SYSTEMATIC BIOLOGY

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PHYLUM CRANIATA

INTRODUCTION TO THE CRANIATA

Craniata (cran-e-A-ta) is derived from the Greek work cranio (kpavío), which literally means head or skull.  In its Latinized form, the word becomes cranium, a term that refers to the braincase.

The craniates occupy almost every environment on the surface of the earth.  They are among the most obvious animals in marine and freshwater habitats; also, they occur in a bewildering array of terrestrial habitats where they may be found on the ground, in trees, underground, in the air, etc.  They range in size from animals of just a few grams to giants like elephants and whales.  Indeed, the mass of the female blue whale is truly astounding and marks it as the largest vertebrate that has ever lived.  Furthermore, we find the vertebrates particularly intriguing because our species is found among them.

As the name Craniata implies, these animals have an anterior brain that is encased, in part, by a cranium of cartilage or bone.  The most primitive members of the phylum,  the hagfishes (Figure A), have the typical chordate features plus the cranium.  Most members of the phylum have articulating skeletal elements (the vertebral column) that develop with or supplant the notochord.  Those with a vertebral column are called Vertebrata.  The internal skeleton developed by the vertebrates has been highly successful and allowed for the development of frames that swim, walk, run, fly, etc. (Figures B-L).  They likely appeared in the early Paleozoic and became dominant aquatic organisms by the Devonian Period (sometimes called the Age of Fishes).  Tetrapods (vertebrates with four legs) appeared also in the upper Devonian.

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A. Hagfish

B. Lamprey

C. Basking Shark

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D. Polypterus

E. Fish from the Great Barrier Reef

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F. Lungfish

G. Xenopus, the African Clawed Frog

H. Spider Monkey in Costa Rica

I. Green Sea Turtle

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J. Gecko

K. Crocodilian from Costa Rica

L. Cockatiel

Images taken from:
A: http://oceanexplorer.noaa.gov/explorations/lewis_clark01/logs/jul08/media/r609hagfish_532.jpg 
B: http://alpha1.fmarion.edu/~bio106lab/Resources/lamprey.jpg
C: http://www.ucmp.berkeley.edu/vertebrates/basalfish/chondrolh.html
D:  http://www.geol.umd.edu/~jmerck/eltsite01/projects/group2/polypterus.jpg 
F: http://mit.biology.au.dk/zoophysiology/education/cources/evolution/evol_of_airbreathing_lungfish2.jpg 
E, G-L: The Systematic Biology Diversity Collection

SYNOPTIC DESCRIPTION OF THE PHYLUM VERTEBRATA

The following information came from Margulis and Schwartz (1998),  Barnes (1984),  Storer and Usinger (1965), Colbert and Morales (1991), Tudge (2000), Nelson (2006), and Benton (2005).

 

I. SYNONYMS: vertebrates.  Craniata (Janvier 1981)

II. NUMBER: >54,703 living species known.

III. PHYLUM CHARACTERISTICS:

A. Structure

Symmetry: Bilateral.

Body Cavity: Eucoelomate, enterocoelic.

Body Covering: Covered with well-developed epidermis; often with protective dermal plates, bones, scales, epidermal scales or epidermal derivatives such as feathers or hair.

Support: Endoskeleton of bone.

Digestive System: Elaborate; most with jaws at mouth and terminal or subterminal anus.

Circulatory System: Blood system generally closed; well-developed heart. Blood elements with hemoglobin.

Locomotion: Fins and segmented myomeres in fishes. Tetrapods usually with muscles acting on jointed internal skeleton to cause movement.

Excretory System: Glomerular; operate by ultrafiltration.

Nervous System: Hollow dorsal nerve cord surrounded by segmented, bony elements (vertebrae). Anterior brain encased by bony elements (cranium).

Endocrine System: complex

Respiratory System: Gills or lungs (associated with the gut).

B. Reproduction:

Reproductive System: Sexes separate. Internal or external fertilization. Oviparous, viviparous or ovoviviparous.

Development: Larval stages or direct development.

C. Ecology: Occupy almost all free-living habitats.

SYSTEMATICS OF THE CRANIATA

Although the craniates have left a rich fossil legacy, their relationships have been clouded by preconceptions of what is advanced and what is not.  That has been complicated by our own inclusion in the phylum so that concepts of advanced become more human.  The standard class structure of the craniates can be seen below:

AGNATHA -Jawless Fishes
CHONDRICHTHYES -Sharks, skates, rays and chimeras
OSTEICHTHYES -Bony Fishes
AMPHIBIA -Amphibians
REPTILIA -Reptiles
MAMMALIA -Mammals
AVES -Birds

Growing bodies of molecular (Tudge 2000) and paleontological (Benton 2005) evidence suggest a much more complex relationship between the classes.  The agnathans represent a grade in form and include at least five independent lines.  Gnathostomes represent another structural grade and require different comparative criteria for their taxonomy, which causes the number of class-level groups to increase.  Not only are there many other groups at the class level, but the distinctions between some of the groups disappear entirely.  For example, a group like the reptiles is a grade of paraphyletic taxa if the mammals and birds are not considered to be reptiles.  Also, the bony fishes merge with the amphibians in a continuum of body form from the sarcopteryigians to the earliest tetrapods and thence to all other tetrapod decendants.  The mammals merge with the basal synapsids (traditionally an extinct group of reptiles).  Birds merge with the archosaurian reptiles (a group that contains the living crocodilians).  The turtles stand alone as a separate group as do most of the modern reptiles in a collective group (lizards, amphisbaenas, snakes, and tuataras).  Despite the cladistic structure, a classification system that places mammals within the bony fish is unsatisfactory from the perspectives of organizing biodiversity and its usefulness as a retrieval system.  So, in my opinion, a strict cladistic application is inappropriate.  However, the classical approach is equally unacceptable in that reptile is a form and not a taxonomic unit.  I have taken the organization of the following system from Benton (2005), which was written in a modified cladistic format, and adapted it to conform to a standard taxonomic system.  The following system stresses both the diversity of craniates and retains the retrieval function of a classification system.  I know full well that the following system will be unacceptable in some way to almost anyone who studies vertebrates.  So, I offer it as a way to enter the conversation about craniate systematics, not to present a system in which I am fully invested.

HIERARCHICAL CLASSIFICATION OF THE CRANIATA

The structure of the following system is a modification of Benton (2005) and Nelson (2006).  Follow the links to expanded descriptions of the taxa, many to family-level.

 

SUBPHYLUM MYXINOMORPHI

CLASS MYLLOKUNMINGIIDA+

CLASS MYXINI (HAGFISHES)

SUBPHYLUM VERTEBRATA

SUPERCLASS PETROMYZONTOMORPHI 

CLASS PETROMYZONTIDA (LAMPREYS)

SUPERCLASS CONODONTOMORPHI+

CLASS CONODONTA+

SUPERCLASS PTERASPIDOMORPHI+

CLASS PTERAPSIDA+ 

SUPERCLASS ANASPIDOMORPHI+

CLASS ANASPIDA

SUPERCLASS THELODONTOMORPHI+

CLASS THELODONTI

SUPERCLASS OSTEOSTRACOMORPHI

CLASS CEPHALASPIDOMORPHI

SUPERCLASS GNATHOSTOMATA

CLASS PLACODERMI+

CLASS CHONDRICHTHYES (SHARKS, SKATES, RAYS, CHIMERAS)

CLASS ACANTHODII+

CLASS OSTEICHTHYES (THE BONY FISHES)

CLASS STEGOCEPHALI+ -(THE TRANSITIONAL EARLY TETRAPODS)

CLASS BATRACHOMORPHA (CONTAINS EXTANT AMPHIBIA)

CLASS PROAMNIOTA+ (LEPOSPONDYLI-REPTILIOMORPHA CLADE)

CLASS EOSYNAPSIDA+- THE MAMMAL-LIKE REPTILES

CLASS MAMMALIA (CONTAINS EXTANT MAMMALS)

CLASS ANAPSIDA (CONTAINS TURTLES)

CLASS EODIAPSIDA+ (CONTAINS STEM DIAPSIDS AND ICHTHYOSAURS)

CLASS LEPIDOSAUROMORPHA (CONTAINS EXTANT LIZARDS, SNAKES, AND TUATARAS)

CLASS ARCHOSAUROMORPHA (CONTAINS CROCODILIANS & BIRDS)


This page is maintained by Jack R. Holt.  Last modified 03/10/2008.