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THE PLATYPUS: THE ANIMAL OF ALL TIME Tyler Tadian A Disbeliever in everything beyond his own reason, might

exclaim, Surely two distinct Creators must have been at work. Charles Darwin, Diary, 1836 Charles Darwin, the father of modern evolution, scribbled these profound words in his diary during his visit to the Australian Continent, unable to reassure his doubts that such a creature as the platypus could possibly fit into his understanding of monophyletic evolution. Today, even though biologists are far from understanding the platypus complete evolutionary history, modern forms of genetic and broader phylogenetic research have supported its place in the monophyletic evolutionary system. While the outer appearance of platypuses and monotremes may raise today the same doubt it did for Darwin more than hundred years ago, a thorough and interesting look into the history and biology of the platypus and its curious Order brings about a greater respect for the evolutionary process and an appreciation of the truly bizarre yet completely reasonable and practical creatures the process of evolution can produce. The platypus is one of three extant species of the egg-laying order Monotremata, the other two being the short and long-beaked echidna. The almost universally accepted phylogeny of the platypus and monotremes describes them as the first branch off therapsids (early mammallike reptiles), which eventually went on to spawn marsupials and modern-day mammals (Berczi, et. al. 1998). While some scientists believe monotremes branched off from the therapsid line with marsupials, recent amino acid sequencing by (Messer et. al. 1998) of milk proteins shows that marsupials and monotremes diverged on different branches over 20 million years apart. Even though they did not branch off the therapsid line together, monotremes and marsupials genetic and anatomical similarities can be seen in early embryonic development, with the only significant differences in monotremes appearing after hatching (Hughes and Hall, 1998). However, when compared to modern-day mammals, monotremes may appear quite bizarre in that fact that they include species of mammals that have many reptilian traits, predominantly their mode of reproducing: laying eggs. However, monotremes birthing by laying eggs, along with its other reptilian traits (a complex pectoral girdle, limbs oriented with the humerus and femur held lateral to the body, as well as the presence of a cloaca [the meaning of monotreme single hole]) all were predominant traits in their direct ancestors, therapsids (Sorin and Myers,

2000). This connection between what scientists know today about therapsid and synapsid (earlier mammal-like reptiles) anatomy and biology and the nature of modern-day monotremes is by far the strongest support for the platypus position in evolutionary development. While the structure of monotremes phylogeny is supported by ample evidence, the actual evolutionary processes that fueled the development of such bizarre creatures are worthy to be explored. First, it is important to picture the habitat and geographical location at the birth of the earliest monotremes over 100 million years ago: the earth was covered with two giant landmasses, Laurasia and Gandwana. Platypuses and other monotremes developed on Gandwana and then eventually, as the landmasses shifted to form the current continents, were isolated in both southern South America (as the discovery of an early platypus fossilized tooth supports (Pascual et. al. 1992)) and in Western Australia (Nature Video, 2008). While the South American platypuses and monotremes died out, the Australian monotremes became isolated in unique geographical locations, resembling their current habitat of rivers, inlets, lagoons and ponds in the temporal forest of Western Australia (Ojo and Omland, 2008). Theses specific habitats were also isolated by the Great Dividing Range, which still divides much of the western Australian coast. This isolation in specific habitats, following the accepted processes of evolution, quickly allowed the platypus to become specialized through natural selection; and because of small populations brought about by geographical isolation, genetic drift also most likely played a role in specializing the platypus to its aquatic habitat. Even though its ways of reproducing are somewhat simpler than the more complex placental mammals, the platypuses that evolved in the isolated biomes of Western Australia developed highly sophisticated and complex sensory receptors located in their bills used to detect the electromyographic activity from moving prey in the water and for obstacle avoidance (Proske et. al. 1998), a complex evolutionary development that took millions of years to perfect. This kind of specialization driven by evolution is jaw-dropping, but it begs the question: If platypuses could evolve and specialize to the extent their electro-sensitive bills did, why have they not moved on beyond the simpler, reptilian method of reproduction? (Phillips et. al. 2009) explains that the retention of egg-laying in early and later monotremes, while marsupials develop placental reproduction, was a response to open niches marsupials were leaving open due to their different method of reproduction. Wet, moist pond and river environments are not ideal areas for still developing, unprotected newborn marsupials to be living, due to the variability in

temperature and humidity; however, developing platypus fetuses sheltered by a leathery reptilian shell can withstand the imperfect environment of river and aquatic habitats. This use of an unrealized niche created an incentive to retain the egg-laying reproductive system (and the monotremes unique mammalian cloaca). Echidnas also show a pattern of realizing unused niche sin the Australian environment, developing a tough and rugged outer protection, while maintaining the safer and more durable egg-laying process. As (Phillips et. al. 2009) explains: Monotremes might have survived the invasion of marsupials into Australasia by exploiting ecological niches in which marsupials are restricted by their reproductive mode. Monotremes did survive, and became highly specialized even as they retained an ancient, reptilian mode of reproduction. Just two years ago, the early results of the dive into the platypus genome were published in Nature, one of the leading science journals in the global scientific community. One of the most enlightening discoveries to come out of the genome project was the analysis of platypus venom. While venom itself is thought to be a reptilian trait, the platypus DNA shows that the duplications in each of the -defensin, C-type natriuretic peptide and nerve growth factor gene families (Warren et. al. 2008) that produce the platypus venom arose independently in monotremes. This genetic information reveals a process of convergent evolution between most extant venom-producing animals, revealing a kind of evolutionary pathway by which species in different environments and under different natural pressures arrive at, by convergent evolution, more or less the same venom structure. In addition to this discovery, the platypus DNA had two matches for the ZPAX gene and another gene for the production of an egg-yolk protein called a vitellogenin, which were both thought to exist only in birds (Brown, 2008). Again, the DNA of the platypus demonstrates the manner in which genes are not created or destroyed, but merely turned on, off, or slightly altered in miniscule ways. These discoveries also further the understanding that even though monotremes are not directly linked to birds as to have inherited the use of these genes directly, they still have inherited the base of genetic material that birds and other species utilized in their evolutionary development. On the other hand, the sequencing of the platypus DNA also supports the findings of (Grutzner et. al. 2004) that point towards, not an independent convergence, but a common origin of the particular differing sex chromosomes found in birds and mammals. The X chromosome found in humans and most mammals and the Z chromosome found in birds are both found in the 10 sex chromosomes of the platypus, the Z

appearing on one end of the chain of other X chromosomes. According to (Grutzner et. al. 2004), this suggests an evolutionary link between mammal and bird sex chromosome systems, which were previously thought to have evolved independently. So while the platypus venom

shows convergent evolution at work, its sex-determining chromosomes possibly reveal a common origin of bird and mammal chromosomes that deepen the application of the understanding of descent with modification, even on the chromosomal level. Merv Griffiths, one of the foremost authorities on its biology, describes the platypus as the animal of all time (Grant and Temple-Smith, 1998), and after plunging into the science of this creature, it does not disappoint. It is almost humorous how the platypus began as the chimera of the theory of evolution, when in reality, it is the most incredible personification of evolution, how it works and what it can produce. Not only is the platypus suited perfectly for its environment and highly evolved in that way, but it is a living fossil that first appeared on the earth over 100 million years ago and yet is still alive and viable today. Its encompassing of both reptilian and mammalian traits, as well as its use of genes that were thought to be species specific, has revealed those very rare and much sought after glimpses of living evolution at its best. By some random coincidence, this prized creature found an isolated niche on the Australian continent and survived, almost entirely preserved, for millions of years, a gift to science and the inquisitive scientific mind that simply cannot be understated.

Literature Cited Berczi Sz. , Holba A. , Luckas B. , Papp E. Monotremes: About the Role of Permotriassic and Jura- Cretaceous Bounderies in the Atherian-Therian Divergence Side Tracks in Evolution. 1998 pp. 20-41 Brown, Susan. Top billing for platypus at end of evolution tree. Nature Online. May 2008. Grant, T.R., Temple-Smith, P.D.. Field Biology of the Platypus (Ornithorhynchus Anatinus): Historical and Current Perspectives. Philosophical Transactions: Biological Sciences, Vol. 353, No. 1372, Platypus Biology: Recent Advances and Reviews (Jul. 29, 1998), pp. 1081-1091 Grutzner, Frank et. al.. In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes. Nature Vol. 432, 913-917, 16 December 2004.

Hughes, Leon. , Hall, Leslie. Early Development and Embryology of the Platypus. Philosophical Transactions: Biological Sciences, Vol. 353, No. 1372, Platypus Biology: Recent Advances and Reviews (Jul. 29, 1998), pp. 1101-1114 Messer, Michael. , Weiss Anthony. , Shaw, Denis. , Westerman, Michael. Evolution of the Monotremes: Phylogenetic Relationship to Marsupials and Eutherians, and Estimation of Divergence Dates Based on a-Lactalbumin Amino Acid Sequences. Journal of Mammalian Evolution. Vol. 5, No. 1, 1998 Moyal, Ann. Platypus: The Extraordinary Story of How a Curious Creature Baffled the World. Smithsonian Institution Press, 2001. Pg. 102 (Source for opening quote) Nature VIDEO Genome Analysis of the Duck-Billed Platypus. topic from: Vol 453, Issue 7192. 08 May 2008. -------------- (I know this is a lame source : ), but it had a lot of good information that the actual journal article lacked. . .) Ojo, E. and K. Omland. 2008. "Ornithorhynchus anatinus" (On-line), Animal Diversity Web. Accessed March 25, 2010 at us_anatinus.html. Pascual, Rosendo. , Archer, Michael. , Jaureguizar, Edgardo. , Prado, Jose. , Godthelp. , Hand, Suzanne. First Discovery of Monotremes in South America. Nature 356, 704-706 (23 April 1992). Phillips, Matthew., Bennett, Thomas., Lee, Michael. Molecules, morphology, and ecology indicate a recent, amphibious ancestry for echidnas. Proceedings of the national Academy of Sciences of the United States. 14 August 2009. Proske, U., Gregory, J.E., Iggo, A. Sensory Receptors in Monotremes. Philosophical Transactions: Biological Sciences, Vol. 353, No. 1372, Platypus Biology: Recent Advances and Review (Jul. 29, 1998), pp. 1187-1198. Sorin, A. and P. Myers. 2000. "Monotremata" (On-line), Animal Diversity Web. Accessed March 24, 2010 at: Warren, Wesley., et. al.. Genome analysis of the platypus reveals unique signatures of evolution. Nature, Vol. 453, 8 May 2008. pp. 175-256.