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Vertebrate Term Paper

Vertebrates It is common knowledge that the human body consists of about 65% water. People cannot live any longer than five days without H20. Individuals of all ages love to sail the oceans, swim in the sea and soar under or speed across the waves. It comes as no surprise, then, that some part of the human psyche remembers millions and millions of years ago before animals came on shore. What is still questionable is how or why these animals made the move from water to land. The journal articles discussed below give some of the latest findings on this topic.

Early in the Devonian Era, close to 400 million years ago, all the continents were grouped closely together and surrounded by the seas. The climate ranged from dry weather to torrential rains as some tropical areas do today. Even flowers had not yet evolved on land, let alone vertebrates. Many of the sealife were preparing for that next big step onto land with lung-like organs that would later evolve into swim bladders to control buoyancy. Some of these creatures moved on lobed fins or fleshy appendages that supported their weight while crawling underground. In time, they adapted to terrestrial life and evolved into amphibians with fully developed legs.

In what kind of environment did the transition to lobed fin first occur? This has recently been a "bone" of contention. Marine biologists Graham and Lee understand that air-breathing fishes may be seen as possible models for the Paleozoic evolution of vertebrate air breathing and the transition to land. They note how recent studies suggest that marine air-breathing amphibious fish in tropical, high intertidal zone habitats are analogs of early tetrapods and that the intertidal zone are feasible early habitats for the Devonian land movement by vertebrates.

However, in response to such scientists, Graham and Lee argue that selection pressures imposed by life in these intertidal zones are insufficient to have led to the necessary respiratory capacity or break from water required for the vertebrates to move to land. The marine amphibious fishes, which occur mainly on rocky shores or mudflats, have reached what the authors call "their land-penetration" limits and remain linked to water by their respiratory structures that are less efficient in air and more vulnerable to desiccation than lungs. Such fish definitely cannot succeed in the biologically complex terrestrial environment that awaits them on the seashore. They are just too tied to the water, even as adults.

Because of the proximity of reefs and mudflats to dry land, as well as the exposure to tidal cycles and wave action, fish that live in these habitats will sometimes occasionally be exposed to air. It is not surprising then that intertidal fish tolerate air exposure. In fact, as scientists as Graham have seen previously, activities by these fish include laying eggs in the upper intertidal splash zone, riding waves to feed at the upper limits of an algal ridge, developing mud turrets or territorial boundary walls, and feeding with the moving tides.

Along with these activities come a variety of specializations regarding respiration, vision and terrestrial locomotion. However, even though they do spend some of their time breathing air, their link to the water has not decreased. For example, during high tide, one of these fish called the mudskipper revert back to water burrows to remoisten their skin and respiratory systems. Further, their reproductive systems are closely tied to the water. Besides serving as a refuge, the burrow serves as a nursery for developing eggs. Lee and Graham also stress that other physiological changes also weigh against these animals in terms of being able to move onto land. It is highly likely, they say, that earlier forms of intertidal marine life would have developed lungs.

Graham and Lee conclude, therefore, that "The unique combinations of changing habitat, exploitable land resources, and the breathing and locomotory capacities of the ancestral tetrapods drove selection for a group of organisms that could, in effect, come farther onto the shore. In the absence of extensive environmental change, this evolutionary tape cannot be replayed for the modern amphibious marine fishes."

How, then, did the first transition develop from water to land? Scientists know that sometime during the Devonian period the harsh sun caused severe droughts. Fish would have been trapped in drying pools and faced death. To survive, a few eusthenopterons must have dragged themselves on their fins, as mudskippers do today, out of the puddles in search of deeper water. Some could then have evolved on land. Their fins became legs, they grew five fingers and toes, they started...

They became tetrapods, the ancestors of all four-legged animals today. The fossil of this ancient ancestor, according to Erik Jarvik, is believed to be ichthyostega.
Ichthyostega was a relatively large, about 4 feet, with a stout body. Although it was originally considered to be the transitional form between fishes and Carboniferous amphibians, its skull possess several primitive, fish-like features. The spine of Ichthyostega is notochordal, instead of being based on a series of loosly jointed but interlocking vertebrae. It most probably did not have internal gills, its tail bore fish-like supraneural spines, and it had a massive ribcage with thoracic ribs that were long, flattened bones that overlapped with other ribs to protect the body from being injured. The large pelvis was attached to the spine, the hindlimbs were significantly smaller than the forelimbs and most likely functioned more like paddles than legs. The knee was flexible without evidence of an ankle joint. It had seven toes.

Since Jarvis' find, other scientists such as Jennie Clack have developed some ideas about similar early tetrapods that have been uncovered. She found, for example, another species of Devonian tetrapod called Acanthostega. Although different from Jarvik's fossil, it clearly shared the same ancestry. The forelimb of Acanthostega had eight digits instead of seven. Both came from the Upper Devonian of East Greenland, similar to the only other known Devonian tetrapod limb, Tulerpeton from Russia, which has six digits.

Clack and others such as Coates questioned if these tetrapods were meant to walk, why did they have different numbers of digits and why were they paddle shaped? The conclusion: The limbs may have been adaptations to an aquatic rather than a terrestrial environment. During this period of time, instead of walking on firm land, the tetrapods had to move around in swampy wet environments. The pattern of digits also altered the proposed model for limb development where digit number is unspecified, rather than earlier models that are rejected because they postulate a fixed number of elements in the ancestral limb. Clack and Coates thus challenged pentadactyly (five fingered) as primitive for tetrapods. The form of these limbs rather suggested early specialization in the evolution of the tetrapod limb bud.

Such findings led to a widely agreed-upon transition in which tetrapods diverged from their lobe-fin ancestors sometime during the Frasnian, and were broadly dispersed throughout tropical and subtropical localities by the end of the Devonian. Most importantly, the Devonian tetrapods appear to have been mostly if not exclusively aquatic. In other words, they were like fishes with legs.

Although Clack had significantly advanced the idea of the early tetrapods, scientists were still concerned about what they called the Tournaisian Gap, or about 20 million years that yielded very few fossils between the latest Devonian tetrapods and a very wide variety of primitive aquatic, secondarily aquatic and terrestrial tetrapods from the Middle and Upper Visean. In 2002, Clack looked once again at a skeleton found in 1971. She realized that it was not a fish as thought, but actually another tetaprod. This named Pederpes finneyae, at least functionally pentadactyl, was really the first to show changes for land locomotion. With a later found American Whatcheeria, it represented the next most primitive tetrapod after those of the Late Devonian. Thus, it bridged the gap that previously existed between Late Devonian and mid-Carboniferous tetrapods.

Pederpes finneyae and the other similar tetaprods were not actually made for walking. Paleontologists believed that most of them plodded along very slowly for millions of years and did not pick up the pace until about 210 million years ago. A skeletal reconstruction of the first four-legged land animal that crawled onto land suggests that it was not very comfortable in its new setting as it crept or scuffled wormlike along.

As noted, even though it was some sort of amphibian, Pederpes finneyae's bone structure was more similar to a fish. Yet it had very strong shoulders and hips that could support the body's weight without the buoyancy of water. Scientists propose two ways that it may have moved: Perhaps it did a type of forward walk with the body held rigid and the limbs moving in alternating diagonal sequence from front left and hind right and front right and hind left. The muscular forelimbs would have bent elbows, but the hind limbs would appear more flipper-like, so the pelvic region dragged along on the ground. Or, perhaps this unique animal moved along similar to an inchworm, pulling its hips and back…

Sources used in this document:
References Cited

Clack, J.A. "An Early Tetrapod from Romer's Gap." Nature (2002) 418: 72-76. [electronic version]

Clack, J.A. "From Fins to Fingers." Science 304.5667 (2004): 57-59. [electronic version]

Coates, M.I, and J.A. Clack. "Polydactyly in the Earliest Known Tetrapod Limbs"

Nature. (1990) 347: 66-69. [electronic version]
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