Podokesaurus

Source: Wikipedia, the free encyclopedia.

Podokesaurus
Temporal range:
Ma
↓
Holotype specimen
, with tail and uncertain bones at left, and body at right
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Superfamily: †Coelophysoidea
Genus: †Podokesaurus
Talbot, 1911
Species:
†P. holyokensis
Binomial name
†Podokesaurus holyokensis
Talbot, 1911
Synonyms
  • Coelophysis holyokensis (Talbot, 1911)
    Colbert
    , 1964

Podokesaurus is a

state dinosaur
of Massachusetts in 2022.

Estimated to have been about 1 m (3 ft) in length and 1–40 kg (2–90 lb) in weight, Podokesaurus was lightly constructed with hollow bones, and would have been similar to Coelophysis, being slender, long-necked, and with sharp, recurved teeth. The vertebrae were very light and hollow, and some were slightly concave at each end. The cervical (neck) vertebrae were relatively large in length and diameter compared to the dorsal (back) vertebrae, and the caudal (tail) vertebrae were long and slender. The humerus (upper-arm bone) was small and delicate, less than half the length of the femur (thigh-bone). The pubis (pubic bone) was very long, expanding both at the front and hind ends. The femur was slender, nearly straight, had thin walls, and was expanded at the back side of its lower end. The three metatarsals of the lower leg were closely appressed together forming a compact structure.

Since it was one of the few small

proto-bird. It was suggested it was a synonym of Coelophysis and a natural cast specimen was assigned to it, but these ideas are not currently accepted. The family Podokesauridae is not in use anymore, having been replaced by Coelophysidae, and Podokesaurus is thought to have been a coelophysoid. As such, Podokesaurus would have been a fleet-footed predator, with powerful forelimbs and grasping hands. It is estimated it could have run at 15–20 km/h (9–12 mph). Podokesaurus is thought to have been collected from the Portland Formation, the age of which has long been unclear, but is currently believed to date to the Hettangian–Sinemurian
stages of the Early Jurassic, between 201 and 190 million years ago.

Discovery and history

Connecticut Valley from 1915; 22 is where Podokesaurus was found, 12 is Mount Holyoke (some areas marked as Triassic are now thought to be Early Jurassic
in age)

In 1910, the American geologist

Connecticut Valley (that she had often taken her students to see), but few actual dinosaur skeletons, and none at Mount Holyoke. She was granted permission by the land owner to collect the specimen for Mount Holyoke College (an all-women's college a few miles from there), where she was head of the geology department.[1][2][3][4]

The next day she brought a group of workmen to collect the specimen, and found another piece of sandstone that contained the rest of the fossil as well as impressions of those in the first slab. The specimen appeared to have been exposed for years with no one noticing it, the boulder having been broken open by people or frost. The fossil was brought to the laboratory where it was prepared and photographed.[1] The incomplete specimen preserved 5 cervical (neck), 11 dorsal (back) and 24 caudal (tail) vertebrae, a fragment of the left scapula (shoulder blade) and right coracoid (part of the pectoral girdle), a partial left humerus (upper arm bone), phalanx bones of 3 fingers, including 2 unguals (claw bones), ribs, the pubis and ischium (bones forming the lower front and lower back of the pelvis, respectively), the femora (thigh bones), the left tibia (lower leg bone), a fragment of the right astragalus (a bone in the ankle), articulated metatarsals (foot bones) of the left foot, followed by 3 pedal phalanges (toe bones), partial metatarsals and possible phalanges of the right foot, and fragments of uncertain identity.[5][6] The light and delicate bones were in their natural position or nearly so within the rock, except for the tail and uncertain fragments, which were a few centimeters away from the skeleton, following a long gap in the skeleton behind the pelvis. The front half of the neck was missing.[1][7]

1915 skeletal reconstruction by Richard Swann Lull; shaded areas were known from the fossil, missing parts were reconstructed after Compsognathus
Model by Lull at Peabody Museum of Natural History

The significance of the fossil was confirmed at an intercollegiate meeting of geology departments, and when the American paleontologist

scientific description, in which she made the specimen the holotype of the new genus and species Podokesaurus holyokensis.[1][8] The generic name is derived from the Ancient Greek words podōkēs (ποδώκης), which means "swift (or fleet)-footed", an epithet commonly used in reference to the Greek hero Achilles, and saura (σαύρα) meaning "lizard", while the specific name refers to Holyoke. In full, the name can be translated as "swift-footed lizard of Holyoke".[8][2] The discovery and naming of Podokesaurus made Talbot the first woman to find and describe a non-bird dinosaur.[9] The American paleontologist Robert T. Bakker stated in 2014 that while old professors grumbled that women were unfit for working with fossils during his time at university, Talbot's discovery of Podokesaurus was a counterargument to that.[10]

By the time the description was published, Talbot had sent the fossil to the

Longmeadow Sandstone.[6][2][11][7]

Mignon Talbot (second from left) and students searching the rubble of the burned down Williston Hall in 1917

Lull had sent his manuscript to the Danish ornithologist Gerhard Heilmann for criticism prior to publication. Heilmann published his response in a 1913 article in which he included previously unpublished photos of the fossil received from Talbot, as well as his own restorations. He disagreed with some of Lull's anatomical interpretations, and had corresponded regarding the fossil with the American zoologist Robert Wilson Shufeldt about his contentions. Heilmann's article was one in a series about the origin of birds, wherein he examined the skeletons of prehistoric reptiles to find traits that may have been ancestral to birds.[12] In a 1916 Washington Academy of Sciences meeting, Shufeldt gave an account of his correspondence with Lull, Talbot, and Heilmann, and agreed with the latter in some of his criticisms of Lull's restoration of Podokesaurus.[13]

Talbot wanted the fossil to stay at Yale or Washington on permanent exhibit, where it could "be with its kind", but it was kept at Mount Holyoke in the old science building, Williston Hall, as a local specimen, where it became a "pet curiosity" for the students. During the Christmas break of 1917, Williston Hall burned down, and no remains of the Podokesaurus fossil were found in the rubble. The American writer Christopher Benfey pointed out in 2002 that Podokesaurus therefore had the peculiar distinction of being the dinosaur that vanished twice.[2][1] The German paleontologist Friedrich von Huene was the last to study and describe the holotype specimen, in a 1914 publication before it was destroyed (he had studied it at Yale before Talbot's description was published).[7][14] While the college's fossil collections were almost entirely destroyed by the fire, its facilities and collections continued to grow and improve afterwards due to Talbot's efforts.[15]

No other unequivocal Podokesaurus specimens have since been found, but cast replicas of the type specimen remain at the Peabody Museum of Natural History and the

sauropodomorph Anchisaurus, Podokesaurus was among the first substantial collections of dinosaur skeletal material known from the East Coast of North America. Apart from a tooth (assigned to Coelophysis in 1976), all other Early Jurassic theropod records on the East Coast are footprints.[16][17][18]

In 2021, Podokesaurus and Anchisaurus were proposed as contenders for

state dinosaur of Massachusetts by Representative Jack Patrick Lewis, with Podokesaurus winning the online poll.[19][20][21] On May 2, 2022, Governor Charlie Baker signed a law making it the official state dinosaur.[22]

Description

Size of Podokesaurus compared to a human and a cat

The preserved body of the only known Podokesaurus fossil was 18 cm (7 in) long.

coelophysoid, it would likely have been similar in build to for example Coelophysis, being slender, long-necked, and with sharp, recurved teeth.[16]

Since few other small theropods were known at the time Podokesaurus was described, it was mainly compared to the later genera Compsognathus,

bilaterally symmetric, and one was broadly convex with a sulcus (or furrow) at the midline, and according to Lull, was possibly placed at the upper wall of the skull, not far from the frontal bones. Lull thought other of these elements may have been part of the nasal bones.[6][17] Von Huene instead interpreted these bones as caudals from the middle of the tail.[7]

Postcranial skeleton

Photo of a partial skeleton
Drawing of the skeleton
Photo and interpretative drawing of the front part of the specimen from 1911 (above), photo of uncertain bone fragments that may belong to either the skull or tail (bottom left), drawing of the tail and uncertain bone fragments (lower middle), and cross-section of a vertebra (lower right)

The vertebrae were very light and hollow, and some were slightly concave at each end. The cervical vertebrae were relatively large in length and diameter compared to the dorsals, and had styliform (resembling a pen or bristle in shape) cervical ribs which were long, straight, and narrow, as in Anchisaurus. The front dorsals were much shorter than the cervicals, then again increased in size towards the pelvis. They measured 12–13 mm (0.47–0.51 in) in length. The neural spines of the dorsals were narrow, high, and curved backwards, but not as expanded from front to back as in Compsognathus. The dorsal ribs were strongly curved and very slender and hollow, reminiscent of those in Saltopus and Hallopus, the largest being 52 mm (2.0 in) long and 2 mm (0.1 in) wide. The transverse processes (that connected with the ribs on each side of a vertebra) formed a circular arc downward from each side of a dorsal, extending about as far from the centrum (the "body" of a vertebra) as the spinous process was high. The zygapophyses (processes that connected adjacent vertebrae) were well-developed. The centra were relatively longer than those of Ornithomimus, while those of Compsognathus resembled those of Podokesaurus more, but were shorter.[8][6][7]

While the hind part of the tail was located some distance away from the rest of the skeleton, Lull believed it was in its natural position and that it would have been very long, the intermediate vertebrae having been swept away. The caudal vertebrae were very long and slender, and along with their

sternal ribs.[8][6][7]

The humerus was small and very delicate, 42 mm (1.7 in) long, less than half the length of the femur. It had a strong processus lateralis (a process directed to the side) 30 mm (1.2 in) above its lower end. Lull found it similar to that of Ornitholestes, apart from the radial crest (that connected with the radius bone of the lower arm) not being as high, showing little muscular power. Von Huene estimated the radius to have been 30 mm (1.2 in) long. Lull found the hand more similar to that of Compsognathus than of Ornitholestes.[8][6][7] Only slender phalanx bones of the three fingers were preserved, including sharp, curved claws.[7]

life restoration

The pubis was very long, expanding both at the front and hind ends. It was 6 mm (0.24 in) wide and 95 mm (3.7 in) long, and its lower expansion was 10 mm (0.39 in) in diameter. Talbot found the pubis similar to that of the then undescribed Procompsognathus. Lull argued that the pubis lay approximately in position in the fossil, pointing forwards.[8][6] Heilmann instead believed the bone had been moved out of position before burial, and would have been directed backwards.[12] Colbert and Baird suggested the pubis was directed forwards, but that it curved slightly upwards instead of downwards due to natural warping and cracking of the bone. The ischium was 55 mm (2.2 in) long, 15.5 mm (0.61 in) wide at the end closest to the body, and its slender shaft was 4 mm (0.16 in) in diameter, and somewhat thickened at its lower end. Lull found it similar to those of Compsognathus and Ornitholestes. Below the acetabulum (where the femur connects with the pelvis) there was a broad lamella running towards the pubis.[6][7]

The femur was slender, nearly straight, had thin walls, and was expanded at the back side of its lower end. It was 86 mm (3.4 in) long and 6.5 mm (0.26 in) in diameter just below the

condyles (round prominences) of the femur's lower end protruded strongly hindwards. The tibia was a thin, narrow shaft, 104 mm (4.1 in) in length, and about 7 mm (0.28 in) in diameter. Talbot stated the very thin fibula lay close to the tibia and was of almost equal length, while von Huene thought this was instead part of the crushed tibia. The three metatarsals of the lower leg were closely appressed together forming a compact structure, similar to what was seen in Ornithomimus, but not fused into a tarsometatarsus, as seen in birds. The length of the metatarsals was 75 mm (3.0 in), while their width was 9 mm (0.35 in). The most complete pedal phalanx was very slender and 12 mm (0.47 in) long.[6][8][7]

Classification

Early interpretations

Lull's 1915 illustration of the pelvic region, showing the forwards directed pubis (Pu) as preserved

When reading her preliminary description in 1910, Talbot suggested Podokesaurus to have been an "herbivorous dinosaur", but further work at Yale University showed that some bones had been incorrectly identified, and the ischium of the pelvis with a well-developed ridge was found to resemble that of Compsognathus. She refrained from making a definite classification of the specimen in her 1911 description due to the lack of jaw and foot-bones which could have aided in this, but concluded it would have belonged to a "carnivorous dinosaur" based on the shape and position of the pubis, as well as the absence of a

family Podokesauridae, wherein he, in addition to Podokesaurus, included Saltopus, Procompsognathus, Coelophysis, and Tanystropheus. He placed this group under Coelurosauria, which at the time was used to include small theropods in general.[25][7][26]

Heilmann considered Podokesaurus very similar to the early bird

proto-bird, with not yet fused middle-foot bones. The only feature he found inconsistent with this interpretation was the short finger with a claw, if it was not instead part of a toe. He suggested that the issue could be determined if the skull, sternum, and clavicle were found in the rock, bones that would be important clues for classification.[12]

dinosaurs

Heilmann wished for a clearer explanation of the placement of the middle-foot bones in relation to each other, and found it surprising that Lull's reconstruction of Podokesaurus showed an even more appressed middle-foot than was present in the later supposed descendant Ornithomimus, which he thought would have developed a completely bird-like middle-foot by that time. Heilmann found Podokesaurus very unusual for a dinosaur from the Triassic, and thought it was one of the earliest bipedal vertebrates. He suggested that since it was so different from

parasuchians, but found its fossils too insufficient to say anything definite about the issue.[12]

Shufeldt elaborated on his and Heilmann's interpretation of the pubis in 1915, and stated the bone had probably been displaced during fossilization. He pointed out that if it had faced forward, it would have been in forcible contact with the abdominal ribs that would have been dangerous for internal organs during movement.

ornithosuchian pseudosuchians rather than from dinosaurs, due to their bipedality and bird-like legs.[27]

The paleontologist

Procerosaurus, Saltopus, and Tanystropheus.[29] Von Huene grouped Podokesaurus, Coelophysis, Spinosuchus, Halticosaurus, Saltopus, Avipes, and Velocipes in Podokesauridae in 1932.[30]

Relation to Coelophysis

Replica of a natural cast of bones assigned to C. holyokensis in 1964 (when Podokesaurus was suggested to be a synonym of Coelophysis)

In 1958, Colbert and Baird described a theropod specimen consisting of natural casts of bones in sandstone (probably formed when the bones were dissolved by acidic water, leaving molds of the bones), including a pubis, tibia, and some ribs. They found the specimen similar to Coelophysis and Podokesaurus, referring to it as C. sp. (of unknown species). They considered the natural cast and Podokesaurus to be from the

new combination C. holyokensis. He also suggested that the natural cast belonged to C. holyokensis. Colbert stated that Podokesaurus could only be distinguished from Coelophysis by the neural spines of its vertebrae not being as long from front to back, and because the ischium differed in shape. He found these differences to be similar to those seen between modern reptile species within the same genus, and that they represented eastern (C. holyokensis) and western (C. bauri) species of the same genus. While he admitted that these conclusions were not ironclad, and that Podokesaurus may indeed have been distinct, he said the burden of proof should be on the proponents of such a view.[17]

In 1977, the paleontologists Paul E. Olsen and

Syntarsus, Olsen suggested that Podokesaurus should be retained as a separate genus of indeterminate theropods, to avoid overextending the temporal range of Coelophysis. He also applied this argument to the natural cast, regarding it as incertae sedis (with uncertain relationships) among theropods rather than as Podokesaurus or Coelophysis.[34]

The paleontologist

Samuel P. Welles stated in 1984 that the family Podokesauridae had become a "catch-all" for most Triassic theropods. While he found Coelophysis the most similar to Podokesaurus among theropods, he thought the two differed greatly in that the fourth trochanter of the latter was below midheight on the femur (unlike the higher position in most other theropods), and its metatarsals were of equal length.[14] The paleontologist Kevin Padian stated in 1986 that while Colbert's suggestion of synonymy was possible, the discernible similarities between Podokesaurus and Coelophysis were primitive theropod features, and the two were not as close in time as once thought.[35] Paul said in 1988 that while the family Podokesauridae was still used for Coelophysis and kin, Podokesaurus was not based on good remains to begin with, and with only bad cast replicas remaining, he thought there would always be disagreement about the taxon. He found this to be too much ambiguity to put up with, and proposed the name Coelophysidae should be used instead (Halticosauridae, another contender, was also based on too fragmentary remains).[36] In 1989, Colbert also doubted the synonymy of Podokesaurus with Coelophysis, and that the natural casts belonged to the latter, due to their revised Early Jurassic ages, and he made no attempt to resolve the status of Podokesaurus due to the only specimen being lost. He retained the name Podokesauridae for the family.[37]

Mounted skeleton of Coelophysis, which is thought to have been similar to Podokesaurus

In 1990, the paleontologists Timothy Rowe and

ceratosaurian affinities for Podokesaurus in 1996.[40]

The paleontologist Ronald S. Tykoski and Rowe noted in 2004 that while Podokesaurus had coelophysoid features (such as a small, knob-like expansion on the lower part of the pubis), it did not have any

dissertation, but left Podokesaurus out of his analysis.[45] Paul suggested in 2016 that Podokesaurus may have formed the family Coelophysidae with Coelophysis, Panguraptor, and Procompsognathus.[24]

Paleobiology

Heilmann's outdated 1913 restoration of Podokesaurus running near a lake and other dinosaurs

Talbot suggested that the short, slender humerus, long, straight hindlimb bones, and the well-developed fourth trochanter of the femur indicated that Podokesaurus was bipedal. She found the fact that the tibia was much longer than the femur, that the metatarsals were very long, over half the length of the tibia, and the skeleton's light construction, were indicative of rapid locomotion. Talbot stated that Lull thought this was an adaptation to climatic conditions, as the animal must have been able to travel fast and far for water in its

semi-arid region. Talbot also reported a small piece of smooth, polished quartz among the ribs, and suggested it could have been a gastrolith (stomach stone), and so the first time these were found in association with a carnivorous dinosaur.[8] Von Huene agreed the stone was a gastrolith, distinct from the surrounding sandstone, and added it was 11 mm (0.43 in) long.[7]

Lull found the animal to have been essentially a slender, cursorial (adapted for running) animal, with carnivorous habits, but that the slenderness that made it swift also confined it to small prey.[6] In 1932, von Huene proposed that small coelurosaurs had a jumping gait, due to their lower legs being longer than their upper legs, contrasting with the alternating steps of carnosaurs. With its large fourth trocanther, he thought Podokesaurus had probably abandoned this jumping gait, instead moving with rapid, alternating steps similar to ratite birds.[30]

Grallator cursorius
 footprint, which Lull correlated with Podokesaurus

In 1982, the paleontologist

locomotor ability and physiology. He cautioned that comparison was difficult because dinosaur anatomy differed significantly from that of mammals and birds. Differences included the massive tails of dinosaurs, and hindlimb retractor muscles that originated behind the femur, while these muscles originate more forward in mammals and extant birds.[46] Weishampel stated in 2006 that as a coelophysoid, Podokesaurus would have been a fleet-footed predator, with powerful forelimbs and grasping hands.[16]

Lull found that fossil footprints named

Brunswick Formation of New York previously attributed to Coelophysis may instead have been produced by Podokesaurus, based on its Jurassic age.[49] This formation is now known as the Passaic Formation, and instead thought to be Late Triassic in age.[50]

Paleoenvironment

1917 restorations of animals from the Connecticut Valley then thought to have lived during the Triassic. The Portland Formation, which Podokesaurus is from, was redated to the Early Jurassic in 1977

The only known specimen of Podokesaurus is thought to have been collected from the Portland Formation (earlier known as the Portland Arkose) in the

geological group within the Newark Supergroup (as the Portland Group), and thereby replacing the former name "Agawam Group". They also reinstated the Longmeadow Sandstone, where Podokesaurus was found, as a formation (within the uppermost Portland Group); it had earlier been considered identical to the Portland Formation.[51][6]

The Portland Group represents the uppermost part of the Newark Supergroup, and was deposited after the

Anchisauripus, Anomoepus, Eubrontes, and Grallator.[52]

See also

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External links

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