Ornithopoda

Ornithopods include heterodontosaurs, known from southern Africa; the slightly larger hypsilophodontids, about three metres in length; the much larger iguanodontids, about nine metres long, mostly from North America and Europe; and the large duck-billed hadrosaurs of North America and Eurasia. In all these forms, the front teeth are set slightly lower than the cheek teeth; the jaw joint is set lower than where the teeth meet in the jaws (the occlusal plane); and the nasal bone is excluded by a separate bone (the premaxilla) from contacting the upper jaw (maxilla).

The postcranial anatomy of the ornithopods reflects the bipedal ancestry of the group, but the giant hadrosaurs and some iguanodontids may have been as comfortable on four legs as on two, especially while feeding on low vegetation. All members had hind legs that were much longer and sturdier than their forelegs. The thighbone (femur) was nearly always shorter than the shinbones (tibia and fibula), especially in all but the largest forms, and it usually bore a prominent process, called the fourth trochanter, just above mid-length for the attachment of the retractor, or walking, muscles. The pelvis was expanded, usually with an elongated and broad blade of the ilium for the attachment of the protractor, or recovery, leg muscles. The pubis, as in all ornithischians, had migrated backward to lie parallel to the ischium, as described above; but in all but the most basal forms, a new prepubic process began to grow forward from the pubis, eventually reaching far in front of the forward edge of the ilium and becoming expanded into a paddlelike shape in hadrosaurs. It is generally thought that this process supported abdominal muscles and connective tissues of internal organs, but little is demonstrably known. The tail was long and sometimes quite deep and flat-sided. The vertebral spines of the tail and trunk region were reinforced by a rhomboidal latticework of bony (ossified) tendons running in criss-cross fashion between adjacent spines. They suggest a certain degree of stiffening of the tail and backbone, which were balanced over the massive hips.

Ornithopod feet were modified from the primitive five-toed pattern in a way that resembled similar modifications in theropod feet. The three middle toes served as the functional foot; the inside toe was shortened and often held off the ground, and the outside toe was greatly reduced or absent altogether. The resemblance to theropod feet is so strong that the footprints of the two groups are easily confused, especially if poorly preserved. The toes of all but the most basal ornithopods terminated in broad, almost hooflike bones, especially in the duckbills, as opposed to the sharp claws of theropods, and this is one way to distinguish their footprints. The hand reflected the primitive five-digit design, and, as was generally true in archosaurs, the fourth and fifth digits were shorter than the other three, with the third being longest. In iguanodontids and hadrosaurs, the fingers ended in broad, blunt bones rather than in claws, much like the toes. It is thought that these middle fingers and toes were covered by blunt, hooflike structures. In the duckbills the fingers apparently were encased in a mittenlike structure that could have broadened the hand for better support of the animal’s weight on soft ground.

The Ornithopoda differ from one another mainly in the structure of their skulls, their jaws and teeth, their hands and feet, and their pelvises. Ornithopods constitute an excellent case study in evolution because, as the various lineages arise and die out from the latest Triassic to the latest Cretaceous, trends in size, complications and elaborations of teeth and chewing mechanisms, adaptations for quadrupedal posture in some forms, and other changes emerge clearly from their phylogenetic patterns.

In the fabrosaurids the teeth were simple leaf-shaped, laterally compressed elements arranged in a single front-to-back row in each jaw. They were not set in from the outer cheek surface as in most ornithopods. Small incisor-like teeth were borne on the premaxillary bones above, but (as always) no teeth were present on the predentary below. One pair of incisors had been lost. The lower jaw had no coronoid process for large muscle attachment, and the upper temporal opening (the jaw muscle site), like the mandibular opening, was relatively smaller than in theropods and other archosaurs. Upper and lower teeth alternated in position when the jaw was closed; they did not occlude directly.

In heterodontosaurs the cheek teeth were crowded together into long rows and set inward slightly from the outer cheek surface. The inset, which persisted through all later ornithopods, has been interpreted to suggest the presence of cheeks that may have held plant food in the mouth for further processing by the cheek teeth. They occluded directly to form distinct chisel-like cutting edges with a self-sharpening mechanism maintained by hard enamel on the outer side of the upper teeth and the inner side of the lower. There were prominent upper and lower tusklike teeth at the front of the mouth (the upper set in the premaxillary bones, the lower on the dentary bones). At least two pairs of incisors seem to have been retained. Certain features of the skull suggest much larger jaw muscles in heterodontosaurs than in the fabrosaurids.

The hypsilophodonts had cheek teeth arranged in tightly packed rows set well inward from the outer cheek surfaces. The teeth occluded directly, and the opposing rows formed a long shearing edge similar to that of the heterodontosaurs. There was, however, no “tusk” either above or below. The premaxillaries had small simple incisor-like teeth above the beak-covered, toothless predentary. Strong projections of bone extended up from the lower jaw toward the moderate-size upper temporal fenestrae.

The skulls of iguanodonts accommodated still larger jaw muscles, but the cheek teeth were less regular and compacted than in the primitive ornithopods and consequently did not occlude as uniformly. Both the premaxillaries and the predentary were toothless but probably were sheathed in horny beaks.

Specialization of the teeth and jaws reached a pinnacle in the hadrosaurs, or duck-billed ornithopods. In this group a very prominent, robust projection jutted from the back of the stout lower jaw. Large chambers housing muscles were present above this process and beneath certain openings in the skull (the lateral and upper temporal fenestrae). These chambers are clear evidence of powerful jaw muscles. The dentition consisted of numerous tightly compacted teeth crowded into large grinding batteries. The battery in each jaw was composed of as many as 200 functional and replacement teeth with distinct, well-defined wear, or grinding, surfaces that resulted from very exact occlusion. As teeth were lost from the front of the jaws in iguanodontids and hadrosaurs, the snouts expanded into a bulbous shape, especially in the “duck-billed” hadrosaur, and may have been covered by a horny beak that improved feeding. These bills apparently had edges sharp enough to shred and strip leaves or needles from low shrubs and branches. Pine needles have been identified in duck-billed dinosaur remains and presumably represent stomach contents.

Other interesting specializations may have assisted iguanodontids and hadrosaurs in feeding. In both groups there was a marked increase in mobility (kinesis) among the joints of the bones of the facial region. As the jaws clamped down, some cheek bones were allowed to rotate outward slightly, perhaps to cushion the stress of chewing tough foods. The hands were also unusually modified in the two groups, though in different ways. In iguanodontids the wrist bones were coalesced into a single blocky structure that was less mobile than in more primitive wrist configurations. The joints of the thumb were similarly coalesced into a single conelike spike that had limited mobility on the wrist. The middle three digits flexed in the normal way and bore broad flat, spatulate claws. The fifth digit actually had two additional joints and became somewhat opposable to the rest of the hand. It is thought that the hands may have been adapted to grasp and strip vegetation, and the spikelike thumb has been suggested to have been an effective weapon against predators. These features were more or less continued in hadrosaurs, except in this group the blocky wrist was reduced and the thumb was lost completely.

Some varieties of hadrosaurs are also noted for the peculiar crests and projections on the top of the head. These structures were expansions of the skull composed almost entirely of the nasal bones. In genera such as Corythosaurus, Lambeosaurus, Parasaurolophus (and a few others), the crests were hollow, containing a series of middle and outer chambers that formed a convoluted passage from the nostrils to the trachea. Except for passing air along to the lungs, the function of these crests is not widely agreed upon. Sound production (honking), an improved sense of smell, and a visually conspicuous ornament for species recognition are some suggestions. Because these animals are no longer considered to have been amphibious, ideas such as snorkeling and extra air storage space have generally been discarded. Besides, the crests had no opening at their ends and consequently would not have been able to work as snorkels; even the largest crests held only an estimated 2 percent of the volume of the lungs, hardly enough to justify the construction of such an elaborate structure.