I also speculate on the ways in which the red mullet/sea bream interaction might be thought of as gregarious behavior according to Aristotle. KEY WORDS: Lesvos – natural history – Aristotelian philosophy – Aristotelian zoology – feeding behavior – Pisces. RED MULLET IN ANTIQUITY It is interesting and illuminating for contemporary biologists and philosophers of Aristotle to look closely at the detailed observations made by him. That great advocate of studying Aristotle’s biological works, David Balme (1987), discovered that observing the organisms and traits of interest to Aristotle actually helps in the understanding of his works in general (Balme 1987: 16–17): Much of this criticism [of these biological writings] arose in the nineteenth and early twentieth centuries from armchair naturalists who disbelieved Aristotle’s reports and thought them too silly for a great philosopher ... I confess that I was still blaming Aristotle for swallowing the story about buffaloes projecting their dung at enemies, until in 1983 I saw a picture on television of hippopotamuses doing just that. In a similar vein, Lennox’s (1983) examination of echinoderm reproductive parts is so illuminating because of its careful analysis of the empirical side of Aristotle. Buddington and Diamond (1986) revisited Aristotle’s ruminations on the function of the pyloric caeca in fish. Voultsiadou and Vafidis (2007) survey Aristotle’s knowledge of marine invertebrates. Brock (2004) has recently pointed out that Aristotle’s discussion of sperm competition in birds is largely accurate, despite the scathing remarks of the translators and commentators. This examination focuses on Aristotle’s observations of a small fish found in the Aegean. Red and striped red mullets (Mullus spp), sometimes called goatfish, are famous food-fishes in the Mediterranean, from ancient times up to the present day. There is not sufficient evidence to determine whether Aristotle distinguished between the red mullet (M. surmuletus) and the striped red mullet (M. barbatus); in what follows, I will assume that the discussion applies equally to both. Red mullets are also known as goatfish because of Archives of natural history 35 (1): 164–171. 2008 # The Society for the History of Natural History DOI: 10.3366/E0260954108000156

Aristotle's observations of the foraging interactions of the red mullet (Mullidae: Mullus spp) and sea bream (Sparidae: Diplodus spp

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Aristotle’s observations of the foraging interactionsof the red mullet (Mullidae: Mullus spp) and sea bream(Sparidae: Diplodus spp)

JASON A. TIPTON

St John’s College, Box 2800, Annapolis, Maryland 21404, USA (email: [email protected]).

ABSTRACT: A brief but detailed observation which Aristotle made regarding the foraging behavior

of the red mullet (Mullus surmuletus) is examined closely in order to better understand the substance

of the discussion. Aristotle’s description of the heterospecific red mullet/sea bream foraging behavior

is evaluated in light of contemporary observations and life history characteristics. Possible methods,

including underwater observations with the use of diving equipment and observations in fish ponds,

employed by Aristotle in making these observations are discussed. I also speculate on the ways in

which the red mullet/sea bream interaction might be thought of as gregarious behavior according to

Aristotle.

KEY WORDS: Lesvos – natural history – Aristotelian philosophy – Aristotelian zoology – feeding

behavior – Pisces.

RED MULLET IN ANTIQUITY

It is interesting and illuminating for contemporary biologists and philosophers of Aristotle to

look closely at the detailed observations made by him. That great advocate of studying

Aristotle’s biological works, David Balme (1987), discovered that observing the organisms

and traits of interest to Aristotle actually helps in the understanding of his works in general

(Balme 1987: 16–17):

Much of this criticism [of these biological writings] arose in the nineteenth and early twentieth centuries from

armchair naturalists who disbelieved Aristotle’s reports and thought them too silly for a great philosopher . . . I

confess that I was still blaming Aristotle for swallowing the story about buffaloes projecting their dung at

enemies, until in 1983 I saw a picture on television of hippopotamuses doing just that.

In a similar vein, Lennox’s (1983) examination of echinoderm reproductive parts is so

illuminating because of its careful analysis of the empirical side of Aristotle. Buddington

and Diamond (1986) revisited Aristotle’s ruminations on the function of the pyloric caeca

in fish. Voultsiadou and Vafidis (2007) survey Aristotle’s knowledge of marine

invertebrates. Brock (2004) has recently pointed out that Aristotle’s discussion of sperm

competition in birds is largely accurate, despite the scathing remarks of the translators and

commentators.

This examination focuses on Aristotle’s observations of a small fish found in the

Aegean. Red and striped red mullets (Mullus spp), sometimes called goatfish, are famous

food-fishes in the Mediterranean, from ancient times up to the present day. There is not

sufficient evidence to determine whether Aristotle distinguished between the red mullet

(M. surmuletus) and the striped red mullet (M. barbatus); in what follows, I will assume that

the discussion applies equally to both. Red mullets are also known as goatfish because of

Archives of natural history 35 (1): 164–171. 2008 # The Society for the History of Natural History

DOI: 10.3366/E0260954108000156

the two stout barbels they possess under their chins, which give them a goat-like beard

(Figure 1); Pliny (Rackham 1940: 205–207)1 said that these mullets are distinguished by

a double beard on the lower lip. Pliny also identified several varieties of the red mullet as

some of the best food fish and even described some of the sauces the epicureans of the day

recommended with its consumption (Rackham 1940: 205–207). Seneca (Gummere 1925:

85)2 told of a large mullet given to Tiberius:

A mullet of monstrous size was presented to the Emperor Tiberius. They say it weighed four and one half

pounds (and why should I not tickle the palates of certain epicures by mentioning its weight?). Tiberius ordered

it to be sent to the fish-market and put up for sale, remarking: ‘I shall be taken entirely by surprise, my friends,

if either Apicius or P. Octavius does not buy that mullet.’ The guess came true beyond his expectation: the two

men bid, and Octavius won, thereby acquiring a great reputation among his intimates because he had bought for

five thousand sesterces a fish which the Emperor had sold, and which even Apicius did not succeed in buying.

To pay such a price was disgraceful for Octavius, but not for the individual who purchased the fish in order to

present it to Tiberius, – though I should be inclined to blame the latter as well; but at any rate he admired a gift

of which he thought Caesar worthy.

As is clear from both Pliny and Seneca, the red mullet was, in antiquity, a great delicacy

(Davidson 1997: 8) and continues to be a highly prized fish today in the Mediterranean.

Figure 1. Striped red mullet (Mullus surmuletus) in a fish market in Mytilini, Greece. Note the robust chin barbels

under the mouth. The bulk of Aristotle’s work in natural history was done in the North Aegean, centered around

Mytilini on the island of Lesvos. In fact, Lesvos is among the few place names mentioned by Aristotle (HA 621b12,

621b22). It is generally acknowledged that Aristotle traveled to Lesvos with Theophrastus shortly after Plato’s

death. All photographs taken by J. Tipton.

ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM 165

ARISTOTLE’S INTEREST IN THE RED MULLET

While they are renowned for the table, Aristotle made no mention of this fact in his

observations of the red mullet (trigle) scattered through the History of animals (HA). Among

other things, Aristotle discussed the fact that the red mullet occurs in lagoons (HA 598a20)3,

has a diet consisting of sea-weed and shellfish (HA 591b12), and he noted some

characteristics of its spawning (HA 543a5, HA 570b25). And while most carnivorous fish

tend to wander, the red mullet does not (HA 621b7); this is an exception to Aristotle’s

general observations that fishes “feed in those places in which they were formed and in

places like them, for the food proper to each is in these places” (HA 621b3). Aristotle also

described its interactions with another fish, the sargue or bream: “The sargue encroaches on

the red mullet, and when the red mullet has stirred up the mud (for it can dig) and has gone

away, the sargue comes down and feeds, and prevents those weaker than itself from coming

down there too” (HA 591b19).

The sargue identified by Aristotle as encroaching on the red mullet is most likely a sea-

bream of the Sparidae (Thompson 1947); it seems probable that in this passage Aristotle was

describing one or more of the fishes modern ichthyologists recognize as species of Diplodus,

for example D. vulgaris, D. sargos and D. annularis.

Aristotle’s discussion of the interaction between the red mullet and the sargue comes in a

discussion of certain feeding behaviors of various types of fishes. For example, Aristotle

noted that the parrot-wrasse (skaros) was thought to ruminate like a quadruped. This

discussion of the different ways in which fishes and other marine animals (for example,

cephalopods and marine mammals) feed and behave is part of Aristotle’s general inquiry

that included articulating the differences with respect to the lives and activities of animals

(HA 487b34). In several restatements of this aspect of his zoological inquiry, Aristotle

included the dispositions (ethe) alongside the lives and activities of animals (for example,

HA 487a14).

Aristotle was correct in suggesting the red mullet feeds by stirring up the mud or sand.

It is perhaps a bit surprising that Thompson, in his excellent reference A glossary of Greek

fishes (1947), does not report this aspect of the red mullet’s behavior nor Aristotle’s thoughts

on the feeding interactions with other fish. I would suggest that the family now known as

Triglidae was associated in Aristotle with the red mullet (trigle) because of their similar

life histories. Triglids use free pelvic fins to walk along the sea floor searching for food,

similar to the red mullet’s use of chin barbels. The striped red mullet (Mullus surmuletus)

and the red mullet (M. barbatus) use modified, freely moving chin barbels (Figure 1) to

probe the sand, searching for buried invertebrates including crustaceans, amphipods,

polychaetes and mollusks (Hureau 1996). Not only is the red mullet connected to the sandy

bottom for its food, it also lays its eggs, according to Aristotle, in the sand or mud after it

has warmed (HA 570b23). And confirming Aristotle’s own observations, I have observed

M. surmuletus foraging in conjunction with other fishes including breams (Diplodus spp) and

wrasses (Labridae) (see Figures 2 and 3). It is worth noting that Pliny suggested that it was

one of the varieties of the red mullet that followed a bream digging in the sand (Rackham

1940: 207)4, which is to reverse the foraging relation that Aristotle suggests. The breams,

such as Diplodus vulgaris, generally occupy littoral waters over rocky or sandy bottoms in

depths to 90 meters and consume crustaceans, mollusks and worms (Bauchot and Hureau

1996). Wrasses, such as the gray wrasse (Symphodus cinereus) observed foraging with

the red mullet, are generally in the vicinity of eel grass beds in 1 to 20 meters of water

166 ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM

and eat shrimps, amphipods, isopods, small gastropods and bivalves (Quignard and Pras

1996).

Aristotle emphasized that the sargue encroached on the red mullet, and chased away

potential competition. This behavior, along with the mullet’s foraging or “stirring up the

Figure 2. Mullus surmuletus (right) foraging with

Diplodus vulgaris and Symphodus cinereus in approx-

imately 3.5m of water; about 2km north of Thermi,

Lesvos, Greece.

Figure 3. Mullus surmuletus (in the middle) flanked by

Diplodus vulgaris (foreground) and Symphodus cinereus

(background) in a co-foraging aggregation; Lesvos,

Greece (as Figure 2).

Figure 4. Note the chin barbels extended as Mullus surmuletus forages for food. Symphodus cinereus accompanies

the striped red mullet in a search of the sandy bottom; Lesvos, Greece (as Figure 2).

ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM 167

bottom” (HA 591b19) seems to describe a particular case of what has been termed the

scrounger-producer relationship (Barnard and Sibly 1981; see also De Pirro et al. 1999).

Scroungers, like the Diplodus vulgaris (Figures 2, 3 and 4), appear to reduce the costs of

exploiting a resource by letting producers, such as Mullus surmuletus in this case, invest the

necessary time and energy in foraging and then usurping the results of those efforts (Barnard

and Sibly 1981). Red mullet are prone to infestation by sea-lice (HA 557a26); lice, Aristotle

stated, are not carnivorous but live on the juices of living flesh, they are parasitic.

Analogously, the sargue parasitizes the work done by the red mullet.

De Pirro et al. (1999) observed Mullus surmuletus foraging in conjunction with two

wrasses (Coris julis and Symphodus cinereus). They reported that heteroaggregation, the co-

foraging of individuals of different species, prevailed in M. surmuletus. M. surmuletus, in

feeding, was more likely than not to be observed in a group with individuals of different

species.

If Mullus surmuletus can be said to form heteroaggregations, it is not clear to me whether

Aristotle would describe groups of red mullet and bream as gregarious or social. In some

sense, they share a common work or activity, which is said of social animals in general (HA

488a8). Saying an animal is gregarious (agelaios) is a more general claim than saying it is

social (politicos); hence, some of the gregarious animals are social (HA 488a3). In the waters

around Lesvos, I have often observed four to eight striped red mullet (M. surmuletus)

foraging in close association, which agrees with Aristotle’s observation that they are

generally shoaling fish (HA 610b3). The sargue also was described as generally shoaling. My

sense is that this red mullet/sea bream association would not be considered as social because

it is composed of different kinds or species and is transitory and opportunistic, they do not

live in close communities; however, the association might be one that exhibits

gregariousness according to Aristotle. Such a conjecture is supported by the fact that

Aristotle did observe that the sea basse (labrax) and the grey mullet shoal together, are

gregarious, despite the hostility between the kinds. Fishes of different kinds will shoal

together and form associations when their habitats and prey items overlap (HA 610b13),

which seems to be the case for the red mullet and sargue.

SOME THOUGHTS ON ARISTOTLE’S “METHODOLOGY”

This kind of co-foraging of the red mullet and sea bream is remarkable. What seems to me

equally remarkable is that Aristotle recorded this aspect of the life history of these fish. How

did he, or his source, make such observations? Did they observe the sandy bottom from

shore? While I have observed different fish from shore, noting their movements and

behaviors in reference to Aristotle’s own work (Tipton 2006), it is difficult for me to imagine

someone observing this cooperative foraging from shore. Did they see these fish interact in

cisterns set up to hold fish (Figure 5)? Given its status as a delicacy, one could easily imagine

considerable resources being devoted to the maintenance of a fresh supply of red mullet.

Pliny described the extraordinary efforts made in the construction and maintenance of oyster

and fish ponds (Rackham 1940: 277).5 Perhaps if they were in such an enclosure or pond of

some kind, Aristotle or his source could more easily have observed their behavior. Could it

be that this remarkable behavior was observed and noted while diving? Anyone who has

swum without a mask could report that it would be very difficult to see with enough clarity

to make the detailed observations of Mullus species foraging in even three or four meters of

168 ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM

water. Perhaps Greek divers had a capacity or a technique that allowed such a feat. In his

thoughts on ancient diving, Frost (1968: 181) made the following observation:

Both Minoan and Mycenaean graphic arts reveal a lively appreciation for undersea life: octopuses, shellfish,

sponges, and their rocky environment are depicted with charm and a high degree of accuracy as well. Sponges,

it might be added, are almost never identified as such by modern commentaries on vase paintings, but anyone

who has dived into the Aegean recognizes them immediately for what they are.

Aristotle did describe some of the equipment that might have been available to divers during

his time. In his Parts of animals (PA), he described a device which resembles a snorkel:

“Some divers, when they go down into the sea, provide themselves with a breathing

machine, by means of which they can inhale the air from above the surface while they

remain for a long time in the water” (PA 659a8–13). He also described the way in which a

cauldron can be lowered into the water, upside down, keeping some air inside which could

be used by a diver, a sort of diving bell (Problems 960b32).

Both of these devices could aid a diver in antiquity but would do nothing to correct that

individual’s extremely blurry vision under the water. With this problem in mind, it is

interesting to note that lenses made of rock crystal have been discovered at archaeological

sites across the Mediterranean (James and Thorpe 1994: 157–163). They are thought to have

been useful as an aid to the eyes in doing detailed engraving and sculpting. While such

lenses could be used to magnify or concentrate the rays of the sun to burn – as Strepsiades

described in Aristophanes’ Clouds – I wonder if such things could have been made into a

crude mask for diving. The detailed description of the feeding habits of the red mullet and

the striped red mullet might suggest the use of something like a mask. If what Aristotle

described was a kind of snorkel-like instrument, one might speculate on the existence of an

accompanying mask.

In the description of the interaction between the sargue and red mullet, Aristotle observed

that the former can take advantage of the work of the latter. That work includes digging or

stirring up of the mud, which is made possible by means of the chin barbels. Aristotle not

only provided many aspects of the life history of the red mullet and sea bream, he also

described this peculiar interaction, described as heterospecific aggregation by researchers,

for example De Pirro et al. (1999), studying some of the same species and problems today.

Figure 5. An ancient holding tank for fish erected near the south harbor in Mytilini, Greece, and dating from the

first century CE. The holes seen one third of the way up the wall are thought to be hiding places and habitat niches

for fish and invertebrates.6

ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM 169

ACKNOWLEDGEMENTS

I wish to thank Hector Williams for sharing his experience with the archaeological remains of fish tanks on Lesvos.

I also appreciate the helpful comments of an anonymous reviewer that served to improve my manuscript.

NOTES

1 Pliny’s discussion of the red mullet comes in Natural history, Book IX, Chapter XXX.2 This discussion is to be found in Epistle XCV.3 All references to the works of Aristotle are based on the conventional Bekker numbers. The Bekker numbers

identify the page and line numbers used by all modern scholarly editions of Aristotle. With respect to the History of

animals, translations by Balme (1991) and Peck (1965, 1970) are used. For the Parts of animals, the translation by

Lennox (2001) is used here. I also reference Hett’s (1937) edition of Problems.4 Natural history, Book IX, LXXIX–LXXX.5 Natural History, Book IX, XXX.6 Hector Williams, per. comm., July 2005.

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Received 27 September 2007. Accepted 5 December 2007.

ARISTOTLE’S OBSERVATIONS ON RED MULLET AND SEA BREAM 171