SYSTEMATIC BIOLOGY

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SUPERCLASS OSTEOSTRACOMORPHI+

Osteostracomorphi (O-ste-o-strak-o-MORF-i) is formed from three Greek roots that mean bone [osteon (οστούν)] shell [ostraco (όστρακo)] forms [morphi (μορφή)].  The name is a reference to the bony shell in which the head is encased.

These jawless armored fish were characterized by having bony head shields like the Anaspidomorphi.  However, they had cellular bone and some had paired fins in addition to in-line fins.  The heavy armor and dorsoventral flattening of the fish suggest that the animals were bottom-dwellers.  These taxa were generally recognized as the agnathans that are closest to the gnathostomes; however, they did not have gnathostome-like gill arches or sensory line system.  They appeared in the Ordovician period and disappeared in the Devonian.

The pectoral fins of the Osteostraci bear important similarities to the paired pectoral fins of the gnathostomes.  Thus, Wilson et al. (2007) consider the paired fins of the Osteostraci to be homologous to those of the gnathostomes.  They also suggest that the galeaspids lost their paired fins.  The homology also seems to extend to other agnathans like the Thelodontomorphi.

HIERARCHICAL TAXONOMY OF THE SUPERCLASS OSTEOSTRACOMORPHI+. The following descriptions come from Benton (2005) and Nelson (2006).  The structure of the following system is also based, in part, on the phylogenetic treatment by Mikko's Phylogeny Web (from the Finnish Museum of Natural History).     Taxa in red are extinct.

 Drawing of a cephalaspid from: http://www.copernico.bo.it/subwww/lavoro%20evoluzione/agnati/Immagini/image07.jpg

SUPERCLASS OSTEOSTRACOMORPHI+

The superclass has a single class with three orders.

CLASS CEPHALASPIDOMORPHI (MONORHINA)+

The three orders of this class are united in having large bony head shields that cover the gills, which exit to the ventral side.  They had two semicircular canals, a pineal eye, and a single nostril (monorhine condition).  There is evidence the the bone was cellular.  The Cephaloaspidiformes had true paired pectoral fins.

Ordovician to Devonian

ORDER CEPHALASPIDIFORMES (OSTEOSTRACI)

Elaborate head shields. Most with paired pectoral fins.

Ordovician to Devonian.

NON-CORNUATE TAXA

Ateleaspis, Hirella, Hemicyclaspis

CORNUATA (CORNUATE TAXA-8 FAMILIES)

FAMILY BENNEVIASPIDAE

FAMILY CEPHALASPIDAE

FAMILY DARTMUTHIIDAE

FAMILY KIAERASPIDAE

FAMILY SCLERODIDAE (=SCLERODONTIDAE)

FAMILY THYESTIIDAE

FAMILY TREMATASPIDAE

FAMILY ZENASPIDAE

GENERA INCLUDE:

Trematapsis, Boreaspis, Benneviaspis, Thyestes, Sclerodus

ORDER GALEASPIDIFORMES (GALEASPIDA)

Head shields, many with elaborate processes.  No pectoral fins.  Up to 45 gill pouches.

Silurian to Devonian.

Unassigned Genera:

Macrothyraspis, Pentathyraspis.

FAMILY HANYANGASPIDAE

Hanyangaspis, Latirostraspis, Xiushuiaspis.

FAMILY DAYONGASPIDAE

Dayongaspis.

FAMILY DUYUNOLEOIDAE

Duyunolepis, Neoduyunaspis, Paraduyunaspis.

FAMILY LUNGMENSHANASPIDAE

Lungmenshanaspis, Neoduyunaspis, Qingmenaspis.

FAMILY TRIDENSASPIDAE

Tridentaspis.

FAMILY NANPANASPIDAE

Nanpanaspis.

FAMILY GALEASPIDAE

Galeaspis.

FAMILY EUGALEASPIDAE

Eugaleaspis, Sinogaleaspis, Yunnanogaleaspis.

FAMILY POLYBRANCHIASPIDAE

Cyclodiscaspis, Damaspis, Diandongaspis, Dongfangaspis, Kwangnanaspis, Laxaspis, Polybranchiaspis, Siyingia.

FAMILY HUNANASPIDAE

Antquisagittaspis, Asiaspis, Hunanaspis, Sanchaspis, Sangiaspis, Szechuanaspis.

ORDER PITURIASPIDIFORMES (PITURIASPIDA)

Known only from fragmentary fossils from Australia .  Large head shields with openings beneath the eyes.  They likely had paired pectoral fins.

Lower Devonian.

Neeyambaspis, Pituriaspis.