Taxa modified

Nectaspis areolate renamed Nectaspis areolata

Vernicomacanthus waynensis renamed Dobunnacanthus waynensis

 

Taxa added

Ateleaspis tessellata

Kolymaspis sibirica

Minjinia turgenensis

 

Characters deleted (numberings from King et al. 2017):

Character 22 (Elongate preorbital region between orbits and nasal capsules) deleted.

Character 173 (ZHU169, GFB60 Type of dermal neck-joint) deleted.

Character 174 (Type of ginglymoid neck joint) deleted.

Character 192 (ZHU168 Dermal neck-joint between paired main-lateral-line-bearing bones of skull) deleted.

 

Characters added:

Character 7 (Laminar basal layer) added.

Character 8 (Cancellous central layer) added.

Character 9 (Dentinous external layer) added.

Character 25 (Extended preorbital region between eyes and nasal capsule) added.

Character 52 (Crus commune) added.

Character 177 (ZHU168* Dermal cranio-thoracic articulation between paired main-lateral-line-bea) added.

Character 178 (Dermal neck joint facet morphology) added.

Character 179 (Para-articular process of dermal neck joint) added.

Character 180 (Endoskeletal craniothoracic (sixth branchial) facet) added.

 

 

Table of state score changes

 

Taxon

 Orig. char. no.

 New char. no.

 Character label

 Old state

 New state

Justification

Acanthodes bronni

140

144

(DFC37 Basihyal)

1 (absent, hyoid arch articulates directly with basibranchial)

0 (present)

We code Acanthodes as having a basihyal after Dearden et al. (2019)

Achoania jarvikii

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Achoania jarvikii

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Achoania jarvikii

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Benneviaspis holtedahli

96

100

(Nasal capsules in anterlolateral corners of orbit)

0 (no)

- (inapplic.)

Although olfactory bulbs are present, discrete nasal capsules appear to be absent in osteostracans.

Benneviaspis holtedahli

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Boreaspis macrorhynchus

96

100

(Nasal capsules in anterlolateral corners of orbit)

0 (no)

- (inapplic.)

Although olfactory bulbs are present, discrete nasal capsules appear to be absent in osteostracans.

Boreaspis macrorhynchus

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Brindabellaspis stensioi

48

51

(DFC86 Sinus superior)

0 (absent or indistinguishable from union of anterior and posterior canals with saccular chamber)

1 (present)

Updated based on Zhu et al. (2021, Curr. Biol.)

Brindabellaspis stensioi

80

84

([GFB141]Subcranial ridges)

- (inapplic.)

1 (present)

We consider the laterobasal angles of brindabellaspidids to be primary homologues of these ridges based on topological position.

Brindabellaspis stensioi

97

101

(vagal process)

- (inapplic.)

0 (forked)

A vagal process (or its homologue) is evidently present as documented by Young (1989. Palaeontographica A, figs. 7, 13)

Brindabellaspis stensioi

114

118

(Hypophysial foramen in braincase)

1 (present)

0 (absent)

There is no evidence of a foramen according to Young (1980) and Zhu et al. (2021)

Brindabellaspis stensioi

187

193

(ZHU159 Parietals (preorbitals of placoderms) surround pineal foramentotoeminence)

1 (no)

0 (yes)

Updated based on Zhu et al. (2021, Curr. Biol.)

Brindabellaspis stensioi

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Brindabellaspis stensioi

248

252

(Preorbital recess)

- (inapplic.)

0 (absent)

Character is not applicable to taxa without an upper lip/premedian area.

Cephalaspis lyelli

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Cephalaspis lyelli

423

428

(DFC106 Dermal shoulder girdle forming a complete ring around the trunk)

0 (present)

1 (absent)

 

Cheirolepis canadensis

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Cheirolepis canadensis

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Cheirolepis canadensis

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Cheirolepis trailli

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Cheirolepis trailli

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Cheirolepis trailli

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Diabolepis speratus

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Diabolepis speratus

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Diabolepis speratus

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Entelognathus primordialis

84

88

([GFB166]Branchial ridges)

2 (absent (articulation made with bare cranial wall))

1 (reduced to vagal process)

Zhu et al. (2013) figure a small posterior postorbital process in Entelognathus which is homologizeable with the vagal process (after Giles et al. 2015, Nature).

Escuminaspis laticeps

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Eusthenopteron foordi

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Eusthenopteron foordi

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Eusthenopteron foordi

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Gemuendina stuertzi

186

192

(ZHU158 Location of pineal foramen/eminence)

1 (well posterior of orbits)

0 (level with posterior margin of orbits)

The pineal opening of rhenanids is situated between the orbits and therefore cannot be considered well posterior to them (Gross 1963; Stensiö, 1969)

Gemuendina stuertzi

246

251

(Preorbital depression)

- (inapplic.)

1 (present)

Character is not applicable to taxa without an upper lip/premedian area.

Glyptolepis groenlandica

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Glyptolepis groenlandica

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Glyptolepis groenlandica

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Gogonasus andrewsae

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Gogonasus andrewsae

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Gogonasus andrewsae

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Guiyu oneiros

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Guiyu oneiros

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Guiyu oneiros

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Hemicyclaspis murchisoni

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Hemicyclaspis murchisoni

423

428

(DFC106 Dermal shoulder girdle forming a complete ring around the trunk)

0 (present)

1 (absent)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Howqualepis rostridens

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Howqualepis rostridens

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Howqualepis rostridens

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Jagorina pandora

186

192

(ZHU158 Location of pineal foramen/eminence)

1 (well posterior of orbits)

0 (level with posterior margin of orbits)

The pineal opening of rhenanids is situated between the orbits and therefore cannot be considered well posterior to them (Gross 1963; Stensiö, 1969)

Kenichthys campbelli

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Kenichthys campbelli

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Kenichthys campbelli

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Kentuckia deani

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Kentuckia deani

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Kentuckia deani

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Lunaspis broili

158

162

(DFC25 Pineal opening perforation in dermal skull roof)

1 (absent)

0 (present)

A pineal opening is present in Lunaspis (Gross 1961, fig. 1)

Lunaspis broili

450

455

(DFC136 Number of dorsal fins, if present)

0 (one)

- (inapplic.)

Dorsal fins are absent in Lunaspis (Gross 1961)

Microbrachius dicki

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Microbrachius dicki

248

252

(Preorbital recess)

- (inapplic.)

0 (absent)

Character is not applicable to taxa without an upper lip/premedian area.

Mimipiscis toombsi

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Mimipiscis toombsi

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Mimipiscis toombsi

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Moythomasia durgaringa

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Moythomasia durgaringa

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Moythomasia durgaringa

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Norselaspis glacialis

96

100

(Nasal capsules in anterlolateral corners of orbit)

0 (no)

- (inapplic.)

Although olfactory bulbs are present, discrete nasal capsules appear to be absent in osteostracans.

Norselaspis glacialis

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Norselaspis glacialis

423

428

(DFC106 Dermal shoulder girdle forming a complete ring around the trunk)

0 (present)

1 (absent)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Onychodus jandemarrai

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Onychodus jandemarrai

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Onychodus jandemarrai

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osorioichthys marginis

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osorioichthys marginis

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osorioichthys marginis

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osteolepis macrolepidotus

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osteolepis macrolepidotus

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Osteolepis macrolepidotus

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Porolepis sp

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Porolepis sp

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Porolepis sp

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Powichthys thorsteinssoni

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Powichthys thorsteinssoni

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Powichthys thorsteinssoni

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Procephalaspis oeselensis

96

100

(Nasal capsules in anterlolateral corners of orbit)

0 (no)

- (inapplic.)

Although olfactory bulbs are present, discrete nasal capsules appear to be absent in osteostracans.

Procephalaspis oeselensis

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Psarolepis romeri

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Psarolepis romeri

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Psarolepis romeri

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Pterichthyodes milleri

247

253

(Preorbital recess)

1 (present)

0 (restricted to premedian plate)

Character is not applicable to taxa without an upper lip/premedian area.

Pterichthyodes milleri

248

252

(Preorbital recess)

0 (restricted to premedian plate)

1 (present)

Character is not applicable to taxa without an upper lip/premedian area.

Remigolepis walkeri

247

253

(Preorbital recess)

1 (present)

0 (restricted to premedian plate)

Character is not applicable to taxa without an upper lip/premedian area.

Remigolepis walkeri

248

252

(Preorbital recess)

0 (restricted to premedian plate)

1 (present)

Character is not applicable to taxa without an upper lip/premedian area.

Romundina stellina

48

51

(DFC86 Sinus superior)

1 (present)

0 (absent or indistinguishable from union of anterior and posterior canals with saccular chamber)

Updated based on Dupret et al. (2017, PLOS ONE)

Romundina stellina

131

135

(DFC55 Median dermal bone of palate (parasphenoid))

1 (present)

0 (absent)

There is no concrete evidence of a parasphenoid in Romundina. Dupret et al. (2017) identified some thickened perichondrium in the hypophyseal region, but stop short of identifying it as a parasphenoid.

Romundina stellina

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Romundina stellina

248

252

(Preorbital recess)

- (inapplic.)

0 (absent)

Character is not applicable to taxa without an upper lip/premedian area.

Shuyu zhejiangensis

48

51

(DFC86 Sinus superior)

0 (absent or indistinguishable from union of anterior and posterior canals with saccular chamber)

1 (present)

Gai et al. (2011) supplementary movie shows what is a distinct sinus superior, contra Zhu et al. (2021. Curr. Biol.)

Styloichthys changae

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Styloichthys changae

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Styloichthys changae

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Tremataspis mammillata

96

100

(Nasal capsules in anterlolateral corners of orbit)

0 (no)

- (inapplic.)

Although olfactory bulbs are present, discrete nasal capsules appear to be absent in osteostracans.

Tremataspis mammillata

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Waengsjoeaspis excellens

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)

Youngolepis praecursor

246

251

(Preorbital depression)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Youngolepis praecursor

247

252

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Youngolepis praecursor

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Yunnanolepis sp

247

253

(Preorbital recess)

0 (absent)

- (inapplic.)

Character is not applicable to taxa without an upper lip/premedian area.

Yunnanolepis sp

248

252

(Preorbital recess)

- (inapplic.)

0 (absent)

Character is not applicable to taxa without an upper lip/premedian area.

Zenaspis salweyi

422

427

(DFC105 Dermal shoulder girdle composition)

0 (ventral and dorsal (scapular) components)

- (inapplic.)

We code this character as inapplicable for osteostracans because our comparative anatomical model proposes that a dorsal area is not yet present in these taxa (and forms during a dorsal expansion of the pharynx)