Nothing but the Tooth: A Dental Odyssey

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The Jaws of the Piranha

Natural History of the Piranha

The name ‘piranha’ conjures up an image of a steaming Amazon River in which a hapless animal straying into it suddenly disappears underwater in a broiling mass of blood. A few minutes later, all is calm – the only trace remaining of the animal is a skeleton clinically stripped of its flesh. This was the type of description given by U.S. President Theodore Roosevelt in his 1914 book Through the Brazilian Wilderness. He stated ‘They will snap the finger off a hand incautiously trailed in the water…They are the fish that eats men when it can get the chance…Blood in the water excites them to madness…’. However, this spectacle could have been purposely orchestrated for him with a shoal of piranhas held in the river without food for some time. This episode convinced Roosevelt that piranhas are ‘the most ferocious fish in the world’.

The fearsome reputation of the piranha is matched today only by that of the great white shark, all the more impressive given the piranha’s diminutive size (usually between 15 and 25 cm in length; Figure 1.1) compared to several metres of shark. Unlike the shark, piranhas usually attack prey larger than themselves.

Figure 1.1 A 19-cm long piranha showing the general body features. Note the deep-bodied outline, the forward-jutting lower jaw and the wide gape of the jaws.

The piranha’s aggressive image has proved highly marketable, giving its name to, among other things, a thirst-slaying, fizzy drink with its ‘bone-crushing citrus’ (Figure 1.2); a U.S. women’s ice hockey team, the ‘Pittsburgh Piranhas’; and two well-known rock bands.

Figure 1.2 A can containing a fizzy drink called Piranha with the logo ‘bone-crushing citrus’.

Piranhas have also inspired Hollywood. In the 1978 cult horror film Piranha, a shoal of piranhas, genetically engineered by the military to be oversized, fast-breeding and flesh-crazy, is accidentally released into a river leading to a holiday resort, with predictably gory results. In the 2010, 3-D film Piranha, a shoal of piranhas thought to be extinct for over 2 million years is released from an underground lake into a river following an earthquake. They reach a town where a large number of teenagers, swimming in the river, meet with a violent end. In the James Bond film You Only Live Twice, supervillain Ernst Stavro Blofeld feeds one of his female agents to his pet piranhas as a punishment for failing to kill Bond.

Much ferocious piranha behaviour has been wildly exaggerated. Of the more than 30 different species, many do not kill other fish, but merely ‘graze’ off them, taking small pieces of the fin or scales, which will later regenerate. Some are vegetarians. Only a few species are considered fierce and aggressive. Attacks on humans are extremely rare, and deaths are virtually unknown. Amazonian children swim quite safely in streams and rivers inhabited by piranhas, as has the occasional intrepid, documentary filmmaker. A recent exception occurred in 2002 in southeast Brazil, where over 50 attacks were recorded (the most serious requiring a toe amputation). This unusual aggression seemed to be a consequence of damming the local river for flood protection, resulting in a large number of understandably irritable piranhas being trapped in the still waters, compared with the flowing white waters of their native environment.

The giant feeding frenzy is also a myth: some piranhas hunt individually, whereas shoals typically contain only 20–30 individuals. Piranhas tend to pair up when swimming in shoals, and if there is an odd number of fish, the unlucky unpaired single one may be eaten by its fellows. Although previously regarded as an attacking strategy, recent research has suggested that shoaling behaviour may also represent a defence against predators (i.e. safety in numbers).

Piranhas are bony fish belonging to the genus Serrasalmus, meaning saw salmon and referring not to the teeth but to the notched appearance of the enlarged scales running in the midline across the belly. The origin of the word piranha is uncertain but may be derived from a local Amazonian tribal word meaning either toothed fish or scissors. In support of the latter, piranha jaws are used as cutting or shearing tools, even being employed to cut hair or to sharpen poisonous darts.

Although their main prey is other fish, piranhas are opportunistic and will eat whatever is available. Analysis of their stomach contents reveals insects, crustaceans, molluscs, lizards and rodents. As the rivers flood into the forests, piranhas incorporate seeds and other vegetable matter into their diet. Like carnivorous predators on the African savannah, piranhas seek out and dispatch the weak and injured animals in Amazonian rivers. Their extreme sensitivity to the scent of blood helps them in this pursuit. Birds, such as young herons and egrets, falling from their nests into the river below supplement the piranha diet.

Piranhas themselves have many predators such as larger fish, birds, caiman, freshwater dolphins, turtles and the indigenous people. A new tourist attraction of piranha fishing is being promoted. In the dry season, piranhas may become concentrated in isolated, shrinking, shallow pools that provide particularly rich pickings for fish-eating birds (including herons).

Teeth of the Piranha

The success of piranhas in obtaining food depends not only upon cooperating as a pack, like wolves, but also in their possession of an extremely specialised set of teeth. The unique arrangement of their teeth makes the piranha very effective as a predator. Whereas the majority of fish have large numbers of teeth (in many instances reaching a hundred or more, as in the barracuda; see Figure 4.10), the piranha has an upper row of only six and a lower row of seven on each side, and this number is retained throughout their life. Unlike most fish, whose teeth are conical and spaced out (Figure 1.3), piranha teeth are sharp, triangular, narrow and bladelike with razor-sharp edges and are in contact with each other (Figures 1.4 and 1.5). In addition to the central, prominent, main cusp, there is a small cusp at the back of each tooth, which slots neatly into a depression at the front of the tooth behind. This results in the teeth being totally interlocked (Figures 1.5 and 1.6). Instead of a number of separate teeth in each jaw, each piranha jaw could be considered as containing a single, continuous, serrated cutting edge.

Figure 1.3 The teeth of the short-nosed gar fish. Note the large number of spaced, conical teeth, with gaps along the dentition where replacing teeth will soon erupt.

Source: Courtesy of the Hunterian Museum at the Royal College of Surgeons.

Figure 1.4 View of the mouth of the piranha from the front, showing the razor-sharp pointed teeth.

Figure 1.5 The complete upper and lower teeth seen from the left side of the piranha. Note there are six teeth in the upper jaw and seven in the lower. The enlarged upper sixth tooth is arrowed.

Source: From R.P. Shellis and B.K.B. Berkovitz, 1976. Courtesy of the Journal of Zoology.

Figure 1.6 High-power view of Figure 1.5 showing the interlocking of the back four teeth in the lower jaw of the piranha. The teeth are attached to the jaw by a thin, dense fibrous ligament (arrows). The teeth lack roots.

Source: From R.P. Shellis and B.K.B. Berkovitz, 1976. Courtesy of the Journal of Zoology.

In contrast to mammals, whose teeth are secured by roots embedded in bony sockets in the jaws, fish teeth consist only of crowns that are attached to the surface of the bones of the jaw. In the piranha, the bases of the teeth are fixed to the crest of the jaws by a short, inflexible, strong, fibrous ligament. Together with the interlocking of the teeth and the saddle shape of the bone to which the teeth are attached, this provides great firmness and stability (Figure 1.6).

In other species of fish, such as the short-nosed gar, whose teeth are typically conical and spaced apart (Figure 1.3), the upper and lower teeth do not meet, and they are unable to cut up food into smaller, manageable pieces. The teeth function merely to grip the prey, which is usually swallowed whole and head first. A shark can rip chunks of flesh from its prey, but this is achieved more as a result of using twisting movements of its body once the prey has been grasped between its sharp teeth.

The piranha has a characteristic, forward-jutting, powerful lower jaw and is deep-bodied. It has a large gape (the distance between the jaws when the mouth is fully open) relative to its total body length (Figure 1.1); a piranha 10 cm long has a 1 cm gape. Together with its strong jaw muscles, this wide gape helps to generate a very large force when biting. As the mouth is snapped shut, the tips of the lower teeth, which slope increasingly backwards towards the rear of the jaw (Figure 1.5), puncture the prey, and the puncture points are elongated into cuts by the razor-sharp edges of the teeth moving upwards. The back four lower teeth slice against the large, back (sixth) upper tooth, which is greatly expanded in length to form an elongated blade (Figure 1.5), like a pair of scissors. A single bite from a piranha is sufficient to sever a portion of flesh from its prey. This slicing ability, where the teeth shear past each other, is unique among fish.

Figure 1.7 shows the unfortunate result of a piranha bite to a human hand. The individual concerned was transferring a piranha from one tank to another when the fish wriggled free from its net and landed on the floor. Without thinking, the person used their bare hand to push the fish back into the net. The piranha struck instantly, neatly excising a small, circular piece of skin the size of a five-penny piece, with resulting profuse bleeding. When the victim arrived at Accident and Emergency, the doctor, on hearing the cause of the injury, was far from being sympathetic and had difficulty keeping a straight face.

Figure 1.7 The palm of the hand of an unwary aquarist bitten by a piranha. The diameter of the wound was about 1 cm.

Tooth Replacement in the Piranha

In most fish (as well as in amphibians and reptiles), the teeth are constantly being replaced throughout life (see Chapter 9); as it may take a few weeks for a new tooth to erupt, temporary gaps are seen along the tooth row (Figure 1.3). In observing the skulls of piranhas, however, gaps are never seen. Invariably, the piranha seems to possess a complete set of teeth, despite the fact that another set of replacing teeth is always present within the jaws beneath the functioning teeth.

The replacing teeth in any one jaw are all at the same stage of development, although the precise stage of development will vary between jaws (Figures 1.8 and 1.9). Clearly, there must be tooth replacement, and yet no one had ever reported a situation where teeth were missing in the functional tooth row. The only possible way to explain these findings was to suggest that the teeth are shed simultaneously in complete rows and replaced so rapidly that a jaw without teeth would be rarely encountered in a museum collection. Due to the close interlocking of the teeth, it would be impossible for a tooth in the middle of the row to drop out on its own, as it was mechanically linked to its neighbouring teeth.

Figure 1.8 An X-ray from one side of a piranha lower jaw, showing the seven functional teeth, beneath which is situated a complete replacing set in which all the teeth are at the same stage of development.

Source: From R.P. Shellis and B.K.B. Berkovitz, 1976. Courtesy of the Journal of Zoology.

Figure 1.9 Tracings from X-rays of six different lower jaws representing a complete cycle of tooth replacement and illustrating the development, overlapping, eruption and tilting of the teeth necessary to establish the interlocked dentition.

Stage 1. With a new set of teeth newly erupted and attached to newly formed bone beneath them, the replacing teeth lying below them are at an early stage of development with just the tips of the teeth mineralised and visible on an X-ray. Note that all the replacing teeth are at the same stage of development.

Stage 2. Mineralisation is more advanced in the developing replacement teeth and has spread downwards from the tips of the cusp to involve the subsidiary cusps at the base of the crown. The two sets of teeth are separated by a reasonable thickness of bone.

Stage 3. The replacing teeth now become tilted in a backward direction so that their open bases face forwards. The sixth and seventh teeth now lie behind the last functional tooth. It is clear that the underlying replacing teeth are erupting upwards as the bone separating them from the functioning teeth above is getting thinner (as it is eaten away, resorbed, by special cells called osteoclasts) and the two sets of teeth are closer together.

Stage 4. The functional teeth have been shed and the bone to which they were attached has been completely resorbed. The replacing teeth have erupted further upwards (as indicated by the increased distance between them and the base of the bony socket in which they were housed). In this stage, the teeth are still tilted backwards.

Stage 5. The final phase in the cycle involves the teeth erupting forwards and rotating into the upright position, thereby ensuring that they interlock correctly.

Stage 6. The teeth then become attached to new bone forming immediately beneath them. A new set of replacing teeth will develop (Stage 1) and the whole cycle is repeated.

Source: From B.K.B. Berkovitz and R.P. Shellis, 1978. Courtesy of the Journal of Zoology.

The only way to test the hypothesis of extremely rapid replacement was to study live piranhas in an aquarium and to examine their teeth continuously, every few days, over a period of about a year. If caught in a net, a piranha will soon become still and allow its jaws to be gently prised open with a blunt probe, enabling its teeth to be observed and counted (Figure 1.10). The first important observation from this study confirmed the hypothesis that all the teeth in any one row are shed simultaneously. Such a pattern of tooth replacement is unique to the piranha. The second important observation was that the replacing teeth erupt into the mouth remarkably quickly, within just a few days. This very rapid rate of tooth replacement and eruption explains why no museum specimens had previously been found with missing teeth. In the piranha, replacing teeth erupt vertically from below. Although sharks and rays also tend to lose their teeth in rows, their teeth are not interlocked and their replacing teeth tend to move up from behind, like an escalator.

Figure 1.10 The right side of a live piranha, with the teeth exposed by a blunt, flat probe. All the teeth are present in the lower jaw but have just been shed in the upper jaw.

Source: From B.K.B. Berkovitz and R.P. Shellis, 1978. Courtesy of the Journal of Zoology.

The next question to be considered is whether the order in which the rows of teeth fall out and are replaced is the same for all piranhas. You might think that the most efficient pattern of replacement would be for all the teeth on one side of the jaw to be replaced first, followed later by those on the other side. This would allow the piranha the use of the teeth on one side while those on the other side were being replaced. Indeed, fish with this pattern of replacement were encountered and were seen able to feed. However, many other different patterns of tooth replacement were encountered. One piranha had the upper-right and lower-left tooth rows replaced simultaneously followed later by the lower right and upper left, while in another fish all the teeth fell out at the same time. When meat chunks were dropped into the tank of this particular temporarily toothless fish, it initially attacked the food in its usual manner, oblivious to the fact that it had no teeth. After a few attempts at trying to cut up the food into smaller, more manageable portions, it soon gave up and retreated to a corner of the tank to await the eruption of its new teeth (seemingly with a resigned look on its face!).

The fish whose teeth had all dropped out at the same time might be thought of as disadvantaged (at least temporarily) in the quest to obtain food, as for a short period of time it was completely without teeth. However, the very existence of such a pattern implies that it cannot be a serious disadvantage, as natural selection would have eradicated such a pattern over time. Just as with other carnivorous fish (such as the pike) and many animals, such as snakes, a few days without food would not be a problem for a piranha.

Piranhas grow throughout life, yet the number of their teeth remains the same. To keep pace with jaw growth, the size of the teeth increases very slightly with each new replacement. The number of tooth replacements that occur during the life of a piranha can be estimated using a mathematical formula (known as a Strasbourg plot). For this, it is necessary to measure the length of a tooth (for convenience in measuring teeth from X-rays, the third tooth along the row in the lower jaw) and that of the slightly wider replacing tooth beneath it. These measurements are then related to the length of the jaws in a large series of piranhas, ranging from very young to very old. Such data indicate that a piranha will have had up to 30 sets of teeth in its lifetime. Also with time, the speed of replacement slows down: as the teeth get larger, they last longer. For example, in one medium-sized piranha, the teeth last for about 100 days before being replaced, but 4 years later in the same fish they are replaced after 120 days. Generally, the teeth show little signs of wear at the time they are shed, so that tooth replacement is more related to the increasing size of the jaws than tooth wear.

Is it possible that for some geriatric piranhas, tooth replacement may eventually cease altogether? The probable answer is yes, as one isolated, large, lower jaw has been examined in which the normally sharp teeth were heavily worn down on one side of the jaw, a feature not previously observed. Furthermore, X-rays showed there was no evidence of replacement teeth waiting to erupt beneath these old, worn teeth (Figure 1.11). For this side of the lower jaw at least, tooth replacement had come to an end. On the opposite side of the jaw there were replacing teeth, and the functional teeth had not been worn