19.1.13: Vertebrates - Biology

19.1.13: Vertebrates - Biology

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During their embryonic development, all chordates pass through a stage called the pharyngula with these features:

  • a dorsal, tubular nerve cord ("1") running from anterior to posterior. At its anterior end, it becomes enlarged to form the brain.
  • a flexible, rodlike notochord ("2") that runs dorsal to the digestive tract and provides internal support. In vertebrate chordates, it is replaced by a vertebral column or backbone long before maturity.
  • pairs of gill pouches. These lateral outpocketings of the pharynx are matched on the exterior by paired grooves. In aquatic chordates, one or more pairs of gill pouches break through to the exterior grooves, forming gill slits ("3"). These provide an exit for water taken in through the mouth and passed over the gills.
  • a tail that extends behind the anus
  • There are three subdivisions of the chordates:
    1. Cephalochordata
    2. Tunicata

      The cephalochordates and tunicates never develop a vertebral column. They are thus "invertebrates" and are discussed with the other invertebrates.

    3. Craniata The vast majority of chordates have a skull enclosing their brain, eyes, inner ear, etc.). All but one group of these (the hagfishes) also convert their notochord into a vertebral column or backbone thus qualifying as vertebrates.


Although hagfishes, never replace their notochord with a vertebral column, and thus might seem not to qualify as vertebrates, they share a number of other features with other vertebrates and certainly should be classified with them. Still uncertain is whether they represent the most primitive vertebrates or are simply degenerate vertebrates (probably the latter).

All the other members of the craniata convert their notochord into a vertebral column or "backbone" (even though in some it is made of cartilage not bone). They also differ from all other animals in having quadrupled their HOX gene cluster; that is, they have 4 different clusters of HOX genes (on 4 separate chromosomes). Perhaps this acquisition played a key role in the evolutionary diversity that so characterizes the vertebrates.

The vertebrates are subdivided into the

  • jawless vertebrates (Agnatha)
  • jawed vertebrates (Gnathostomata)


Lampreys and hagfishes are the only jawless vertebrates to survive today. They both have a round mouth and for this reason are often referred to as cyclostomes. They are the most primitive of the vertebrates. By "primitive", a biologist means that they are the least changed from the first vertebrates. Besides lacking jaws,

  • They have no paired pectoral (shoulder) or pelvic (hip) fins.
  • Their notochord persists for life, never being completely replaced by a backbone even in the lampreys.
  • They have no scales.
  • The axons of their neurons are unmyelinated (like those of all invertebrates).
  • Lampreys have both an innate immune system and an adaptive immune system, but the latter is entirely different from that found in the jawed vertebrates.

The photo (courtesy of the Carolina Biological Supply Company) is of the West Coast lamprey. Note the gill slits and the absence of paired pectoral and pelvic fins.


As well as having jaws, all the members of this group have

  • Myelin sheaths around the axons of their neurons. This permits much more rapid transmission of nerve impulses - a trait probably as important for active vertebrates as their jaws.
  • An adaptive immune system backing up their innate immune system.

Cartilaginous Fishes (Chondrichthyes)

Fossils of cartilaginous fishes become abundant in deposits dating to the Devonian period. They were very much like the sharks of today. The group, which today is made up of some 1,188 species of sharks, skatesand rays gets its name from the fact that their skeleton is made of cartilage, not bone.

With their gills exposed to sea water, all marine fishes are faced with the problem of conserving body water in a strongly hypertonic environment. Sea water is about 3.5% salt, over 3 times that of vertebrate blood. The cartilaginous fishes solve the problem by maintaining such a high concentration of urea in their blood (2.5% — far higher than the ~0.02% of other vertebrates) that it is in osmotic balance with - that is, is isotonic to - sea water.

This ability develops late in embryology, so the eggs of these species cannot simply be released in the sea. Two solutions are used:

  • Enclose the egg in an impervious case filled with isotonic fluid before depositing it in the sea.
  • Retain the eggs and embryos within the mother's body until they are capable of coping with the marine environment.

Both these solutions require internal fertilization, and the cartilaginous fishes were the first vertebrates to develop this. The pelvic fins of the male are modified for depositing sperm in the reproductive tract of the female.

Bony Fishes (Osteichthyes)

As their name indicates, the skeleton in this group is made of bone. The group is subdivided into the

  • ray-finned fishes (Actinopterygii)
  • lobe-finned fishes (Sarcopterygii)

Ray-finned fishes

  • Their fins are thin and supported by spines.
  • There are over 30,000 species (representing more than half of all living vertebrates).
  • They are an important part of the human diet in many areas of the world and, in affluent nations, support a large sports fishing industry.

Although the earliest bony fishes may have appeared late in the Silurian period, their fossils become abundant in freshwater deposits of the Devonian period. In addition to gills, these fishes had a pair of pouched outgrowths from the pharynx which served as lungs. They were inflated with air taken in through the mouth and may have provided a backup gas exchange organ when the water became too warm and stagnant to carry enough dissolved oxygen. Their kidneys were adapted for the hypotonic environment in which they lived.

These animals diversified through the remainder of the Devonian period (which is often called the "Age of Fishes"). Some migrated to the oceans. In this more stable environment, their lungs became transformed into a swim bladder with which they could alter buoyancy. Their kidneys became transformed as well adapting them to their new - hypertonic - surroundings.

Lobe-finned fishes

The only ones to survive today are:

  • two species of coelacanths. Coelacanths were long thought to have become extinct at the end of the Mesozoic era, some 70 million years ago. But in December 1938, a living coelacanth, Latimeria chalumnae, was pulled up from the depths of the ocean off the east coast of Africa. Since then, over 200 additional specimens have been caught.
  • several species of lungfish found in Africa, South America, and Australia.

The nostrils of bony fishes open only to the outside and are used for smelling. Some of the lobe-finned fishes developed internal openings to their nostrils. This made it possible to breath air with the mouth closed as modern lungfishes do.

Judging from present-day lungfishes, two other significant adaptations evolved in this group:

  • two atria and a partial septum in the ventricle of the heart (similar to the frog heart). This permitted a partial separation of oxygenated blood returning from the lung(s) and the deoxygenated blood returning from the rest of the body.
  • an enzyme system to convert ammonia into the less toxic urea. This mechanism is highly-developed in the African and South American lungfishes. While in the water, these fishes excrete their waste nitrogen as ammonia, just as most ray-finned fishes do. In time of drought, these animals burrow in the mud and switch to urea production.

With their bony limbs and lungs inherited from their lobe-finned ancestors, amphibians were so successful during the Carboniferous (Mississippian and Pennsylvanian periods) that these periods are known as the Age of Amphibians.

The Carboniferous was followed by the Permian, when the earth became colder and dryer. The fortunes of the amphibians began to decline until only three groups - totaling about 6500 species - remain today:

  • frogs and toads (Anura) (The one pictured is Rana pipiens, the leopard frog.)
  • salamanders and newts (Urodela)
  • caecilians (Apoda), which are rare, limbless, tropical animals.

As the name suggests, amphibians are only semiterrestrial:

  • Their skin is soft and moist so they are at risk of desiccation in dry surroundings.
  • Their eggs have no waterproof covering so
    • they must be laid in water (which makes them useful animals for studying embryonic development) where they are fertilized or
    • placed within the mother's body (some use a pouch in the skin, some use their mouth, some even use their stomach — which stops secreting acid and enzymes for the duration!) after external fertilization.

Amniotes (Amniota)

Some 310 million years ago (in the Pennsylvanian), some amphibians evolved the ability to lay shelled, yolk-filled eggs. The embryo developing within the egg produces 4 extraembryonic membranes:

  • amnion, which surrounds the embryo with a fluid as watery as the pond water around a frog's egg (and accounts for the name amniota)
  • chorion, which serves for gas exchange
  • allantois, which serves both for gas exchange and to store metabolic wastes
  • yolk sac, which supplies the embryo with food

With the arrival of the cold, dry Permian, reptiles were well-adapted to survive because of their development of a shelled, yolk-filled egg which could be deposited on land without danger of drying out. The photo (courtesy of the Carolina Biological Supply Company) shows an American chameleon emerging from its egg.

Other adaptations that enabled the reptiles to flourish for the next 220 million years were:

  • a dry, water-impermeable skin
  • lungs inflated by expansion of the rib cage
  • a partial septum in the ventricle reducing the mixing of oxygenated and deoxygenated blood

Beginning late in the Paleozoic era and exploding in the Triassic period, the reptiles underwent a remarkable adaptive radiation producing the diapsids. This group developed the ability to convert their nitrogenous waste into uric acid. Uric acid is almost insoluble in water so its excretion involves little loss of water. (It is the whitish paste that pigeons leave on statues.) This modification largely freed the diapsids and their descendants from a dependence on drinking water; the water in their food is usually sufficient.

Diapsid evolution soon produced:

  • lizards and snakes (Squamata - some 6,300 species survive today);
  • turtles
  • thecodonts.

Thecodonts were able to run fast by rising up on their hind legs, which became larger than their front legs, and using their long tail for balance. The group diversified into:

  • crocodiles and alligators (Crocodilia — 22 species survive today)
  • an extraordinary array of dinosaurs from some of which evolved today's birds.

Feathers are the feature that most clearly distinguishes the birds from their dinosaur ancestors. These scaly skin outgrowths provide a light, strong surface for the wings;

  • heat insulation, making it possible to be small but still warm-blooded.

Other adaptations are:All of these adaptations help birds to fly (to escape predators and find suitable food and nesting sites). Almost 10,000 species are known today.

March 2021

Wodarz Dominik, Komarova Natalia L, Schang Luis M. Role of high-dose exposure in transmission hot zones as a driver of SARS-CoV-2 dynamics. J R Soc Interface. 2021 Mar18(176):20200916.

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350 Publications

Our company is committed to advancing knowledge and conservation. This bibliography lists the contributions to scientific literature we have made over the course of our history: more than 350 peer-reviewed articles, book chapters and books (authored in whole or part) by our team members. (The names in bold indicate an affiliation with our company.) These publications include research on a wide variety of marine and terrestrial animals in numerous arenas of investigation. These studies reflect the knowledge, skill, commitment and passion of our staff. Beyond these direct contributions, still more manuscripts were authored by outside researchers that utilized our animals and data. Those outside studies benefited from access to our animals in a controlled environment that is impossible to duplicate in the wild.


Ambati, B. K., Nozaki, M., Singh, N., Takeda, A., Jani, P. D., Suthar, T., Albuquerque, R. J. C., Richter, E., Sakurai, E., Newcomb, M. T., Kleinman, M. E., Caldwell, R. B., Lin, Q., Ogura, Y., Orecchia, A., Samuelson, D. A., Agnew, D. W., St. Leger, J., Green, W. R., Mahasreshti, P. J., Curiel, D. T., Kwan, D., Marsh, H., Ikeda, S., Leiper, L. J., Collinson, J. M., Bogdanovich, S., Khurana, T. S., Shibuya, M., Baldwin, M. E., Ferrara, N., Gerber, H., De Falco, S., Witta, J., Baffi, J. Z., Raisler, B. J. and Ambati, J. 2006. Corneal avascularity is due to soluble VEGF receptor-1. Nature, 443(7114): 993-997.

Anderson, K., Garner, M., and Stedman, N. 2016. Retrospective Evaluation Of Histopathologic Findings In Captive Gazelle Species. Journal of Zoo and Wildlife Medicine, 47(1):113-119.

Andrews, B., Davis, W. and Parham, D. 2001. Corporate response and facilitation of the rehabilitation of a California gray whale calf. Aquatic Mammals. 27(3):209-211.

Anthony, S.J., St. Leger, J. A., Pugliares, K., Ip, H.S., Chan, J.M., Carpenter, Z.W., Navarette-Macias, I., Sanchez-Leon, M., Saliki, J.T., Pedersen, J., Karesh, W., Daszak, P., Rabadan, R., Rowles, T. and Lipkin, W.I. 2012. Emergence of fatal avian influenza in New England harbor seals. mBio. 3(4): e00166-12.

Anthony, S.J., St. Leger, J. A., Navarrete-Macias, I., Nilson, E., Sanchez-Leon, M., Liang, E., Seimon, T., Jain, K., Karesh, W., Daszak, P., Briese, T. and Lipkin, W.I. 2013. Identification of a novel cetacean polyomavirus from a common dolphin (Delphinus delphis) with tracheobronchitis. PLoS ONE. 8(7): e68239.

Antonellis, G. A., S. Leatherwood, L. H. Cornell, and J. E. Antrim 1981. Activity cycles, feeding rates and food preferences of captive sea otters, Enhydra lutris. The Murelett., 62:6-9.

Antrim, J. 2001 Logistics of maintenance, rehabilitation and return to the pacific ocean of a California gray whale calf. Aquatic Mammals. 27(3):228-230.

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Antrim, J., J. F. McBain, and D. Parham 1998. Rehabilitation and release of a gray whale calf: J.J.&rsquos story. Endangered Species Update. 15(5):84-89.

Argade, S., Smith, C. R., Shaw, T., Zupkas, P., Schmitt, T. L., Venn-Watson, S. and Sur, R. L. 2013. Solubility of ammonium acid urate nephroliths from bottlenose dolphins (Tursiops truncatus). Journal of Zoo and Wildlife Medicine. 44(4), 853-858.

Asper, E. D. 1975. Techniques of live-capture of smaller Cetacea. Journal of the Fisheries Research Board, Canada. 32(1):191-196.

Asper, E. D., and J. H. Prescott 1970. Courtship and nesting of captive black-footed albatrosses, Diomedia nigripes. International Zoo Yearbook. 10: 90-92.

Asper, E. D., L. H. Cornell, D. A. Duffield, and L. Dimeo-Ediger 1988. Marine mammals in zoos, aquaria and marine zoological parks in North America: 1983 census report. International Zoo Yearbook. 27:287-294.

Asper, E.D. and Cornell, L.H. 1977. Live capture statistics for the killer whale (Orcinus orca) 1961-1976 in California, Washington and British Columbia. Aquatic Mammals. 5(1):21-26.

Asper, E.D., Cornell, L.H., Duffield, D.A., Odell, D.K., Joseph, B.E., Stark, B.L. and Perry, C.A. 1990. Hematology and serum chemistry values in bottlenose dolphins. pp. 479-485 In: Leatherwood, S., Reeves, R. (Eds). The Bottlenose Dolphin. Academic Press, Inc., San Diego, CA.

Asper, E.D., Young, G.W. and Walsh, M.T. 1988. Observations on the birth and development of a captive-born killer whale. International Zoo Yearbook. 27:295-304.

Barros, N. B., and D. K. Odell 1990. Food habits of bottlenose dolphins (Tursiops truncatus) in the southeastern United States. pp. 309-328. In: S. Leatherwood and R. Reeves (Eds.), The Bottlenose Dolphin. Academic Press: San Diego, CA.

Bechert. U., Christensen, M.J., Poppenga, R., Le, H., Wyatt, J. and Schmitt, T. 2010. Pharmacokinetics of orally administered terbinafine in African penguins (Spheniscus demersus) for potential treatment of aspergillosis. Journal of Zoo and Wildlife Medicine. 41(2):263-274.

Begeman, L., St. Leger, J.A., Blyde, D.J., Jauniaux, T.P., Lair, S., Lovewell, G., Raverty S., Seibel H., Siebert U., Staggs, S.L., Martelli P. and Keesler, R.I. 2013. Intestinal volvulus in cetaceans. Veterinary Pathology. 50(4): 590-596.

Bejarano, A. C., Gulland, F. M., Goldstein, T., St. Leger, J., Hunter, M., Schwacke, L. H., Van Dolah, F. M. and Rowles, T.K. 2008. Demographics and spatio-temporal signature of the biotoxin domoic acid in California sea lion (Zalophus californianus) stranding records. Marine Mammal Science. 24(4): 899-912.

Beldegreen, R.A., and E.D. Asper 1980. Captive Breeding and Husbandry Techniques for Adelie Penguins. International Zoo Yearbook. 20:197-200.

Benirschke, K., and L. H. Cornell 1987. The placenta of the killer whale, Orcinus orca. Marine Mammal Science. 3(1):82-86.

Benz, G. W., P. J. Mohan, and G. L. Crow 1992. Developmental stages of Paralebion elongatus from aquarium-held reef whitetip sharks (Triaenodon obesus) captured in Hawaiian waters. Journal of Parasitology. 78(6):1027-1035.

Bergfelt, D.R., Blum, J.L., Steinetz, B.G., Steinman, K.J., O&rsquoBrien, J.K. and Robeck, T.R. 2015. Relaxin as a hormonal aid to evaluate pregnancy and pregnancy loss in bottlenose dolphins (Tursiops truncatus). General and Comparative Endocrinology, doi: .

Berman-Kowalewski, M., Gulland, F.M.D., Wilkin, S., Calambokidis, J., Mate, B., Cordaro, J., Rotstein, D., St. Leger, J., Collins, P., Fahy, K. and Dover, S. 2010. Association between blue whale (Balaenoptera musculus) mortality and ship strikes along the California coast. Aquatic Mammals. 36(1): 59-66.

Beusse, D. O., E. D. Asper, J. N. Baucom, and S. W. Searles 1977. Diethylcarbamazine citrate for prevention of heartworm (Dirofilaria immitis) in the California sea lion (Zalophus californianus). Veterinary Medicine &ndash Small Animal Clinician. 1977:470-471.

Bissell, H. 2014. Nutritional implications of the high-elevation lifestyle of Rhinopithecus bieti. pp.199-210 In: Grow, N.B., Gursky-Doyen, S., Krzton, A. (Eds.) High Altitude Primates, Developments in Primatology: Progress and Prospects 44. Springer-Verlag, New York.

Black, P. A., Macek, M., Tieber, A. and Weber, M. 2013. Reference values for hematology, plasma biochemical analysis, plasma protein electrophoresis, and aspergillus serology in elegant-crested tinamou (Eudromia elegans). Journal of Avian Medicine and Surgery. 27(1): 1-6.

Black, P.A., Trent, A.M., Bauman, J.E., Farnsworth, R.J., Monfort, S.L..2011. Fecal steroid analysis for evaluating ovarian function in the greater kudu (Tragelaphus strepsiceros) and lesser kudu (Tragelaphus imberbis). Journal of Zoo and Wildlife Medicine. 42(4):558-64.

Black, P.A., D. L. McRuer, and L. Horne. 2011. Hematologic parameters in raptor species in a rehabilitation setting before release. Journal of Avian Medicine and Surgery, 25(3):192-198.

Blankenship, B., Dold, C., Jensen, E., Smith, C., Van Bonn, W. and Ridgway, S. 2008. Neuronal migration defect: A case of subcortical heterotopia in a California sea lion. Veterinary Pathology 45(3): 412-416.

Bodley, K., and T.L. Schmitt. 2014. Penguins, pp. 435-443. In G. West, D.J. Heard, and N. Caulkett (Eds), Zoo Animal and Wildlife Immobilization and Anesthesia, Second Edition. John Wiley & Sons, Inc., Ames, IA.

Bossart, G. D., T. Reidarson, L. A. Dierauf, and D. Duffield 2001. Pathology of Marine Mammals: Clinical Pathology. In: L. A. Dierauf and F. M. D. Gulland (Eds.), CRC Handbook of Marine Mammal Medicine. CRC Press: Boca Raton, FL.

Bowles, A. E., W. G. Young, and E. D. Asper 1988. Ontogeny of stereotyped calling of a killer whale calf, Orcinus orca, during her first year. Rit Fiskideildar., 11:251-275.

Branstetter, B.K., St. Leger, J., Acton, D., Stewart, J., Houser, D., Finneran, J.J., and Jenkins, K. 2017. Killer whale (Orcinus orca) behavioral audiograms. Acoustical Society of America. 141(4)2387-2398. doi:10.1121/1.4979116.

Brodie, E. C., Gulland, F.M.D., Greig, D.J., Hunter, M., Jaakola, J., St. Leger, J., Leighfield, T.A. and Van Dolah, F.M. 2006. Domoic acid causes reproductive failure in California sea lions (Zalophus californianus). Marine Mammal Science. 22(3):700-707.

Browning, N. E., Dold, C., I-Fan, J., and Worthy, G. A. J. 2014. Isotope turnover rates and diet-tissue discrimination in skin of ex situ bottlenose dolphins (Tursiops truncatus). The Journal of Experimental Biology. 217: 214-221.

Bruehler, G. L., DiRocco, S., Ryan, T. and Robinson, K. 2001. Husbandry and hand-rearing of a rehabilitating California gray whale calf. Aquatic Mammals. 27(3): 222-227.

Buck, C., G. P. Paulino, D. J. Medina, G. D. Hsiung, T. W. Campbell, and M. T. Walsh 1993. Isolation of St. Louis encephalitis virus from a killer whale. Clinical Diagnostic Virology. 1:109-112

Buckles, E. L, Lowenstine, L. J., Funke, C., Vittore, R. K., Wong, H. N., St. Leger, J., Greig, D. J., Duerr, R. S., Gulland, F. M. D. and Stott, L. J. 2006. Otarine herpesvirus-1, not papillomavirus, is associated with endemic tumours in California Sea Lions (Zalophus californianus). Journal of Comparative Pathology. 135(4): 183-189.

Buckles, E. L., Lowenstine, L.J., DeLong, R.L, Melin, S.R., Vittore, R.K., Wong, H.N., Ross, G.L, St. Leger, J.A., Greig, D.J., Duerr, R.S., Gulland, F.M.D. and Stott, J.L. 2007. Age-prevalence of otarine herpesvirus-1, a tumor-associated virus, and possibility of its sexual transmission in California sea lions. Veterinary Microbiology. 120: 1-8.

Burch, L. E., and C. Gailband 2000. Comparison of hand-reared Caribbean flamingos and lesser flamingos at SeaWorld California. Waterbirds., 23 (Special Publication 1):193-197.

Burton, M., J. Olsen, R. Ball, and G. Dumonceaux. Mycobacterium aviaum subsp. Paratuberculosis infection in an addax (Addax nasomaculatus). Journal of Zoo and Wildlife Medicine. 32(2): 242-244.

Butterworth, A., S.C. Kestin, and J.F. McBain. 2004. Evaluation of baseline indices of sensibility in captive cetaceans. Veterinary Record 155: 513-518.

Campbell, T. W. 1990. Common parasites of exotic pets &ndash part I. Veterinary Technician. 11(4):223-225.

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Campbell, T. W., and B. Phillips 1993. Rehabilitation of sea turtles. Veterinary Technician. 14(3):165-169.

Campbell, T. W., J. D. Letcher, and D. H. Nielsen 1993. Diseases of aquarium fish. pp. 45-49. In: P. W. Pratt (Ed.), Review Questions and Answers for Veterinary Boards, Ancillary Topics. American Veterinary Publications, Inc.:Goleta, CA.

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Cassiano, E.J., Wittenrich, M.L., Violetta, G.C., and Watson, C.A. 2012. Growth and survival of poarkfish (Anisotremus virginicus) large: comparing rotifers and copepod nauplii during first feeding. ABAH Bioflux. 4(2):72-78.

Clark, S. 2002. First report of albinism in the white-spotted bamboo shark, Chiloscyllium plagiosum (Orectolobiformes: Hemiscyllidae), with a review of reported color aberrations in elasmobranchs. Zoo Biology. 21(6): 519-524.

Clark, S. T. 2000. Age, growth, and distributions of the giant snake eel, Ophichthus rex, in the Gulf of Mexico. Bulletin of Marine Science. 67(3): 911-922.

Clark, S. T. and D. K. Odell 1999. Allometric relationships and sexual dimorphism in captive killer whales (Orcinus orca). Journal of Mammalogy. 80(3): 777-785.

Clark, S. T., and Odell, D. K. 1999. Nursing parameters in captive killer whales (Orcinus orca). Zoo Biology. 18(5): 373-384.

Clark, S. T., Odell, D. K. and Lacinak, C. T. 2000. Aspects of growth in captive killer whales (Orcinus orca). Marine Mammal Science. 16(1): 110-123.

Clark, S.T. and Odell, D. K. 1999. Nursing behaviors in captive false killer whales (Pseudorca crassidens). Aquatic Mammals. 25(3): 183-191.

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(1) There is significant potential to improve our knowledge of development in Ediacaran macro-organisms, but the synthesis of existing data allows us to refute several previously proposed phylogenetic affinities for key Ediacaran taxa. Analysis of development in rangeomorphs and dickinsoniomorphs reveals congruence with aspects of metazoan development.

(2) We conclude that developmental data alone allow us to identify Dickinsonia, Andiva, Yorgia and the rangeomorphs as early metazoans.

(3) Morphogenesis offers promise for disentangling Ediacaran phylogenetic relationships and the evolution of development. Although the study of ontogeny is the study of change over time, by adopting a largely morphological approach when considering Ediacaran organisms, the ‘change’ has been largely overlooked. Future study of populations of organisms will allow better quantification of this change, as well as the production of growth models, both of which will ultimately increase the precision of phylogenetic resolution of Ediacaran organisms.

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