Palæos:		Unit 50: Thelodonti
The Vertebrates		400: Cephalaspidomorphi

Thelodonti: Cephalaspidomorphi

Galeaspids

CRANIATA
|--CONODONTA
`--+--PTERASPIDOMORPHI
   `--Thelodonti
      |--+--Furcacaudiformes
      |  `--Thelodontida 
      `--+--+--Katoporida  
         |  `--Cephalaspidomorphi 
         |     |--Galeaspida
         |     |  |--Eugaleaspidiformes
         |     |  `--Polybranchiaspidida
         |     |     |--Polybranchiaspidiformes
         |     |     `--Huananaspidiformes
         |     `--+--Pituriaspida
         |        `--Osteostraci
         |           |--Ateleaspis
         |           `--Cornuata
         `--+--Loganiidae            
            `--Gnathostomata
               |--PLACODERMI
               `--+--CHONDRICHTHYES
                  `--TELEOSTOMI

1. Cephalaspidomorphi X

2. Eugaleaspidiformes X

3. Galeaspida X

4. Huananaspidiformes X

5. Polybranchiaspidida X

6. Polybranchiaspidiformes X

Cephalaspidomorphi

"Cephalaspidomorphi" is an old name of rather uncertain meaning.  In some sources, it meant a group consisting -- roughly -- of the Heterostraci and Osteostraci, the Galeaspida and Pituriaspida being largely unknown at the time.  As the name implies, it included any jawless fish with a head shield.  During the last decades of the XXth Century, the term fell out of use because the two groups appeared to be unrelated, and the Osteostraci were suspected to be paraphyletic, or at least the sister group of the Gnathostomata.  The way things are presently moving, it seems more likely that the heterostracans will be shown to be paraphyletic.  To be specific, we suspect that the Thelodonti are Heterostracomorphi, or perhaps even Heterostraci, and that both the osteostracans and gnathostomes are thelodonts.  

However that may be, we now have a perfectly good, and highly descriptive name that isn't being used for anything worthwhile.  We have therefore retooled it to refer to osteostracans + galeaspids, a large but otherwise nameless group of jawless fishes with head shields.  ATW030604.

Galeaspida

The Galeaspida are a strange group of armored fishes possessing a massive, flattened, one-piece bony shield.  The mouth was ventral.  The dorsal shield was pierced by a single, large, nostril-like opening situated in the middle of the head in front of the eyes. The head shield was usually an elongate oval or a simple half-moon, but there were also bizarre (possibly nektonic?) forms with long tubular rostra ("nose spikes") and widely flaring cornual processes ("wings").  The body was covered by numerous ornamented scales. The bone structure of galeaspids shows hollow cavities within ornamental tubercles or blisters.  These animals may have buried themselves in the sand or mud, with only their "blow hole" showing above the surface.

Polybranchiaspis, buried in the sediment, showing the median dorsal inhalant opening at the front (left) of the head, above the eyes.  Illustration from Janvier (1996: 301).

The Galeaspids were geographically limited primarily to the South China terrane (southern China and Vietnam) from the Early Silurian to the Early Devonian.  A few are also found in Northern China, and even in Tibet (all of these areas were part of a single geographic region).   The last known type is from the Late Devonian of Ningxia, northern China.

Cephalaspidomorphi: Osteostraci + Galeaspida

Range: Early Silurian to Late Devonian

Phylogeny: Thelodonti::: Katoporida + *: Galeaspida + (Pituriaspida + Osteostraci)

Characters: massive endo- and exoskeletal head shield.  ATW030605.

Galeaspida: Hanyangaspis. 

Range: Early Silurian to Late Devonian of South China. 

Phylogeny: Cephalaspidomorphi: (Pituriaspida + Osteostraci) + *: Polybranchiaspidida + Eugaleaspidiformes. 

Characters: Overall morphology similar to Osteostraci. Massive endo- & exoskeletal head shield; endoskeletal shield is solid cartilage lined with perichondral bone and some calcified cartilage; length of shield variable; mouth & gill openings ventral; ventral plates small; orbits usually small & dorsal (anterior in Hanyangaspis); $ large anterior median dorsal opening leading to oralobranchial cavity (inhalant duct?); 2 separate nasal olfactory tracts open into this duct; large number of gill openings (up to 45); shield cornua & rostra common (especially in Huananaspidiformes); $ main lateral line has lateral processes on shield, sometimes in scalloped pattern; 2 semicircular canals; body has minute aspidine-like scales with bulging base, usually with enameloid cap; bone acellular with no dentine; boundary between exo- and endo-skeleton is vascularized (subaponevrotic vascularization); shield apparently formed by fusion of scale units & may grade into body squamation. Unlike Osteostraci, no dorsal sensory fields or associated sensory canals, no paired fins; no dorsal or anal fins. Fresh water & marine. 

Links: Galeaspida; Class Galeaspida; Geol 437 jawless fishes; Galeaspida. 

Image: Bannhuanaspis from the Early Devonian Bac Bun Formation of Vietnam, after Janvier (1996). ATW010421.

Eugaleaspidiformes: Eugaleaspis, Sinogaleaspis, Yunnanogaleaspis. 

Range: Early Silurian to Early Devonian. 

Phylogeny: Galeaspida: Polybranchiaspidida + *.

Characters: short, horseshoe-shaped headshield; slit-shaped median dorsal opening;  cornual processes short; 6-8 pairs of gill openings with branchial fossae for gills (primitive number).

Links: Galeaspida

References: Janvier (1996).  ATW020715.

Polybranchiaspidida: 

Range: Early Devonian (to Late Devonian?)

Characters: $ >10 (up to 45) pairs of branchial fossae.

Phylogeny: Galeaspida: Eugaleaspidiformes + *: Polybranchiaspidiformes + Huananaspidiformes.

References: Janvier (1996). 

Polybranchiaspidiformes: Dongfangaspis, Duyunolepis, Polybranchiaspis

Range: Early Devonian (to Late Devonian?)

Phylogeny: Polybranchiaspidida: Huananaspidiformes + *.

Characters: headshield oval or rounded; median dorsal opening oval or slightly bean-shaped; elaborate rostral and cornual processes absent; most have star-shaped pattern at lateral end of transverse sensory lines on headshield. 

Note: [1] Likely paraphyletic stem group of Polybranchiaspidida. [2] includes largest galeaspids (~20 cm). 

Links: Galeaspida; 

References: Janvier (1996).  ATW030404.

Huananaspidiformes: Asiaspis, Gantarostraspis, Lungmenshanaspis, Sanchaspis, Sanqiaspis. 

Range: Early Devonian of South China. 

Phylogeny: Polybranchiaspidida: Polybranchiaspidiformes + *. 

Characters: Median dorsal opening may have unusual shapes; $ long, sometimes spatulate, rostra and slender cornua; numerous branchial openings. Janvier suggests some may have been nectonic based on shield shape. 

References: Janvier (1996).  ATW991113.

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