Old Earth Ministries Online Dinosaur Curriculum
Free online curriculum for homeschools and private schools
From Old Earth Ministries (We Believe in an Old Earth...and God!)
NOTE: If you found this page through a search engine, please visit the intro page first.
Lesson 33- Dromaeosauridae (Raptors)
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'. In informal usage they are often called raptors (after Velociraptor), a term popularized by the film Jurassic Park; a few types include the term "raptor" directly in their name and have come to emphasize their supposed bird-like habits.
Dromaeosaurid fossils have been found in
North America, Europe, Africa, Japan, China, Mongolia, Madagascar,
Argentina, and Antarctica. They first
appeared in the mid-Jurassic Period 167
million years ago and survived until the end of the Cretaceous, 65.5 Ma, existing for over 100 million years, until the
Cretaceous–Tertiary extinction event. The presence of dromaeosaurs as
early as the Middle Jurassic has been confirmed by the discovery of isolated
fossil teeth, though no dromaeosaurid body fossils have been found from this
Date Range: 167 - 65 Ma
The following four lessons focus on individual Dromaeosaurid species:
|Deinonychus antirrhopus (large) and Buitreraptor gonzalezorum (small), Field Museum of Natural History (Picture Source)|
The distinctive dromaeosaurid body plan helped to rekindle theories that at least some dinosaurs may have been active, fast, and closely related to birds. Robert Bakker’s illustration for John Ostrom’s 1969 monograph, showing the dromaeosaurid Deinonychus in a fast run, is among the most influential paleontological reconstructions in history. The dromaeosaurid body plan includes a relatively large skull, serrated teeth, narrow snout, and forward-facing eyes which indicate some degree of binocular vision. Dromaeosaurids, like most other theropods, had a moderately long S-curved neck, and their trunk was relatively short and deep. Like other maniraptorans, they had long arms that could be folded against the body in some species, and relatively large hands with three long fingers (the middle finger being the longest and the first finger being the shortest) ending in large claws. The dromaeosaurid hip structure featured a characteristically large pubic boot projecting beneath the base of the tail. Dromaeosaurid feet bore a large, recurved claw on the second toe. Their tails were slender, with long, low, vertebrae lacking transverse process and neural spines after the 14th caudal vertebra.
It is now known that at least some, and probably all, dromaeosaurids were covered in feathers, including large, vaned, wing and tail feathers. This development, first hypothesized in the mid-late 1980s and confirmed by fossil discoveries in 1999, represents a significant change in the way dromaeosaurids have historically been depicted in art and film (see “Feathers” below).
Like other theropods, dromaeosaurids were bipedal; that is, they walked on their hind legs. However, whereas other theropods walked with three toes contacting the ground, fossilized footprint tracks confirm that most dromaeosaurids held the second toe off the ground in a hyperextended position, with only the third and fourth toes bearing the weight of the animal. This is called functional didactyly. The enlarged second toe bore an unusually large, curved sickle-shaped claw (held off the ground or 'retracted' when walking), which is thought to have been used in capturing prey and climbing trees (see "Claw function" below).
Dromaeosaurids had long tails. Most of the tail vertebrae bear bony, rod-like extensions, as well as bony tendons in some species. In his study of Deinonychus, Ostrom proposed that these features stiffened the tail so that it could only flex at the base, and the whole tail would then move as a single, rigid, lever. However, one well-preserved specimen of Velociraptor mongoliensis (IGM 100/986) has an articulated tail skeleton that is curved horizontally in a long S-shape. This suggests that, in life, the tail could bend from side to side with a substantial degree of flexibility. It has been proposed that this tail was used as a stabilizer and/or counterweight while running or in the air; in Microraptor, an elongate diamond-shaped fan of feathers is preserved on the end of the tail. This may have been used as an aerodynamic stabilizer and rudder during gliding and/or powered flight (see "Flight and gliding" below).
Mahakala is both the most primitive dromaeosaurid ever described and the smallest. This evidence, combined with the small size of other primitive relatives such as Microraptor and the troodontid Anchiornis, indicates that the common ancestor of dromaeosaurids, troodontids, and birds – which is called the ancestral paravian – may have been very small, at around 65 cm in length and 600 to 700 grams of mass.
There is a large body of evidence showing that dromaeosaurids were covered in feathers. Some dromaeosaurid fossils preserve long, pennaceous feathers on the hands and arms (remiges) and tail (rectrices), as well as shorter, down-like feathers covering the body. Other fossils, which do not preserve actual impressions of feathers, still preserve the associated bumps on the forearm bones where long wing feathers would have attached in life. Overall, this feather pattern looks very much like Archaeopteryx.
The first known dromaeosaur with definitive evidence of feathers was Sinornithosaurus, reported from China by Xu et al. in 1999. Many other dromaeosaurid fossils have been found with feathers covering their bodies, some with fully-developed feathered wings. Microraptor even shows evidence of a second pair of wings on the hind legs. While direct feather impressions are only possible in fine-grained sediments, some fossils found in coarser rocks show evidence of feathers by the presence of quill knobs, the attachment points for wing feathers possessed by some birds. The dromaeosaurids Rahonavis and Velociraptor have both been found with quill knobs, showing that these forms had feathers despite no impressions having been found. In light of this, it is most likely that even the larger ground-dwelling dromaeosaurids bore feathers, since even flightless birds today retain most of their plumage, and relatively large dromaeosaurids, like Velociraptor, are known to have retained pennaceous feathers. Though some scientists had suggested that the larger dromaeosaurids lost some or all of their insulatory covering, the discovery of feathers in Velociraptor specimens has been cited as evidence that all members of the family retained feathers.
In Popular Culture
Raptors have captured the imagination of people everywhere. They were a central focus of the Jurassic Park movie trilogy, and have been portrayed in numerous other movies and television shows. One novel, Raptor Red, by famed paleontologist Bob Bakker, portrays a year in the life of a female Utahraptor. It is available from Amazon.
--------------Optional Reading Below this Point---------------
There is currently disagreement about the function of the enlarged "sickle claw" on the second toe. When John Ostrom described it for Deinonychus in 1969, he interpreted the claw as a blade-like slashing weapon, much like the canines of some saber-toothed cats, used with powerful kicks to cut into prey. Adams (1987) suggested that the talon was used to disembowel large ceratopsian dinosaurs. The interpretation of the sickle claw as a killing weapon applied to all dromaeosaurids. However, Manning et al. argued that the claw instead served as a hook, reconstructing the keratinous sheath with an elliptical cross section, instead of the previously inferred inverted teardrop shape. In Manning's interpretation, the second toe claw would be used as a climbing aid when subduing bigger prey and also as stabbing weapon.
Ostrom compared Deinonychus to the ostrich and cassowary. He noted that the bird species can inflict serious injury with the large claw on the second toe. The cassowary has claws up to 125 millimetres (4.9 in) long. Ostrom cited Gilliard (1958) in saying that they can sever an arm or disembowel a man. Kofron (1999 and 2003) studied 241 documented cassowary attacks and found that one human and two dogs had been killed, but no evidence that cassowaries can disembowel or dismember other animals. Cassowaries use their claws to defend themselves, to attack threatening animals, and in agonistic displays such as the Bowed Threat Display. The seriema also has an enlarged second toe claw, and uses it to tear apart small prey items for swallowing.
Phillip Manning and colleagues (2009) attempted to test the function of the sickle claw and similarly shaped claws on the forelimbs. They analyzed the bio-mechanics of how stresses and strains would be distributed along the claws and into the limbs, using X-ray imaging to create a three dimensional contour map of a forelimb claw from Velociraptor. For comparison, they analyzed the construction of a claw from a modern predatory bird, the Eagle Owl. They found that, based on the way that stress was conducted along the claw, they were ideal for climbing. The scientists found that the sharpened tip of the claw was a puncturing and gripping instrument, while the curved and expanded claw base helped transfer stress loads evenly.
The Manning team also compared the curvature of the dromaeosarid "sickle claw" on the foot with curvature in modern birds and mammals. Previous studies had shown that the amount of curvature in a claw corresponded to what lifestyle the animal has: animals with strongly curved claws of a certain shape tend to be climbers, while straighter claws indicate ground-dwelling lifestyles. The sickle-claws of the dromaeosaurid Deinonychus have a curvature of 160 degrees, well within the range of climbing animals. The forelimb claws they studied also fell within the climbing range of curvature.
Paleontologist Peter Mackovicky commented on the Manning team's study, stating that small, primitive dromaeosaurids (such as Microraptor) were likely to have been tree-climbers, but that climbing did not explain why later, gigantic dromaeosaurids such as Achillobator retained highly curved claws when they were too large to have climbed trees. Mackovickey speculated that giant dromaeosaurids may have adapted the claw to be used exclusively for latching on to prey.
Deinonychus fossils have been uncovered in small groups near the remains of the herbivore Tenontosaurus, a larger ornithischian dinosaur. This had been interpreted as evidence that these dromaeosaurs hunted in coordinated packs like some modern mammals. However, not all paleontologists found the evidence conclusive, and a subsequent study published in 2007 by Roach and Brinkman suggests that the Deinonychus may have actually displayed a disorganized mobbing behavior. Modern diapsids, including birds and crocodiles (the closest relatives of dromaeosaurs), display minimal cooperative hunting; instead, they are usually either solitary hunters, or are drawn to previously-killed carcasses, where conflict often occurs between individuals of the same species. For example, in situations where groups of komodo dragons are eating together, the largest individuals eat first and might attack smaller komodo dragons that attempt to feed; if the smaller animal dies, it is usually cannibalized. When this information is applied to the sites containing putative pack-hunting behavior in dromaeosaurs, it appears somewhat consistent with a komodo- or crocodile-like feeding strategy. Deinonychus skeletal remains found at these sites are from subadults, with missing parts that may have been eaten by other Deinonychus, which a study by Roach et al. presented as evidence against the idea that the animals cooperated in the hunt.
In 2007, scientists described the first known extensive dromaeosaur trackway, in Shandong, China. In addition to confirming the hypothesis that the sickle-claw was held retracted off the ground, the trackway (made by a large, Achillobator-sized species) showed evidence of six individuals of about equal size moving together along a shoreline. The individuals were spaced about one meter apart, and retained the same direction of travel, walking at a fairly slow pace. The authors of the paper describing these footprints interpreted the trackways as evidence that some species of dromaeosaurs lived in groups. While the trackways clearly do not represent hunting behavior, the idea that groups of dromaeosaurs may have hunted together could not be ruled out.
Flying and gliding
The ability to fly or glide has been suggested for at least two dromaeosaurid species. The first, Rahonavis ostromi (originally classified as avian bird, but found to be a dromaeosaurid in later studies) may have been capable of powered flight, as indicated by its long forelimbs with evidence of quill knob attachments for long sturdy flight feathers. The forelimbs of Rahonavis were more powerfully built than Archaeopteryx, and show evidence that they bore strong ligament attachments necessary for flapping flight. Luis Chiappe concluded that, given these adaptations, Rahonavis could probably fly but would have been more clumsy in the air than modern birds.
Another species of dromaeosaurid, Microraptor gui, may have been capable of gliding using its well-developed wings on both the fore and hind limbs. A 2005 study by Sankar Chatterjee suggested that the wings of Microraptor functioned like a split-level "biplane", and that it likely employed a phugoid style of gliding, in which it would launch from a perch and swoop downward in a 'U' shaped curve, then lift again to land on another tree, with the tail and hind wings helping to control its position and speed. Chatterjee also found that Microraptor had the basic requirements to sustain level powered flight in addition to gliding.
Powered flight has also been suggested for the species Cryptovolans pauli (the name of which means "hidden flyer"), though Cryptovolans is probably synonymous with Microraptor.
Relationship with birds
Dromaeosaurids share many features with early birds (clade Avialae or Aves). The precise nature of their relationship to birds has undergone a great deal of study, and hypotheses about that relationship have changed as large amounts of new evidence became available. As late as 2001, Mark Norell and colleagues analyzed a large survey of coelurosaur fossils and produced the tentative result that dromaeosaurids were most closely related to birds, with troodontids as a more distant outgroup. They even suggested that Dromaeosauridae could be paraphyletic relative to Avialae. In 2002, Hwang and colleagues utilized the work of Norell et al., including new characters and better fossil evidence, to determine that birds (avialans) were better thought of as cousins to the dromaeosaurids and troodontids.
The current consensus among paleontologists agrees with the findings of Hwang et al. (2002); that dromaeosaurids are most closely related to the troodontids, and together with the troodontids form the clade Deinonychosauria. Deinonychosaurians in turn are the sister taxon to avialans, and therefore the closest relatives of avialan birds. A consensus of paleontologists has concluded that there is not yet enough evidence to determine whether any dromaeosaurs could fly or glide, or whether they evolved from ancestors that could.
Alternative Theories and Flightlessness
Dromaeosaurids are so birdlike that they have led some researchers to argue that they would be better classified as birds. First, since they had feathers, dromaeosaurs (along with many other coelurosaurian theropod dinosaurs) are “birds” under traditional definitions of the word “bird”, or “Aves”, that are based on the possession of feathers. However, other scientists, such as Lawrence Witmer, have argued that calling a theropod like Caudipteryx a bird because it has feathers may stretch the word past any useful meaning.
At least two schools of researchers have proposed that dromaeosaurs may actually be descended from flying ancestors. Hypotheses involving a flying ancestor for dromaeosaurs are sometimes called “Birds Came First” (BCF). George Olshevsky is usually credited as the first author of BCF. In his own work, Gregory S. Paul pointed out numerous features of the dromaeosaurid skeleton that he interpreted as evidence that the entire group had evolved from flying, dinosaurian, ancestors, perhaps something like Archaeopteryx. In that case, the larger dromaeosaurids were secondarily flightless, like the modern ostrich. In 1988, Paul suggested that dromaeosaurids may actually be more closely related to modern birds than to Archaeopteryx. By 2002, however, Paul placed dromaeosaurs and Archaeopteryx as the closest relatives to one another.
In 2002, Hwang et al. found that Microraptor was the most primitive dromaeosaur. Xu and colleagues in 2003 cited the basal position of Microraptor, along with feather and wing features, as evidence that the ancestral dromaeosaur could glide. In that case the larger dromaeosaurs would be secondarily terrestrial—having lost the ability to glide later in their evolutionary history.
Also in 2002, Steven Czerkas described Cryptovolans, though it is a probable junior synonym of Microraptor. He reconstructed the fossil inaccurately with only two wings and thus argued that dromaeosaurs were proper birds, rather than possible gliders. He later issued a revised reconstruction in agreement with that of Microraptor
Other researchers, like Larry Martin believe that dromaeosaurs, along with all maniraptorans are not dinosaurs at all. Martin asserted for decades that birds were unrelated to maniraptorans, but in 2004 he changed his position, and now he agrees that the two are the closest of relatives. Martin believes that maniraptorans are secondarily flightless birds, and that birds evolved from non–dinosaurian archosaurs, so that most of the species formerly called theropods would now not even be classified as dinosaurs.
In 2005, Mayr and Peters described the anatomy of a very well preserved specimen of Archaeopteryx, and determined that its anatomy was more like non-avian theropods than previously understood. Specifically, they found that Archaeopteryx had a primitive palatine, unreversed hallux, and hyper-extendable second toe. Their phylogenetic analysis produced the controversial result that Confuciusornis was closer to Microraptor than to Archaeopteryx, making the Avialae a paraphyletic taxon. They also suggested that the ancestral paravian was able to fly or glide, and that the dromaeosaurs and troodontids were secondarily flightless (or had lost the ability to glide). Corfe and Butler criticized this work on methodological grounds.
A challenge to all of these alternative scenarios came when Turner and colleagues in 2007 described a new dromaeosaurid, Mahakala, which they found to be the most basal and most primitive member of the Dromaeosauridae, more primitive than Microraptor. Mahakala had short arms and no ability to glide. Turner et al. also inferred that flight evolved only in the Avialae, and these two points suggested that the ancestral dromaeosaurid could not glide or fly. Based on this cladistic analysis, Mahakala suggests that the ancestral condition for dromaeosaurids is non-volant.
Return to the Old Earth Ministries Online Dinosaur Curriculum homepage.