Craniate

Source: Wikipedia, the free encyclopedia.
Craniates
Temporal range:
Early Cambrian
- Recent
A Pacific hagfish, an example of (what was thought to be) a "non-vertebrate craniate"
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Olfactores
Clade: Craniata
Lankester, 1877[1]
Included groups
Synonyms
  • Craniota Haeckel, 1866
  • Pachycardia Haeckel, 1866
  • Vertebrata J-B. Lamarck, 1801

A craniate is a member of the Craniata (sometimes called the Craniota), a proposed

Hyperoartia (including lampreys), and the much more numerous Gnathostomata (jawed vertebrates).[3][4] Formerly distinct from vertebrates by excluding hagfish
, molecular and anatomical research in the 21st century has led to the reinclusion of hagfish as vertebrates, making living craniates synonymous with living vertebrates.

The clade was conceived largely on the basis of the Hyperoartia (lampreys and kin) being more closely related to the Gnathostomata (jawed vertebrates) than the Myxini (hagfishes). This, combined with an apparent lack of vertebral elements within the Myxini, suggested that the Myxini were descended from a more ancient lineage than the vertebrates, and that the skull developed before the vertebral column. The clade was thus composed of the Hyperoartia and the vertebrates, and any extinct chordates with skulls.

However recent studies using

Hyperoartia and Myxini) is monophyletic; this suggests that the Myxini are degenerate vertebrates, and therefore the vertebrates and craniates are cladistically equivalent, at least for the living representatives. The placement of the Myxini within the vertebrates has been further strengthened by recent anatomical analysis, with vestiges of a vertebral column being discovered in the Myxini.[5]

Characteristics

In the simplest sense, craniates are chordates with well-defined heads, thus excluding members of the chordate subphyla

sauropsids and mammals. The craniate head consists of a three-part brain, neural crest which gives rise to many cell lineages, and a cranium.[6][7]

In addition to distinct crania (sing.

In general, craniates are much more active than tunicates and lancelets and, as a result, have greater metabolic demands, as well as several anatomical adaptations. Aquatic craniates have gill slits, which are connected to muscles to pump water through the slits, engaging in both feeding and gas exchange (as opposed to lancelets, whose

red blood cells, oxygen transporting hemoglobin as well as myoglobin, livers and kidneys.[3]

Systematics and taxonomy

placoderm fish (Dunkleosteus sp.), are characterized by the presence of a cranium, mandible, and other facial bones.[8]

Linnaeus (1758)[9] used the terms Craniata and Vertebrata interchangeably to include lampreys, jawed fishes, and terrestrial vertebrates (or tetrapods). Hagfishes were classified as Vermes, possibly representing a transitional form between 'worms' and fishes.

Cyclostomes were regarded as either degenerate cartilaginous fishes or primitive vertebrates. Cope (1889)[9] coined the name Agnatha ("jawless") for a group that included the cyclostomes and a number of fossil groups in which jaws could not be observed. Vertebrates were subsequently divided into two major sister-groups: the Agnatha and the Gnathostomata (jawed vertebrates). Stensiö (1927)[9]
suggested that the two groups of living agnathans (i.e. the cyclostomes) arose independently from different groups of fossil agnathans.

Løvtrup (1977)[9] argued that lampreys are more closely related to gnathostomes based on a number of uniquely derived characters, including:

  • Arcualia (serially arranged paired cartilages above the notochord)
  • Extrinsic eyeball muscles
  • Radial muscles in the fins
  • A closely set atrium and ventricle of the heart
  • Nervous regulation of the heart by the vagus nerve
  • A typhlosole (a spirally coiled valve of the intestinal wall)
  • True lymphocytes
  • A differentiated anterior lobe of the pituitary gland (
    adenohypophysis
    )
  • Three inner ear maculae (patches of acceleration sensitive 'hair cells' used in balance) organized into two or three vertical semicircular canals
  • Neuromast
    organs (composed of vibration sensitive hair cells) in the laterosensory canals
  • An electroreceptive lateral line (with voltage sensitive hair cells)
  • Electrosensory lateral line nerves
  • A cerebellum, i.e. the multi-layered roof of the hindbrain with unique structure (characteristic neural architecture including direct inputs from the lateral line and large output Purkinje cells) and function (integrating sensory perception and coordinating motor control)

In other words, the cyclostome characteristics (e.g. horny teeth on a "tongue", gill pouches) are either instances of convergent evolution for feeding and gill ventilation in animals with an eel-like body shape, or represent primitive craniate characteristics subsequently lost or modified in gnathostomes. On this basis Janvier (1978)[9] proposed to use the names Vertebrata and Craniata as two distinct and nested taxa.

Validity

The validity of the taxon "Craniata" was recently examined by Delarbre et al. (2002) using

Cyclostomata
) and the name Craniata, being superfluous, would become a junior synonym.

The new evidence removes support for the hypothesis for the evolutionary sequence by which (from among tunicate-like chordates) first the hard cranium arose as it is exhibited by the hagfishes, then the backbone as exhibited by the lampreys, and then finally the hinged jaw that is now ubiquitous. In 2010, Philippe Janvier stated:

Although I was among the early supporters of vertebrate paraphyly, I am impressed by the evidence provided by Heimberg et al. and prepared to admit that cyclostomes are, in fact, monophyletic. The consequence is that they may tell us little, if anything, about the dawn of vertebrate evolution, except that the intuitions of 19th century zoologists were correct in assuming that these odd vertebrates (notably, hagfishes) are strongly degenerate and have lost many characters over time.[10]

Classification

dagger, †. Some are invertebrates. The positions (relationships) of the Lancelet, Tunicate, and Craniata clades are as reported.[11][12][13]

Chordata

lancelets)

Olfactores

larvacea)

Vertebrata/
Craniata

See also

Notes

  1. S2CID 83266247
    .
  2. .
  3. ^ a b c Campbell & Reece 2005 p. 676
  4. ^ Cracraft & Donoghue 2004 p. 390
  5. S2CID 17652802
    .
  6. ^ Campbell & Reece 2005 pp. 675–7
  7. ^ Parker & Haswell 1921
  8. ^ Chordates OpenStax, 9 May 2019.
  9. ^ a b c d e f Janvier, Philippe. "Craniata - Animals with skulls". Tree of Life Web Project (ToL). Tree of Life Web Project.
  10. ^ "MicroRNAs revive old views about jawless vertebrate divergence and evolution." Proceedings of the National Academy of Sciences (USA) 107:19137-19138. [1]
  11. PMID 18563158
    .
  12. .
  13. .

References