Characterization of the placoderm (Gnathostomata) assemblage from the tetrapod-bearing locality of Strud (Belgium, Upper Famennian)

CHARACTERIZATION OF THE PLACODERM

(GNATHOSTOMATA) ASSEMBLAGE FROM THE

TETRAPOD-BEARING LOCALITY OF STRUD

(BELGIUM, UPPER FAMENNIAN)

by S�EBASTIEN OLIVE1,2*, GA€EL CL�EMENT3, EDWARD B. DAESCHLER4 and

VINCENT DUPRET5,6*1OD Earth and Life History, Royal Belgian Institute of Natural Sciences, Rue Vautier 29, 1000, Brussels, Belgium; e-mail: [email protected] & Diversity Dynamics Lab, Department of Geology, Li�ege University, B18, Boulevard du Rectorat, Sart-Tilman, 4000, Li�ege, Belgium3CR2P, UMR 7207 CNRS/MNHN/Paris6 CR2P, CP 38, 57 rue Cuvier, F75231, Paris Cedex 05, France; e-mail: [email protected] of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103, USA; e-mail: [email protected] of Evolution and Development, Department of Organismal Biology, Uppsala University, Norbyv€agen 18A, SE 752 36, Uppsala, Sweden6Current address: Research School of Physics and Engineering, College of Physical and Mathematical Sciences, Building 84, The Australian National University,

Acton, ACT 2601, Australia; e-mail: [email protected]

*Corresponding authors

Typescript received 14 April 2015; accepted in revised form 27 June 2015

Abstract: The placoderm fauna of the late Famennian tet-

rapod-bearing locality of Strud, Belgium, is studied on the

basis of historical and newly collected material. It includes

the previously described antiarch Grossilepis rikiki, the groen-

landaspidid Turrisaspis strudensis sp. nov. and the actinolepi-

doideid Phyllolepis undulata. P. undulata is thoroughly

described and joins the list of the valid Phyllolepis species

confidently diagnosed. A morphometrical analysis performed

on the centronuchal and anterior ventrolateral plates of the

Phyllolepis material demonstrates that there is only one

species of Phyllolepis in Belgium (thus, Phyllolepis konincki

becomes a junior synonym of P. undulata), that P. rossimon-

tina (Pennsylvania) is a synonym of P. undulata and that the

unity of the genus Phyllolepis is strongly supported, although

the characterization of several species within this genus is

blurred. The strong resemblance between the faunal compo-

sitions in Strud and Red Hill (Pennsylvania, USA) suggests

important faunal exchanges between these regions of the

Euramerica landmass.

Key words: Famennian, palaeobiogeography, Phyllolepis,

Red Hill, Strud, Turrisaspis.

S INCE its rediscovery in 2004, the tiny quarry of Strud

(Namur Province, Belgium) has been the subject of

intensive excavation and the driving force for a reconsid-

eration of the Famennian palaeontological record in

Belgium (see Olive et al. 2015a for a summary). The site

has yielded a very abundant flora (Table 1; Prestianni

et al. 2007) but also an exquisitely preserved invertebrate

fauna (Table 1) including one of the oldest putative

insects (Garrouste et al. 2012, 2013; H€ornschemeyer et al.

2013) and continental crustaceans (Gueriau et al. 2014a,

b; Lagebro et al. 2015). The vertebrate fauna from Strud

(Table 1) consists of sarcopterygian taxa (Lohest 1888a,

b; Leriche 1931; Cl�ement et al. 2004; Cl�ement and

Boisvert 2006), as well as yet undescribed acanthodian

and actinopterygian taxa. The placoderm fauna has been

partly described recently, with the antiarch Grossilepis

rikiki Olive, in press. Phyllolepid placoderms have been

considered in the past (Lohest 1888b; Leriche 1931), but

need a complete re-evaluation, which was one of the aims

of this article.

Phyllolepid placoderms are easily recognizable by the

presence of a ‘centronuchal’ plate of still-debated

homologies (Dupret and Zhu 2008) and a ridged dermal

ornamentation. Their most basal member, Gavinaspis, is

dated from the Lochkovian of China (Dupret and Zhu

2008). Other phyllolepids are known from the Middle

Devonian of Gondwana with the genera Austrophyllolepis,

Placolepis, Cobandrahlepis, Cowralepis and Yurammia. Late

Devonian (Famennian) phyllolepids from Euramerica all

belong to the genus Phyllolepis (Young 2005a, b, c;

Dupret and Zhu 2008; Long and Daeschler 2013).

The arthrodire groenlandaspidid genus Turrisaspis was

first described by Daeschler et al. (2003) from the late

Famennian Catskill Formation at the Red Hill site in

Pennsylvania, USA. They justified the erection of a new

genus based on the high shape of the median dorsal plate

© The Palaeontological Association doi: 10.1111/pala.12190 981

[Palaeontology, Vol. 58, Part 6, 2015, pp. 981–1002]

and the tall and narrow plates of the lateral trunk shield.

To date, it has been considered to be endemic to Pennsyl-

vania. A short history of the family Groenlandaspididae is

available in Olive et al. (2015a).

Here, we present the placoderm assemblage found in the

Famennian tetrapod-bearing locality of Strud. The nomen-

clatural issue of Belgian Phyllolepis species is resolved and

the species definition within this genus questioned. A new

species of the genus Turrisaspis is recognized in Strud.

Institutional abbreviations. ANSP, Academy of Natural Sciences

of Philadelphia, Pennsylvania, USA; IRSNB, Institut royal des

Sciences naturelles de Belgique, Brussels, Belgium; NHMD, Nat-

ural History Museum of Denmark, Copenhagen, Denmark;

PALULG, Palaeontological collections of the Universit�e de Li�ege,

Li�ege, Belgium; UCL, Universit�e Catholique de Louvain-

la-Neuve, Louvain-la-Neuve, Belgium.

Anatomical abbreviations. ADL, anterior dorsolateral plate; AL,

anterior lateral plate; AMV, anterior median ventral plate; APi,

anterior pineal plate; AVL, anterior ventrolateral plate; bhy, basi-

hyal; CeNu, centronuchal plate; chy, ceratohyal; IL, interolateral

plate; MD, median dorsal plate; PDL, posterior dorsolateral plate;

PMV, posterior medioventral plate; PN, postnasal plate; PNu,

paranuchal plate; PrO, preorbital plate; Psph, parasphenoid;

PtO, postorbital plate; PVL, posterior ventrolateral plate; Sp,

spinal plate.

GEOLOGICAL SETTING

The stratigraphical section at Strud belongs to the Haine–Sambre–Meuse Overturned Thrust Sheets (HSM OTS),

which is a Variscan structural element of the Namur–Dinant Basin (Denayer et al. in press; Fig. 1A). During

the late Famennian times, this region was located along

the south-eastern margin of Euramerica, and in particular

in the London–Brabant Peninsula (Ziegler 1990). The

observed facies in the HSM OTS are typically continental

(Thorez et al. 2006). The Strud section is composed of a

TABLE 1 . Strud flora and fauna.

Vertebrata

Placodermi

Arthrodira

Phyllolepididae

Phyllolepis undulata

Groenlandaspididae

Turrisaspis strudensis

Antiarcha

Bothriolepididae

Grossilepis rikiki

Actinopterygii

Actinopterygii indet.

Sarcopterygii

Dipnomorpha

Dipnoi

Soederberghia cf. S. groenlandica

Jarvikia sp.

Dipteridae indet.

Porolepiformes

Holoptychius sp.

Tetrapodomorpha

‘Osteolepiformes’

Litoptychius-like

Tristichopteridae indet.

Cosmine-covered ‘osteolepididae’

Tetrapoda

Ichthyostega-like tetrapod

Acanthodii

Acanthodii indet.

Arthropoda

Hexapoda

Insecta

Strudiella devonica

Crustacea

Malacostraca

Schramidontus labasensis

Tealliocaris walloniensis

Branchiopoda

Strudops goldenbergi

Anostraca indet.

Conchostraca indet.

Chelicerata

Merostomata

Eurypterida indet.

Plantae

Tracheophyta

Zosterophyllopsida

Barinophytales

Barinophyton citrulliforme

Filicopsida

Zygopteridales

Rhacophyton condrusorum

Progymnospermopsida

Archaeopteridales

Archaeopteris halliana

(continued)

TABLE 1 . (Continued)

Gymnospermopsida

Condrusia rumex

Pseudosporogonites hallei

Moresnetia zalesskyi

Incertae sedis

indet. microsporangia

Sphenopteris flaccida

Sphenopteris modavensis

Vertebrata: Cl�ement et al. (2004), Cl�ement and Boisvert (2006),

Cl�ement and Prestianni (2009), Olive (in press). Arthropoda:

Garrouste et al. (2012), Gueriau et al. (2014a, b), Lagebro et al.

(2015). Plantae: Prestianni et al. (2007), Cl�ement and Prestianni

(2009), Denayer et al. (in press).

982 PALAEONTOLOGY , VOLUME 58

road-cut exposure and of two disused quarries. The

smallest quarry (the northern one) was exploited during

the second half of the nineteenth century. It had been

abandoned and subsequently rediscovered in 2004, thanks

to geologists from Li�ege University (Laurent Barchy and

Jean-Marc Marion). In this quarry, the beds are in reverse

succession and the dipping is 80° southward. The fossilif-

erous strata of this quarry (lithological unit UL7) belong

to the Upper Devonian Evieux Formation (Denayer et al.

in press; Fig. 1B, C). They were first considered to be

upper middle Famennian (GF Biozone, Prestianni et al.

2007) because of the absence of some typical palynologi-

cal markers. However, the plant association and the

lithostratigraphy, as well as a revision of the palynological

markers, argue preferentially for a late Famennian age

(VCo ‘rad’ zone, Denayer et al. in press). The sedimentol-

ogy of the lithological unit 7 corresponds to the filling

sequence of a channel in an alluvial plain (Garrouste

et al. 2012; Denayer et al. in press). Placoderm remains

are found throughout the lithological unit UL7, except in

its uppermost part (Fig. 1C). The top of the sequence

corresponds to the final phase of the channel filling. This

corresponds to fresh or brackish water ponds seasonally

dried and flooded, which permitted the preservation of

delicate invertebrates and small actinopterygian remains

unpreserved in the rest of the sequence. Heavier fossils,

such as placoderm remains, are found in the immediately

underlying beds.

The placoderm material discussed herein comes from

Strud, except for a few specimens from Modave, Ch�evre-

mont and Evieux. All these localities belong to the Mont-

fort/Evieux Formation, Famennian. Modave and Evieux

belong to the Dinant synclinorium, whereas Ch�evremont

belongs to the Vesdre area, both being Variscan (Hercy-

nian) structural elements (Fig. 1A). The fossil material

consists of disarticulated head and thoracic elements,

except for one phyllolepid specimen partly preserved in

articulation (PALULG.2014.01.29.16 from Ch�evremont).

MATERIAL AND METHODS

The material used in the morphometric analysis includes

specimens from various international collections. The

material from the ANSP palaeontological collections and

the material from Belgian institutions have been directly

measured, and the rest of the material has been measured

based on figures in the literature (including a few lost

Belgian specimens used by Lohest (1888a) and Leriche

(1931)). Measurement methods are those of Daeschler

et al. (2003, fig. 10) for median dorsal plates of Turrisas-

pis, and those of Figure 2 for anterior ventrolateral and

centronuchal plates of Phyllolepis. As noticed by Young

(2005b, p. 209), Stensi€o (1939, p. 7) did not clearly

defined the anterior ventrolateral plate measurements he

used. Young (2005b, c) clearly defined some of those

measurements but did not take into account all features

used by Stensi€o such as the lengths of the medial or of

the posterior margins for instance. Therefore, we decided

to redefine the measurements used by Stensi€o (1939).

Measurements are given in Olive et al. (2015b, appendices

1–2). Material recently collected has been used in this

analysis as well as Lohest’s and Leriche’s figured and mea-

surable material. Not all measurements were possible

depending on plate preservation (see Olive et al. 2015b,

appendices 1–2) ‘NM’ for nonmeasurable). Thus, sample

size may vary from one graphic to another.

SYSTEMATIC PALAEONTOLOGY

This published work and the nomenclatural act it contains have

been registered in ZooBank: http://zoobank.org/References/

835B725E-7824-442D-B283-2CBC614EE92C

Order ARTHRODIRA Woodward, 1891

Suborder PHLYCTAENIOIDEI Miles, 1973

Infraorder PHLYCTAENII Miles, 1973

Family GROENLANDASPIDIDAE Obruchev, 1964

Genus TURRISASPIS Daeschler et al., 2003

Type species. Turrisaspis elektor Daeschler et al., 2003. Red Hill,

Clinton County, Pennsylvania, USA, Duncannon Member, Cats-

kill Formation, late Famennian (Fa2c substage), Late Devonian.

Turrisaspis strudensis sp. nov.

Figure 3

1965 Tiaraspis sp.; Gross, abb. 4A, taf. 2, figs 2–3

(median dorsal plate PALULG A/2005)

2005 Groenlandaspis thorezi (partim); Janvier and

Cl�ement, pl. 2C (median dorsal plate PALULG

A/2005, the same median dorsal plate that was

attributed to Tiaraspis sp. by Gross (1965)).

LSID. urn:lsid:zoobank.org:act:D4668CEA-9F3C-4BA8-8769-

C45B9579C6E1

Derivation of name. In reference to the locality of Strud, where

the material was found.

Holotype. IRSNB P.9447, a posterior dorsolateral plate (Strud).

Other material. Strud: APi: IRSNB P.9448; MD: IRSNB P.9449,

IRSNB P.9450, IRSNB vert 32.438-001, PALULG A/2005,

UCL.P.V.L.10.532; AL: IRSNB P.9451–9452, IRSNB vert 32.220-

OL IVE ET AL . : P LACODERMS FROM STRUD 983

http://zoobank.org/References/835B725E-7824-442D-B283-2CBC614EE92C

http://zoobank.org/References/835B725E-7824-442D-B283-2CBC614EE92C

A

B

C


001–002, IRSNB vert 32.438-002; AVL: IRSNB P.9453–9454,IRSNB vert 32.164-003, IRSNB vert 32.220-003–004, IRSNB vert

32.745-001; PMV: IRSNB P.9455; PVL: IRSNB P.9456–9457,IRSNB vert 32.745-002; Sp: IRSNB P.9458, IRSNB vert 32.438-

003.

Modave: MD: PALULG A/2005.

Diagnosis. Turrisaspis species with 90° angle of the bend

in the main lateral sensory line groove on the PDL;

overlapping area of the PDL for the ADL posteriorly long;

PVL with a large posterior part of the ventral lamina; Sp

with a large anteriorly directed spinelet at the middle of

the lateral margin.

Localities and horizons. Strud, Haltinne District, Gesves Com-

mune, Namur Province, Belgium; Evieux Formation (Denayer

et al. in press), VCo ‘rad’ zone, late Famennian, Upper Devo-

nian.

Modave, Li�ege Province, Belgium; upper part of the Evieux

Formation, Famennian, Upper Devonian.

Morphological description of the skull roof

Anterior pineal plate. (APi, Fig. 3A) The anterior pineal plate is

a trapezoidal plate with the posterior margin concave and

shorter than the anterior one. The pineal foramen is located at

the geometric centre of the plate. The ornamentation consists of

numerous closely spaced, small tubercles.

Morphological description of the thoracic armour

Median dorsal plate. (MD, Fig. 3B–E) The median dorsal plate

is very high and narrow. The tip of the plate is pointed and

forms an angle varying between 25° (specimen IRSNB P.9449,

Fig. 3C) and 39° (largest specimen UCL.P.V.L.10.532, Fig. 3B)

depending on the specimen. Three of four median dorsal plates

(IRSNB P.9449, P.9450 and PALULG A/2005, Fig 3C–E) are

roughly the same size, and presumably of the same ontogenetic

stage but differ in the tip angle. It is thus unclear whether a rela-

tionship between the size of the plate and tip angle exists. How-

ever, it seems by comparison with the largest median dorsal

plate (UCL.P.V.L.10.532, Fig. 3B) that the angle opened with

growth, as observed in Groenlandaspis riniensis (Long et al. 1997,

p. 258). The posterior edge of the plate displays a gentle slope

and is covered with large, pointed spinelets whose apices are of

a different colour to the plate itself (i.e. darker; on

UCL.P.V.L.10.532 only, Fig. 3B), suggesting that it is likely to be

a different type of tissue. The same observation is made for the

tips of several tubercles covering the same MD. The anteroven-

tral projection of the median dorsal plate is much narrower than

the posterior one. The tip of the plate can also bear spinelets, as

the posterior edge does (see specimen PALULG A/2005, Fig. 3E).

The anterior margin of the median dorsal plate is straight or

slightly concave and devoid of spinelets. The ventral margin of

the plate is deeply indented by overlap areas of the anterior and

posterior dorsolateral plates. The ornamentation of the sides of

the median dorsal plate consists of small tubercles organized in

sinuous rows parallel to the ventral margin.

A

B

F IG . 2 . Measurement methods for the Phyllolepis plates. A,

CeNu. B, AVL. Abbreviations: L, length; L.ant, anterior length;

L.mg.ant, anterior margin length; L.mg.lat, lateral margin length;

L.mg.med, medial margin length; L.mg.post, posterior margin

length; W, width.

F IG . 1 . Location, geological and stratigraphical framework of the Strud locality and location and geological framework of the Mod-

ave, Evieux and Ch�evremont localities. A, geological map of southern Belgium with the location of the Strud, Modave, Evieux and

Ch�evremont localities. B, lithological and lithostratigraphical column of the Strud locality. C, schematic block diagram of the Strud

channel with the location of placoderm remains. Abbreviations: HSM OTS, Haine–Sambre–Meuse Overturned Thrust Sheets; Lux, Lux-

emburg; UL, lithological unit. Modified after Denayer et al. (in press). Colour online.


A

F

K L O

Q

R

NM

G H I

J

P

B C D E

F IG . 3 . Turrisaspis strudensis sp. nov. A–D, F–Q, from Strud, and E, from Modave. A, APi in external and internal views,

IRSNB P.9448a. B, MD in left external view, UCL.P.V.L.10.532. C, MD in left external view, IRSNB P.9449a. D, MD in left

external view, IRSNB P.9450a. E, MD in right external view, PALULG A/2005. F, right PDL in external view, IRSNB P.9447a.

G, right PDL in external view, IRSNB P.9447b. H, right PDL in external view, IRSNB P.9447, interpretative drawing. I, right AL

in external view, IRSNB P.9451. J, left AL in external view, IRSNB P.9452a. K, left AVL in external and internal views, IRSNB

P.9453a. L, left AVL in external and internal views, IRSNB P.9454. M, PMV in external view, IRSNB P.9455a. N, left PVL in

external view, IRSNB P.9457a. O, left PVL in external view, IRSNB P.9456a. P, proximal part of a right Sp in ventral view,

IRSNB P.9458a. Q, distal part of a right Sp in dorsal view, IRSNB P.9458b. R, cross section of a right Sp, IRSNB P.9458a,

indicated by a double arrow on P. Dashed line for assumptions; dashed line plus dots for information provided by counterparts.

Abbreviations: a.b, annular bourrelet/thickening; ad.cr, anterodorsal crest of anterior lateral plate; av.cr, anteroventral crest of

anterior lateral plate; av.s, anteroventral sulcus; emb.ADL, embayment of median dorsal plate for anterior dorsolateral plate;

emb.PDL, embayment of median dorsal plate for posterior dorsolateral plate; f.pi, pineal foramen; gr.Sp, ventral groove of spinal

plate; ins.Sp, insertion area for spinal plate; lat, laterally; oa.ADL, overlap area for anterior dorsolateral plate; oa.AL, overlap area

for anterior lateral plate; oa.AMV, overlap area for anterior medioventral plate; oa.AVL, overlap area for anterior ventrolateral

plate; oa.IL, overlap area for interolateral plate; oa.MD, overlap area for median dorsal plate; oa.PMV, overlap area for posterior

medioventral plate; oa.PVL, overlap area for posterior ventrolateral plate; obs.l, obstantic lamina of anterior lateral plate; pbr.l,

postbranchial lamina of anterior lateral plate; pd.cr, posterodorsal crest of anterior lateral plate; pec.n, pectoral notch. Scale bar

represents 1 cm. Colour online.


Posterior dorsolateral plate. (PDL, Fig. 3F–H) The only available

posterior dorsolateral plate (part and counterpart) is slightly

deformed but appears narrower than the material from Pennsyl-

vania (although this could be due to an earlier ontogenetic

stage). The posterodorsal process of the ornamented area is pro-

nounced and angles upward, forming an apron and thus resem-

bling the condition observed in the material of Groenlandaspis

potyi (see Olive et al. 2015a, fig. 2) as well as that of Turrisaspis

elektor from Pennsylvania (although to a lesser extent; see

Daeschler et al. 2003, fig. 9E). The posterior edge of the over-

lap area for the MD plate is situated anteriorly to the posterior

edge of the plate, while it is at the same level in T. elektor.

Below this apron, the posterior edge of the plate is convex

(while concave in T. elektor). The area overlapped by the med-

ian dorsal plate is high and well preserved on the counterpart

and does not reach the posterior margin (i.e. the apron is free).

The main lateral sensory line groove forms an angle of exactly

90° on the holotype. The area overlapped by the anterior

dorsolateral plate extends more sharply and posteriorly than

in T. elektor. The ornamentation consists of small, evenly

distributed tubercles.

Anterior lateral plate. (AL, Fig. 3I–J) The shape of the AL seems

to vary during ontogeny. The anterodorsal edge is rounder on the

smallest specimen than on the largest plate; this ensures a slightly

convex aspect of the dorsal edge of the smallest plate, while the

largest one displays a straighter dorsal edge. The anteroventral

and posterodorsal crests are sharper on the largest plate. The

ventral edge is straight and longer than the dorsal edge. The

posterior margin is concave, and the posterodorsal corner of the

plate is the most posterior part of the plate. The ornamentation

consists of numerous small tubercles distributed randomly.

Anterior ventrolateral plate. (AVL, Fig. 3K–L) The AVL is the

most frequently found plate and the best preserved. The

V-shaped depression is well defined and opens on the pectoral

notch, which forms a large embayment. The ornamentation of

the AVL consists mostly of small scattered tubercles.

Posterior median ventral plate. (PMV, Fig. 3M) The PMV is

enlarged in its posterior part with wide overlapped areas for all

neighbouring plates. The exposed part of the plate is elongate

and covered with small tubercles.

Posterior ventrolateral plate. (PVL, Fig. 3N–O) The ventral lam-

ina of the posterior ventrolateral is elongated. On the cranial

margin, the area overlapped by the anterior ventrolateral plate

is crescent-shaped and quite long. The plate presents the typi-

cal sigmoid flexure and mutual overlap seen in the Phlyctaenii

(except for the family Phlyctaenididae): the right plate over-

laps the left one, while the latter protrudes a short process

overlapping the right plate. This sigmoid flexure is sharper

and simpler (one process) on T. strudensis than it is on

T. elektor where it is less pronounced (see Daeschler et al.

2003, fig. 9R). The angle between the ventral and lateral lami-

nae cannot be measured because of the strong compression of

the specimen. The ventral surface displays a well-marked

annular bourrelet of Goujet (1984) circling the posterior ring

of the thoracic armour. The dermal ornament consists of scat-

tered tubercles.

Spinal plate. (Sp, Fig. 3P–R) IRSNB P.9458 is broken into two

parts, the posterior one corresponding roughly to the ‘free’ part

of the plate and the anterior one contacting the AVL. The latter

displays two types of ornamentation on the lateral side, divided

by a large anteriorly pointed spinelet at mid-length. Anterior to

the large spinelet, the visible ornamentation consists of three

rows of large tubercles; past the spinelet, two rows remain with

smaller and spiniform tubercles. The free posterior medial mar-

gin displays medially oriented spinelets larger and more widely

spaced than the lateral ones, as in Turrisaspis elektor. The rest of

the plate is smooth. In cross section, the angle made by the

asymptotic lines of the ventral and dorsal laminae is 30°(Fig. 3R). The continuation of the anteroventral sulcus (of the

interolateral plate) is visible on the ventral side of the spinal

plate (see Goujet 1984; Dupret 2003, 2010).

Remarks

The material is referred to the genus Turrisaspis based on

the high and narrow median dorsal and posterior dorso-

lateral plate, as well as a short posterior projection

(‘apron’) of the posterior dorsolateral plate which does

not allow the overlap area for the median dorsal plate to

reach the posterior corner of the plate, a feature shared

by Turrisaspis and Africanaspis (Long et al. 1997). The

attribution to Africanaspis is not considered because the

typical lateral ridge on the posterior dorsolateral plate is

not present in the plate from Strud. Moreover, in Africa-

naspis the dorsal process of its posterior dorsolateral plate

is much lower than that of T. strudensis.

Several characteristics of T. strudensis differ from the

description of T. elektor: (1) the angle of the sensory line

groove on the posterior dorsolateral plate being 90° in

T. strudensis rather than 70° in T. elektor; (2) the over-

lapping area of the posterior dorsolateral plate for the

anterior dorsolateral plate is longer posteriorly in T. stru-

densis; (3) the spines on the posterior margin of the

median dorsal plate are more widely spaced and not as

numerous in the Belgian specimens; (4) the posterior part

of the ventral lamina of the posterior ventrolateral plate is

larger in T. strudensis; and (5) the mainly smooth spinal

plate of T. strudensis displays larger spinelets on the free

posterior medial margin and with a very large spinelet at

the middle of the lateral margin. Although point 3 may be

due to individual variation within T. elektor according to

Daeschler et al. (2003, p. 56: ‘individual variation is also

evident in the median dorsal plate sample with respect to

details of the serrations along the caudal margin’), we

assert here that the other features diverging from the type

species of the genus Turrisaspis are relevant and numerous

enough to justify the erection of a new species.


Olive et al. (2015a) questioned the attribution of the

median dorsal plate published by Gross (1965, abb. 4A)

(Fig. 3E) to Tiaraspis, and later redescribed by Janvier and

Cl�ement (2005) as Groenlandaspis thorezi. Olive et al.

(2015a) did not agree with that assumption because: (1) ‘the

specimens do not match with each other’; and (2) they ‘con-

cur with Gross (1965) who recognized the ventral lamina as

an overlap area’. The newly described material of Turrisaspis

from Strud allows the attribution of this problematic median

dorsal plate to this taxon, based on the numerous shared fea-

tures (i.e. high and narrow shape, posterior margin with a

gentle posteroventral slope and with pointed spines, ventral

margin deeply indented by the posterior dorsolateral plate,

and ornamentation made of small tubercles arranged in

sinuous rows parallel to the ventral margin).

Suborder ACTINOLEPIDOIDEI Miles, 1973

Infraorder PHYLLOLEPIDIDA Stensi€o, 1934

Family PHYLLOLEPIDIDAE Woodward, 1891

Genus PHYLLOLEPIS Agassiz, 1844

Type species. Phyllolepis concentrica Agassiz, 1844. Perthshire and

Rosebrae beds, Morayshire, Scotland, Famennian.

Phyllolepis undulata Lohest, 1888b

Figures 4–7

1888b Phyllolepis undulatus Lohest, p. 48.

1888b Phyllolepis corneti Lohest, p. 48.

1888b Pentagonolepis konincki Lohest, p. 52.

1931 Phyllolepis undulata; Leriche, p. 9.

1939 Phyllolepis konincki; Stensi€o, p. 19.

2005 Phyllolepis rossimontina Lane and Cuffey.

2013 Phyllolepis konicki [sic]; Long and Daeschler, fig. 6.

2013 Phyllolepis undiulata [sic]; Long and Daeschler, fig. 6.

Type specimen. PALULG.2014.04.08.10, median dorsal plate, lec-

totype defined by Leriche (1931, pl. 1, figs 3, 3a) and figured

before by Lohest (1888b, pl. 10, fig. 5).

Other material. Strud: PrO: IRSNB P.9459; PN: IRSNB P.9460,

IRSNB vert 32.745-003–004; PtO: IRSNB P.9461, IRSNB vert

32.220-005; PNu: IRSNB P.9462–9463, IRSNB vert 31.264-001–005, IRSNB vert 31.913-003–005, IRSNB vert 32.164-004, IRSNB

vert 32.164-018, IRSNB vert 32.220-006–011, IRSNB vert 32.438-

004–009, IRSNB vert 32.745-005; CeNu: IRSNB P.9464–9468,IRSNB vert 31.264-006–015, IRSNB vert 31.595-001, IRSNB vert

31.913-006, IRSNB vert 32.164-005–007, IRSNB vert 32.220-012–021, IRSNB vert 32.438-010–014, IRSNB vert 32.745-006,

PALULG.2014.04.08.19, PALULG.2014.04.08.20; bhy, chy & Psph:

IRSNB P.9469; MD: IRSNB P.9470–9473, IRSNB vert 31.264-016–030, IRSNB vert 31.595-002–009, IRSNB vert 31.913-007–008,IRSNB vert 32.048-001, IRSNB vert 32.164-008–009, IRSNB vert

32.220-022–031, IRSNB vert 32.438-015–023, IRSNB vert 32.745-

007–008, PALULG.2014.04.08.11, PALULG.2014.04.08.17; ADL:

IRSNB P.9474, IRSNB vert 31.913-009, IRSNB vert 32.164-010,

IRSNB vert 32.220-032–034, IRSNB vert 32.438-024–026, IRSNBvert 32.745-009; AL: IRSNB P.9475; AVL: IRSNB P.9476–9482,IRSNB vert 31.264-031–049, IRSNB vert 31.595-010–011, IRSNBvert 31.913-010–015, IRSNB vert 32.048-002, IRSNB vert 32.164-

011–013, IRSNB vert 32.220-035–036, IRSNB vert 32.438-027–032, PALULG.2014.04.08.16, PALULG.2014.04.08.15, PALULG.

2014.04.08.18; PVL: IRSNB P.9483–9484, IRSNB vert 31.264-050–056, IRSNB vert 31.595-012–013, IRSNB vert 32.048-003, IRSNB

vert 32.164-014–017, IRSNB vert 32.220-037–041, IRSNB vert

32.438-033–040.

Ch�evremont: Articulated specimen: PALULG.2014.01.29.16;

CeNu: PALULG.2014.04.08.20; MD: PALULG.2014.04.08.12; AVL:

PALULG.2013.05.3.11, PALULG.2014.01.29.19; PVL: PALULG.20

14.01.29.17, PALULG.2014.01.29.18.

Evieux: MD: PALULG.2014.04.08.13; AVL: PALULG.2014.04.

08.14.

Red Hill: See Lane and Cuffey (2005), Long and Daeschler

(2013) and Olive et al. (2015b, appendices 1–2).

Tioga site: CeNu: ANSP 23360 (Olive et al. 2015b,

appendix 1).

F IG . 4 . Phyllolepis undulata, from Strud. A, right PrO in external view, IRSNB P.9459. B, left PtO in external view, IRSNB P.9461a.

C, right PN in internal view, IRSNB P.9460. D, right PNu in external view, IRSNB P.9462. E, PNu in external and internal views,

IRSNB P.9463a. F, CeNu in external and internal views, IRSNB P.9467. G, CeNu in external view, IRSNB P.9466a. H, CeNu in exter-

nal view, IRSNB P.9464. I, CeNu in external and internal views, IRSNB P.9465a. J, CeNu in internal view, IRSNB P.9468. K, MD in

external view, IRSNB P.9470. L, MD in external and internal views, IRSNB P.9471. M, MD in internal view, IRSNB P.9472. N, MD in

internal view, IRSNB P.9473. O, left ADL in external view, IRSNB P.9474a. P, right AL in ventral view, IRSNB P.9475. Q, right PVL

in external and internal views, IRSNB P.9483. R, left PVL in internal view, IRSNB P.9484. Abbreviations: ab.at, abdominal muscle

attachment of median dorsal plate; bu, central bulge of centronuchal plate; csc, central sensory canal; de, central depression on cen-

tronuchal plate visceral surface; dsc, dorsal sensory canal; ifc, infra-orbital canal; ins.Sp, insertion area for spinal plate; lc, main lateral

line; oa.ADL, overlap area for anterior dorsolateral plate; oa.AL, overlap area for anterior lateral plate; oa.CeNu, overlap area for cen-

tronuchal plate; oa.MD, overlap area for median dorsal plate; oa.PDL, overlap area for posterior dorsolateral plate; oa.PNu, overlap

area for paranuchal plate; oc.os, occipital ossification attachment of the centronuchal plate; pbr.l, postbranchial lamina of anterior lat-

eral plate; ppl, posterior pit line; soc, supraorbital canal; syn.at, synarcual attachment of the median dorsal plate; vsc, V-shaped canal

of postnasal plate. Scale bars represent 1 cm. Colour online.


Diagnosis. This is the first formal diagnosis for this spe-

cies, as diagnoses were not compulsory when it was first

named and described in 1888.

V-shaped sensory line canal of the postnasal plate

reaching the centronuchal plate; centronuchal plate shape

displaying a variability stretching from longer than broad

to broader than long (shape variation continuous);

anterolateral margin of the centronuchal plate either well

defined or included in the convex anterior margin which

may be sinusoidal; external surface of the centronuchal

plate bulged in the middle; visceral surface of the cen-

tronuchal plate with a tiny depression in the centre and

with a longitudinal median groove running from the cen-

tre to the posterior margin and corresponding to the

occipital ossification; basihyal elongate and oak-leaf-

shaped; visceral surface of the median dorsal plate with

A

F

K

L

O Q R

NM

G H I

J

P

B C D E

OLIVE ET AL . : P LACODERMS FROM STRUD 989

an anteroposterior ridge corresponding anteriorly to the

synarcual attachment area; anterior ventrolateral plate

slightly longer than broad in the majority of cases; ante-

rior ventrolateral plate following an isometric growth;

ornamentation made of concentric parallel lines on all

plates and of a largely meshwork reticulated pattern in

the anterior part of the centronuchal plate.

Localities and horizons. Strud, Haltinne, Gesves, Namur Pro-

vince, Belgium, Evieux Formation (Denayer et al. in press), VCo

‘rad’ zone, late Famennian, Upper Devonian.

Evieux, Esneux, Li�ege Province, Belgium, Montfort/Evieux

Formation, Famennian, Upper Devonian.

Ch�evremont, Li�ege Province, Belgium, Montfort/Evieux For-

mation, Famennian, Upper Devonian.

Red Hill, road cut on route 120, 1 mile west of Hyner, Clin-

ton County, Pennsylvania, USA, Duncannon Member, Catskill

Formation, late Famennian (Fa2c substage), Late Devonian.

Tioga site, construction zone on route 15, just south of Tioga,

Tioga County, Pennsylvania, USA, Catskill Formation, late

Famennian, Late Devonian.

Morphological description of the skull roof

Preorbital plate. (PrO, Fig. 4A) The only known preorbital plate

is poorly preserved. The supraorbital canal is well marked, and

the overlap area for the centronuchal plate is narrow.

Postnasal plate. (PN, Fig. 4B) The postnasal plate is pentagonal

in shape and displays the typical V-shaped sensory line canal,

which reaches the area overlapped by the centronuchal plate.

Postorbital plate. (PtO, Fig. 4C) The sole specimen of a postor-

bital plate displays the anterior part only. The intersection

between the main lateral line, central sensory and infra-orbital

canals is observable. The areas overlapped by the centronuchal

and paranuchal plates are narrow.

Paranuchal plate. (PNu, Figs 4D–E, 5) The paranuchal plate is

characteristic of the genus Phyllolepis with the anterior extremity

of the main lateral line located at about half-length of the lateral

side and separating a larger area in front of the main lateral

line and a smaller one posteriorly (Long 1984). The postnuchal

process is thin, and the area overlapped by the centronuchal

plate is narrow. No median process (interpreted as a vestigial

endolymphatic duct; see Dupret and Zhu 2008) has been identi-

fied. The ornamentation consists of well-defined concentric

parallel lines.

Centronuchal plate. (CeNu, Figs 4F–J, 5) The centronuchal plate

of P. undulata is slightly broader than long in the majority of

specimens (73%) and sometimes as long as broad (21%). How-

ever, two of the smallest plates of the sample (IRSNB P.9467

and IRSNB vert 32.164-007, Fig. 4F) display a centronuchal plate

longer than broad (6%). The anterolateral corner is either well

defined (Phyllolepis-like) or part of the curved and convex ante-

rior edge (Placolepis-like), which can occasionally be sinusoidal

(see Discussion for further details). A low bulge is consistently

found at the radiation centre in all specimens. The supraorbital

and the central sensory canals are well defined contrary to the

posterior pit lines, which are faint impressions at best or present

in more superficial layers of the dermis. On the smallest individ-

uals, the supraorbital canals and posterior pit lines are consis-

tently invisible, and the central sensory canals are faint. The

visceral surface is smooth except for the shallow central depres-

sion (which corresponds externally to the bulge). Moreover, a

longitudinal median groove runs from this central depression to

the posterior margin as in Phyllolepis orvini (Stensi€o 1934, p. 46),

and may correspond to the occipital ossification attachment

(Ritchie 2005, fig. 13) The ornamentation consists of a large

reticulated pattern in the anterior half and straight ridges, parallel

to posterior margin, in the posterior half.

A B C

F IG . 5 . Phyllolepis undulata, from Ch�evremont. A, partly articulated specimen in external and internal views, PALULG.2014.01.29.16.

B, specimen partly in connection in external and internal views, PALULG.2014.01.29.16, interpretative drawing. C, MD, in external

view, PALULG.2014.04.08.12. Abbreviations: AVL, anterior ventrolateral plate; CeNu, centronuchal plate; dsc, dorsal sensory canal;

MD, median dorsal plate; PNu, paranuchal plate; PVL, posterior ventrolateral plate. Scale bars represent 1 cm. Colour online.


Shape variations of the centronuchal plate are numerous; that

is straight or convex posterolateral margin, anterolateral corner

either well defined (Phyllolepis-like) or part of the curved and con-

vex anterior edge (Placolepis-like). However, the morphometrical

analysis (see below) does not permit discrimination between

different forms and the variations seem continuous. Therefore, all

centronuchal plates are gathered under P. undulata.

Parasphenoid, basihyal and ceratohyals

Three elements of the visceral skeleton are recovered in anatomi-

cal position on the part and counterpart of specimen IRSNB

P.9469 (Fig. 6) and are preserved as extremely flattened, dark

brown bone. The basihyal is on the part, and the parasphenoid

and ceratohyals are on the counterpart. The outlines of the skull

roof are very difficult to discern.

A B

C D

F IG . 6 . Phyllolepis undulata, from Strud. Parasphenoid, basihyal and ceratohyals. A, bhy in ventral view, part, IRSNB P.9469a. B, chy

and Psph in dorsal view, counterpart, IRSNB P.9469b. C, close-up of the bhy in ventral view, part, IRSNB P.9469a. D, three elements in

connection, IRSNB P.9469, interpretative drawing. Abbreviations: bhy, basihyal; chy, ceratohyal; m.ri, median ridge of basihyal; o.g, obli-

que groove of basihyal; Psph, parasphenoid. Scale bars represent: 1 cm (A–B, D); 2 mm (C). Colour online.


The parasphenoid and ceratohyals are preserved as three

small discs; the parasphenoid is approximately four times

bigger than each ceratohyal. The basihyal shows a complex

outline, oak-leaf-like; the anterior tip is slender, while the

posterior part is broad. The only visible reliefs on the basihyal

consist in a median ridge in the anterior half, and an oblique

groove just posterior to it on the right side. The four

elements connect to each other when print and counterprint

are assembled.

Morphological description of the thoracic armour

Median dorsal plate. (MD, Figs 4K–N, 5) The median dorsal

plate is slightly broader than long in P. undulata. Only IRSNB

P.9471 (Fig. 4L) and PALULG.2014.04.08.12 (Fig. 5C) display

respectively the left or the right dorsal sensory line groove run-

ning from the anterolateral corner to the centre of the plate. The

visceral surface exhibits a longitudinal ridge surrounded by a

pair of shallow triangular depressions, forming a hourglass shape

with the thinner part in the middle or in the anterior half of the

plate (as figured by Stensi€o 1934, pl. 10). The anterior portion

of the hourglass shape most likely corresponds to the synarcual

attachment area, and the posterior portion may correspond to

abdominal muscle attachment. The ornamentation consists of

well-defined concentric parallel lines.

Anterior dorsolateral plate. (ADL, Fig. 4O) The anterior dorso-

lateral plate is only known by its smooth visceral side. The over-

lap area for the median dorsal plate is broader than that for the

anterior lateral plate; the overlap area for the paranuchal plate is

narrow.

Anterior lateral plate. (AL, Fig. 4P) The anterior lateral plate is

known from a single specimen. It is quadrangular in shape,

longer than high. The lateral margin is slightly convex, and the

posterior, lateral and anterior margins are straight.

Anterior ventrolateral plate. (AVL, Figs 5A–B, 7) The anterior

ventrolateral plate is slightly longer than broad in the majority

of specimens (87%) and sometimes as long as broad (9%) or

slightly broader than long (4%). The antero-mesial angle

(sensu Young 2005c, fig. 2B) is protruding anteriorly; this

corner can be straight or slightly rounded, implying a limited

contact with the minute anterior median ventral plate (if

present). The area overlapping the posterior ventrolateral plate

is very narrow (Fig. 7E). Growth series are available thanks to

the great number of specimens. The anterior ventrolateral

plate follows an isometric growth (Fig. 8A). The ornamenta-

tion consists of well-defined parallel lines, more widely spaced

anteriorly.

Posterior ventrolateral plate. (PVL, Figs 4Q–R, 5A–B) The poste-rior ventrolateral plate is triangular in shape. The antero-mesial

angle varies between 60 and 70°. The area overlapped by the

anterior ventrolateral plate is narrow mesially and slightly wider

laterally. The ornamentation consists of well-defined parallel lines.

Remarks

Concerning the paranuchal plates, IRSNB P.9463

(Fig. 4E) is very different from all other paranuchal plates

found in Strud (IRSNB P.9462 illustrating the most com-

mon shape; Fig. 4D). IRSNB P.9463 displays a straight

lateral margin and the portion of the plate anterior to the

lateral line canal is shorter than the posterior portion, a

condition absent in other paranuchal plates of Strud and

in other Phyllolepididae species (Long 1984, fig. 2). The

species with a paranuchal plate that is the most similar to

IRSNB P.9463 is Phyllolepis woodwardi, which shares a

similar posterolateral angle and posterior and posterolat-

eral straight edges, but the portion of the plate anterior

to the lateral line canal is much longer in P. woodwardi

(Stensi€o 1939; but for skull roof and paranuchal plate

comparisons, see Long 1984, figs 2A, 25; Ritchie 2005, fig.

20) than in P. undulata. As it is the largest of all phyl-

lolepid plates of the locality, IRSNB P.9463 may corre-

spond to an adult specimen. Whatever the case, as the

sample is small and as the variations could be due to

intraspecific variation, this plate is placed in P. undulata

for convenience.

Long and Daeschler (2013, p. 37) advocated ‘that the

identification of phyllolepid placoderms from disarticu-

lated material is only possible if a large associated

assemblage is known, as for Placolepis from Braidwood,

Australia (Ritchie 1984) or Phyllolepis species from East

Greenland (Stensi€o 1934, 1936, 1939)’. This is now the

case for P. undulata of which most plates of the trunk

shield are known. Long and Daeschler (2013) added

‘the identification of genera from isolated plates

remains uncertain based only on plate shape and

ornamentation unless one of the diagnostic plates is

present (marginal or paranuchal)’. Even if no marginal

plates of P. undulata are currently recognized, the para-

nuchal plates are numerous and correspond (except one

doubtful plate, see above) to Phyllolepis (paranuchal

with the anterior extremity of the main lateral line

located at about half-length of the lateral side (Long

1984)).

Long and Daeschler (2013, p. 37) listed the valid Phyl-

lolepis species that can be confidently diagnosed (i.e.

P. woodwardi, P. orvini, P. nielseni, P. rossimontina (here

put in synonymy with P. undulata) and P. thomsoni).

P. undulata can be added to this list.

DISCUSSION

How many Phyllolepis species in Belgium?

For the material found in Belgium, Lohest (1888b)

erected the genus Pentagonolepis and two new species of


Phyllolepis: P. undulatus and P. corneti. At that time, no

articulated specimens of Phyllolepididae were known,

which led Lohest to define Pentagonolepis konincki with

anterior ventrolateral plates, Phyllolepis undulatus with

median dorsal plates and Phyllolepis corneti on the basis

of a sole centronuchal plate. Thanks to the discovery of a

complete specimen of Phyllolepis (Woodward 1915), Ler-

iche (1931) realized that the various plates were in fact

different parts of the same animal; Leriche thus put these

Lohest taxa in synonymy (under the name Phyllolepis

undulata and no longer ‘undulatus’, which was grammati-

cally incorrect). Later, on the basis of the material figured

by Lohest (1888b), Leriche (1931) and Stensi€o (1939)

assessed that there were two species of Phyllolepis in Bel-

gium. He assigned the material studied by Leriche and

Lohest to Phyllolepis undulata and to Phyllolepis konincki.

According to Stensi€o, P. konincki displayed a shorter and

broader centronuchal plate than that of P. undulata, a

median dorsal plate with the anterolateral margin slightly

forward directed, and an anterior ventrolateral plate

longer, in proportion to its width, than in P. undulata.

He also mentioned the ornamentation as a discriminating

character, with narrow ridges mostly situated further

apart in P. undulata than in P. konincki. This specific dis-

crimination character has recently been questioned by

Long and Daeschler (2013, p. 37), because ‘the nature of

A

F G

B

C

D E

F IG . 7 . Phyllolepis undulata, from Strud. Anterior ventrolateral plates. A, left AVL in external and internal views, IRSNB P.9476a.

B, left AVL in external and internal views, IRSNB P.9480. C, left AVL in internal view, IRSNB P.9479. D, left AVL in internal view,

IRSNB P.9477. E, left AVL in internal view, IRSNB P.9482. F, right AVL in external and internal views, IRSNB P.9481. G, left AVL in

external and internal views, IRSNB P.9478a. Abbreviations: oa.PVL, overlap area for posterior ventrolateral plate; pec.n, pectoral notch.

Scale bar represents 1 cm. Colour online.


A

F

G H

B

C D

E


the dermal ornamentation is quite variable within popula-

tions of phyllolepid species’.

A morphometrical analysis was carried out on cen-

tronuchal and anterior ventrolateral plates (Fig. 8) to

solve the problem of the number of Phyllolepis species

from Belgium. The majority of the material comes from

the locality of Strud, but a few plates from Ch�evremont

(Li�ege Province, Famennian) found in collection were

added to the morphometrical analysis. The centronuchal

and anterior ventrolateral plates were chosen for analysis

because they display a high degree of variability within

the Belgian material and are the most commonly encoun-

tered elements in the Famennian deposits of Belgium.

Moreover, those plates were used by Stensi€o (1939) to

justify the presence of two species of Phyllolepis in Bel-

gium. Thus, they appear to be the most appropriate

material to study the Phyllolepis diversity in Belgium.

Measurements of the centronuchal and anterior ventro-

lateral plates clearly show a distribution of points suggest-

ing that the size and shape variations are continuous

(R2 stretching from 0.8469 to 0.9434, Fig. 8). This result

argues against the interpretation of more than one species

within the sample. For instance, the overall shape of the

centronuchal plate (shorter and broader for P. konincki)

and of the anterior ventrolateral plate (longer and nar-

rower in P. konincki), used by Stensi€o as discriminating

characters (respectively Fig. 8A, D), are actually characters

within a range of continuous variations. Thus, we assume

that variations around the best-fit lines illustrate

intraspecific and ontogenetic variation. Consequently, we

assert that there is a sole species of Phyllolepis in Belgium.

The specific epithet ‘undulata’ is retained to follow

Leriche (1931), who acted as first reviewer, and thus

follow the article 24.2.1 of the International Code of

Zoological Nomenclature and the recommendation 24A,

which allow to ‘best serve stability and universality of

nomenclature’. Pentagonolepis konincki, Phyllolepis corneti,

Phyllolepis undulatus and Phyllolepis konincki are all

considered to be synonyms of P. undulata.

Phyllolepis rossimontina as a junior synonym of

Phyllolepis undulata

Lane and Cuffey (2005) described Phyllolepis rossimontina

from the Famennian deposits of the Catskill Formation,

at the Red Hill site, in north-central Pennsylvania. From

the same site, they recognized other specimens attributed

to various species of Phyllolepis: P. aff. concentrica,

P. aff. nielseni, P. aff. undulata and P. aff. woodwardi.

Later, several of these specimens were identified by Long

and Daeschler (2013) as specimens of P. rossimontina (see

Olive et al. 2015b, appendices 1–2 for details). To clarify

the situation, all the measurable anterior ventrolateral and

centronuchal plates of phyllolepids from the Catskill

Formation (Red Hill and Tioga sites) were integrated in

the morphometrical analysis (Fig. 9).

Figure 9 plots the different measurement ratios used.

The distribution of points for all graphs suggests that the

size variations are continuous, arguing against an inter-

pretation of more than one taxon within the sample,

based on the size. All points representing P. rossimontina

and the other species noticed by Lane and Cuffey (2005),

that is P. aff. nielseni, P. rossimontina?, P. sp. and

P. aff. concentrica, as well as unpublished material found

in the collection of the ANSP, fit within the distribution

of points from those representing P. undulata sensu this

article. This analysis suggests the presence of only one

species in the article of Lane and Cuffey (2005), which

corresponds to P. undulata found in Belgium.

Lane and Cuffey (2005) compared P. rossimontina with

P. undulata sensu Stensi€o (1939) and P. konincki sensu

Stensi€o (1939). According to them, P. rossimontina differs

from P. undulata by: (1) a centronuchal plate slightly

broader than long (roughly square in P. undulata); (2)

the curvature of the posterolateral margins of the cen-

tronuchal plate more pronounced; and (3) the posterolat-

eral corners of the same plate flared outward slightly.

Thanks to the morphometrical analysis (Fig. 9A), it is

clear that the first criterion is not correct, considering the

intraspecific and ontogenetic variations, and the fact that

the sample follows the best-fit line. The second and third

criteria are not directly explained by the morphometrical

analysis but as there are, within the sample, centronuchal

plates broader than long and centronuchal plates longer

than broad (for instance IRSNB P.9464, Fig. 4G), cen-

tronuchal plates displaying curved posterolateral margins

and flared posterolateral corners and some others not, it

could be related to intraspecific variation.

Concerning the anterior ventrolateral plate, Lane and

Cuffey (2005, p. 120) argued that P. undulata also differs

from P. rossimontina by: (1) a broader and more angular

shape; (2) a straight posterolaterally slanting lateral mar-

gin; (3) a sharp angle where the lateral and posterolateral

plate margins join; (4) a proportionally longer posterome-

dial margin; (5) a less well-pronounced curvature to the

F IG . 8 . Graphs of measurement ratios for CeNu (A–C) and AVL (D–H) of Phyllolepis specimens from Belgium. A, length vs width.

B, anterior length vs length. C, posterior margin length vs anterior margin length. D, length vs width. E, anterior length vs length. F,

lateral margin length vs medial margin length. G, medial margin length vs width. H, posterior margin length vs lateral margin length.

Abbreviations: L.ant, anterior length; L.mg.ant/lat/med/post, anterior/lateral/medial/posterior margin length. Colour online.


A

F

G H

B

C D

E


posterolateral margin; and (6) a more posterolaterally

slanting anterior margin. Our morphometrical analysis

shows that criteria 1 and 4 may be invalidated by contin-

uous variation (Fig. 9D, H). The other criteria, even if

not quantified by our morphometrical analysis, are not

sufficient to discriminate between species, as Figure 7

shows anterior ventrolateral plates displaying a strong

variability concerning the criteria 2, 3, 5 and 6.

Lane and Cuffey (2005, p. 120) also compared the cen-

tronuchal plate of P. rossimontina with that of P. konincki

sensu Stensi€o (1939). According to them, that of P. rossi-

montina differs from that of P. konincki in lacking the

ornamentation of concentric ridges. In agreement with

Long and Daeschler (2013), it should not be considered

as a diagnostic character. Moreover, in P. undulata sensu

this article, both ornamentation types are found in the

same specimens, that is concentric ridges and meshlike

ornamentation. Regarding the anterior ventrolateral plate,

Lane and Cuffey (2005, p. 123) argued that P. konincki

sensu Stensi€o (1939) differs from P. rossimontina by: (1)

a much more pronounced curve in the posterolateral

plate margin; (2) a better defined angle where the ante-

rior and lateral plates margins are joined; and (3) a pos-

terolaterally slanting lateral plate margin. As for the

comparison with the anterior ventrolateral plate of P. un-

dulata sensu Stensi€o (1939), Figure 7 illustrates anterior

ventrolateral plates of the Belgian species with a strong

variability for all these criteria, which are therefore insuf-

ficient to discriminate P. rossimontina from P. konincki

sensu Stensi€o (1939).

Therefore, and considering the results of our morpho-

metrical analysis, we consider that P. rossimontina is actu-

ally the same as P. undulata sensu this article. According

to the nomenclatural rule of priority, only the taxon

P. undulata is retained, whereas P. rossimontina becomes

a junior synonym of P. undulata.

Other Phyllolepis species, a preliminary analysis

The figured and measurable anterior ventrolateral and

centronuchal plates of other species (all Famennian in

age) of the genus Phyllolepis have also been integrated in

this study (Fig. 10): P. tolli Vasiliauskas, 1963 (Latvia);

P. concentrica Agassiz, 1844 (Scotland); P. woodwardi

Stensi€o, 1939, and P. orvini Heintz, 1930 (East Green-

land); P. nielseni Stensi€o, 1939 (East Greenland); and

P. thomsoni Long and Daeschler, 2013 (Pennsylvania,

USA). Several remarks can be made:

1. Concerning the centronuchal measurements, Phyllolepis

woodwardi approaches (Fig. 10B–C) or even overlaps

(Fig. 10A) the cloud of points represented by the speci-

mens of P. undulata (which includes all specimens of

P. rossimontina), but diverges more from that group

when considering the AVL measurements (Fig. 10D–E,G–H). However, measurements for the only available

AVL have been taken on an enlarged photography

without scale information (i.e. Stensi€o 1939, text-fig. 2)

and the point representing the AVL of P. woodwardi in

all graphs should therefore appear closer to the cloud

of P. undulata. In any case, the sample of P. woodwardi

is too limited to draw a conclusion.

2. Concerning P. nielseni, measurements made on the

AVL overlap those of P. nielseni and P. undulata

(Fig. 10D–H). Measurements made on the cen-

tronuchal plates show P. nielseni representative

slightly above the point cloud of P. undulata

(Fig. 10A–C). However, as for P. woodwardi (see

above), the sample is too limited.

3. On all graphs (Fig. 10A–H), all specimens of P. orvini

group together, except a centronuchal plate (Stensi€o

1936, pl. 2, fig. 1; Fig. 10A–B), which could either be

misidentified, or have a wrong scale on the publica-

tion, or even be a juvenile form. The points repre-

senting P. orvini clearly fall outside the range of

variation of P. undulata.

4. The only specimen (an AVL) known of P. tolli falls

within the cloud point of P. orvini. This would argue

for a synonymy between both taxa (P. tolli being

junior synonym of P. orvini).

5. Recently, Long and Daeschler (2013) described Phyl-

lolepis thomsoni from the late Famennian (expansa

zone) of Pennsylvania, USA. Only one centronuchal

plate is usable for the morphological analysis. Like for

P. nielseni, it approaches the P. undulata cloud, but

the size of the sample does not permit any further

conclusions.

6. P. concentrica is represented by a single AVL, well iso-

lated of all other species on the graphs.

An interesting feature appears when the best-fit line is

drawn for all species. The R2 ranges between 0.8964 and

0.996 (Fig. 10) and suggests that both size and shape vari-

ations are continuous for all species of Phyllolepis. It con-

firms that the genus Phyllolepis is strongly supported.

F IG . 9 . Graphs of measurement ratios for CeNu (A–C) and AVL (D–H) of Phyllolepis specimens from Belgium and Pennsylvania. A,

length vs width. B, anterior length vs length. C, posterior margin length vs anterior margin length. D, length vs width. E, anterior

length vs length. F, lateral margin length vs medial margin length. G, medial margin length vs width. H, posterior margin length vs

lateral margin length. Abbreviations: L.ant, anterior length; L.mg.ant/lat/med/post, anterior/lateral/medial/posterior margin length;

Pennsyl., Pennsylvania. Colour online.


A

F

G H

B

C D

E


However, recognizing different species within this genus

is more difficult. Three morphogroups may be identified:

one includes P. orvini and P. tolli, another one includes

P. concentric, and a third one includes P. undulata,

P. nielseni, P. woodwardi and P. thomsoni.

Faunal and floral affinities between Belgium and

Pennsylvania

The Red Hill site is a road-cut exposure in the upper part

of the Duncannon Member of the Catskill Formation in

Clinton County, Pennsylvania, USA. The fossil remains at

Red Hill come from a broad but narrow sequence of

strata (Cressler et al. 2010a), which are attributed to the

VCo palynozone (sensu Streel et al. 1987) within the

Famennian stage.

Daeschler and Cressler (2011, table 1) established a

list of the Red Hill fauna and flora contents. The verte-

brate fauna comprises phyllolepid and groenlandaspid

placoderms, gyracanthid acanthodians, primitive chon-

drichthyans, palaeoniscid actinopterygians, dipnoan,

rhizodontid, megalichthyid and tristichopterid sarcoptery-

gians, as well as early tetrapods. The antiarch placoderm

Bothriolepis and the porolepiform Holoptychius have not

been reported from the main fossiliferous zone at Red Hill

although they are very well represented in many other sites

of the Catskill Formation (Daeschler et al. 2003). Their

absence in Red Hill may be due to palaeoenvironmental

conditions (Cressler et al. 2010a). The palaeoenvironment

of the Red Hill site consists of a meandering stream sys-

tem, with frequent avulsion events, in which floodplain

ponds were present (Sevon 1985; Berg et al. 1993; Woo-

drow et al. 1995; Daeschler and Cressler, 1999, 2011;

Cressler et al. 2010a). The presence of terrestrial arthro-

pods (Shear 2000) and of well-preserved terrestrial plants

(Cressler 1999; Cressler et al. 2010b) reinforces the non-

marine interpretation of the Red Hill site (Daeschler et al.

2003).

Kenrick and Fairon-Demaret (1991) and then Cressler

et al. (2010b) and Prestianni and Gerrienne (2010)

noticed that the localities of the Belgian Evieux Forma-

tion, including Strud, display plant assemblages that are

very similar to those from the North American sites,

especially Red Hill. Both sites were located along the

southern margin of the Euramerica and date from the

same VCo palynozone (sensu Streel et al. 1987).

An equivalent observation may be made for the

vertebrate faunas. Phyllolepididae, Groenlandaspididae,

Acanthodii, Actinopterygii, Dipnoi, Megalichthyididae,

Tristichopterididae and Ichthyostegalia are found in both

localities. Similarities even reach the generic level, when

considering Turrisaspis (known in these two localities

only) and even the species level for Phyllolepis undulata

(see above). The main differences between the two locali-

ties are the chondrichthyans that are present in Red Hill,

but lacking in Strud, and the presence of antiarchs and

porolepiforms in Strud, which are absent in Red Hill

(although present in other localities of the Catskill Forma-

tion). An accurate quantification of the fauna similarities

between the Strud and Red Hill localities is difficult to

evaluate so far because taxonomic identification is still in

progress for some taxa at Strud and Red Hill. However,

even if not quantified, the similarities of fauna and flora

between these two late Famennian localities seem strong.

Exchanges between these sites over a single landmass (i.e.

Euramerica) would have been facilitated for plants with

aeolian spore dispersion. The close similarity of the verte-

brate faunas from these sites may be explained by faunal

exchanges. Several of the vertebrate taxa found in both

localities are considered to be non-marine; for example,

groenlandaspidids are at least considered cautiously as

indicators of non-marine rocks in the Famennian

(Daeschler et al. 2003). Thus, either there was a non-

marine link between Pennsylvania and Belgium during the

late Famennian, or the shared taxa were more tolerant of

marine/brackish conditions than previously thought

(Schultze and Cloutier 1996; Anderson et al. 1999).

Long and Daeschler (2013, p. 41) discussed the co-

occurrences of Phyllolepis species with early tetrapods,

bothriolepid antiarchs and groenlandaspid arthrodires in

several localities (notably P. tolli from Latvia, P. nielseni

from East Greenland, ‘P. rossimontina’ and P. thomsoni

from Pennsylvania). The placoderm fauna from Strud

represents a similar association. They also argued that the

presence of Phyllolepis in the Northern Hemisphere dur-

ing the late Famennian ‘may serve as an indicator of

favorable paleoecological conditions in the search for

early tetrapods’ (Long and Daeschler 2013, p. 41), a

remark already made by Young (2005c) and confirmed

herein by the association of Phyllolepis and an ichthyoste-

gid-like tetrapod in the late Famennian of Strud (Cl�ement

et al. 2004). However and to date, the absence of tetrapod

remains among several thousand specimens of vertebrate

F IG . 10 . Graphs of measurement ratios for CeNu (A–C) and AVL (D–H) of all Phyllolepis species. A, length vs width. B, anterior

length vs length. C, posterior margin length vs anterior margin length. D, length vs width. E, anterior length vs length. F, lateral mar-

gin length vs medial margin length. G, medial margin length vs width. H, posterior margin length vs lateral margin length. Abbrevia-

tions: Green., Greenland; L.ant, anterior length; L.mg.ant/lat/med/post, anterior/lateral/medial/posterior margin length; Pennsyl.,

Pennsylvania; Scot., Scotland. Colour online.


fossils in the Kl�unas fossil site, Latvia (T�ervete Fm, mid-

dle Famennian), where Phyllolepis tolli, Bothriolepis ornata

and B. jani have been found together, is a counterexam-

ple of the presence of Phyllolepis as indicator of tetrapods

(Vasilkova et al. 2012).

Acknowledgements. We gratefully acknowledge the Belgian Fed-

eral Science Policy Office for the research financial support

(Doctoral Fellow to SO) and the Jessup Fund (ANSP, Philadel-

phia, USA) for the funding provided to SO for the ANSP collec-

tion visit. This research received support, under the number

DK-TAF-2253, from the SYNTHESYS Project (http://www.syn-

thesys.info/), which is financed by European Community

Research Infrastructure Action under the FP7 Integrating Activi-

ties Programme. The authors thank J. Denayer (University of

Li�ege, Belgium; the University of Queensland, Australia) for his

remarks concerning the geological framework as well as E. Bour-

don (University of Copenhagen) for her help. The ANSP team

and G. Cuny (University of Copenhagen, now University of

Lyon) are thanked for their warm welcome in their respective

institutions. A. Folie, D. Nolf and A. Dreze (IRSNB, Brussels, Bel-

gium) granted access to the collection of the IRSNB and provided

technical support. E. Poty (ULg) granted access to the collection

of the ULg and M.-C. Van Dyck (UCL, Louvain-la-Neuve,

Belgium) to the collection of the University of Louvain-la-

Neuve. We thank the Gesves local council staff for providing

technical support and excavation permission in Strud. We are

indebted to the successive Strud field teams, who helped us since

2004. The authors thank the editor and the reviewers for their

fruitful remarks. The photographs were provided by W. Miseur

(IRSNB) and Philippe Loubry (CNRS/MNHN/UPMC-Paris 6,

Paris, France). This article is a contribution to the ANR TERRES

2010-BLAN-607-03 project.

DATA ARCHIVING STATEMENT

Data for this study are available in the Dryad Digital Repository:

http://dx.doi.org/10.5061/dryad.qb550

Editor. Zerina Johanson

REFERENCES

AGASSIZ, L. 1844. Monographie des poissons fossiles du Vieux

Gr�es Rouge ou syst�eme d�evonien (Old Red Sandstone) des Isles

Britanniques et de Russie. Jent and Gassman, Neuchatel, 171

pp.

ANDERSON, M. E., LONG, J. A., EVANS, F. J., AL-

MOND, J. E., THERON, J. N. and BENDER, P. A. 1999.

Biogeographic affinities of Middle and Late Devonian fishes of

South Africa. Records of the Western Australian Museum,

suppl. 57, 157–168.BERG, T. M., McINERNEY, M. K., WAY, J. H. and

MACLACHLAN, D. B. 1993. Stratigraphic correlation chart

of Pennsylvania. Commonwealth of Pennsylvania, Bureau of

Topographic and Geologic Survey, Pennsylvania, General

Geology Report 75.

CL�EMENT, G. and BOISVERT, C. A. 2006. Lohest’s true

and false ‘Devonian amphibians’: evidence for the rhyn-

chodipterid lungfish Soederberghia in the Famennian of

Belgium. Journal of Vertebrate Paleontology, 26, 276–283.-and PRESTIANNI , C. 2009. Fauna, flora, and paleoenvi-

ronment of the Upper Devonian tetrapod-bearing locality of

Strud, Belgium. 105–115. In GODEFROIT, P. and LAM-

BERT, O. (eds). Tribute to Charles Darwin and Bernissart

iguanodons: New perspectives on vertebrate evolution and Early

Cretaceous ecosystems. Institut royal des Sciences naturelles de

Belgique, Brussels, 145 pp.

-AHLBERG, P. E., BLIECK, A., BLOM, H., CLACK,

J. A., POTY, E., THOREZ, J. and JANVIER, P. 2004.

Palaeogeography: Devonian tetrapod from Western Europe.

Nature, 427, 412–413.CRESSLER, W. L. III 1999. Site-analysis and floristics of the

Late Devonian Red Hill locality, Pennsylvania, an Ar-

chaeopteris-dominated plant community and early tetrapod

site. Published PhD thesis, University of Pennsylvania,

Philadelphia, 156 pp.

- DAESCHLER, E. B., SLINGERLAND, R. and

PETERSON, D. A. 2010a. Terrestrialization in the Late

Devonian: a palaeoecological overview of the Red Hill site,

Pennsylvania, USA. 111–128. In VECOLI , M., CL�EMENT,

G. and MEYER-BERTHAUD, B. (eds). The terrestrializa-

tion process: modelling complex interactions at the biosphere–geosphere interface. Geological Society, London, Special Publi-

cation, 339, 192 pp.

-PRESTIANNI , C. and LEPAGE, B. A. 2010b. Late

Devonian spermatophyte diversity and paleoecology at Red

Hill, north-central Pennsylvania, U.S.A. International Journal

of Coal Geology, 83, 91–102.DAESCHLER, E. B. and CRESSLER, W. L. 1999. Sampling

an early continental ecosystem: Late Devonian bone beds at

Red Hill, Clinton County, PA. Journal of Vertebrate Paleontol-

ogy, 19, 41A.

--2011. Late Devonian paleontology and paleoenviron-

ments at Red Hill and other fossil sites in the Catskill Forma-

tion of north-central Pennsylvania. 1–16. In RUFFOLO,

R. M. and CIAMPAGLIO, C. N. (eds). From the shield to

the sea: geological trips from the 2011 Joint Meeting of the GSA

Northeastern and North-Central Sections. Geological Society of

America, Field Guide, 20, 116 pp.

- FRUMES, A. C. and MULLISON, C. F. 2003.

Groenlandaspidid placoderm fishes from the Late Devonian

of North America. Records of the Australian Museum, 55,

45–60.DENAYER, J., PRESTIANNI , C., GUERIAU, P., OLIVE,

S. and CL�EMENT, G. in press. Stratigraphy and deposi-

tional environments of the Late Famennian (Late Devonian)

of Southern Belgium and characterization of the Strud locality.

Geological Magazine.

DUPRET, V. 2003. Etude anatomique des genres Kuj-

danowiaspis Stensi€o 1942 et Erikaspis nov. gen. (Placodermi

Arthrodira “Actinolepida”) du D�evonien inf�erieur de Podolie

(Ukraine). Nouvelle proposition de la phylog�enie des Arthro-


http://www.synthesys.info/

http://www.synthesys.info/


dira. Unpublished PhD thesis, Mus�eum national d’Histoire

naturelle, Paris, 403 pp.

-2010. Revision of the genus Kujdanowiaspis Stensi€o, 1942

(Placodermi, Arthrodira, “Actinolepida”) from the Lower

Devonian of Podolia (Ukraine). Geodiversitas, 32, 5–63.-and ZHU, M. 2008. The earliest phyllolepid (Placodermi,

Arthrodira), Gavinaspis convergens, from the late Lochkovian

(Lower Devonian) of Yunnan (South China). Geological Maga-

zine, 145, 257–278.GARROUSTE, R., CL�EMENT, G., NEL, P., ENGEL, M.

S., GRANDCOLAS, P., D’ HAESE , C., LAGEBRO, L.,

DENAYER, J., GUERIAU, P., LAFAITE , P., OLIVE,

S., PRESTIANNI , C. and NEL, A. 2012. A complete insect

from the Late Devonian. Nature, 488, 82–85.------------

-2013. Garrouste et al. reply. Nature 494, E4–E5.GOUJET, D. 1984. Les poissons placodermes du Spitsberg –Arthrodires Dolichothoraci de la Formation de Wood Bay

(D�evonien inf�erieur). Editions du CNRS, Paris, 284 pp.

GROSS , W. 1965. €Uber die placodermen-gattungen As-

terolepis und Tiaraspis aus dem Devon Belgiens und einen

fraglichen Tiaraspis-rest aus dem Devon Spitzbergens. Bul-

letin de l’Institut royal des Sciences naturelles de Belgique, 61,

1–19.GUERIAU, P., CHARBONNIER, S. and CL�EMENT, G.

2014a. First decapod crustaceans in a Late Devonian continen-

tal ecosystem. Palaeontology, 57, 1203–1213.---2014b. Angustidontid crustaceans from the Late

Devonian of Strud (Namur Province Belgium): insights into

the origin of Decapoda. Neues Jahrbuch f€ur Geologie and

Pal€aontologie, Abhandlungen, 273/3, 327–337.HEINTZ, A. 1930. Oberdevonische fischreste aus €Ost-

Gr€onlands. Skrifeter om Svalbard og Ishavet, 30, 31–46.H €ORNSCHEMEYER, T., HAUG, J., B�ETHOUX, O.,

BEUTEL, R. G., CHARBONNIER, S., HEGNA, T. A.,

KOCH, M., RUST, J., WEDMANN, S., BRADLER, S.

and WILLMANN, R. 2013. Is Strudiella a Devonian insect?

Nature, 494, E3–E4.JANVIER, P. and CL�EMENT, G. 2005. A new groenlandas-

pidid arthrodire (Vertebrata: Placodermi) from the Famennian

of Belgium. Geologica Belgica, 8, 51–67.KENRICK, P. and FAIRON-DEMARET, M. 1991.

Archaeopteris roemeriana (G€oppert) sensu Stockmans, 1948

from the Upper Famennian of Belgium: anatomy and leaf

polymorphism. Bulletin de l’Institut royal des Sciences naturelles

de Belgique, Sciences de la Terre, 61, 179–195.LAGEBRO, L., GUERIAU, P., HEGNA, T. A., RABET,

N., BUTLER, A. and BUDD, G. E. 2015. The oldest noto-

stracan (Upper Devonian Strud locality, Belgium). Palaeontol-

ogy, 58 (3), 497–509.LANE, J. A. and CUFFEY, R. 2005. Phyllolepis rossimontina

sp. nov. (Placodermi) from the Uppermost Devonian at Red

Hill. North-Central Pennsylvania. Revista Brasileria de Paleon-

tologia, 8, 117–126.LERICHE, M. 1931. Les poissons famenniens de la Belgique –Les faci�es du Famennien dans la r�egion gallo-belge – Les

relations entre les formations marines et les formations

continentales du D�evonien sup�erieur sur la bordure

m�eridionale du continent Nord-Atlantique. M�emoires de la

Classe des Sciences de l’Acad�emie Royale de Belgique, 4, 1–72.LOHEST, M. 1888a. D�ecouverte du plus ancien amphibien

connu et de quelques fossiles remarquables dans le Famennien

sup�erieur de Modave. Annales de la Soci�et�e G�eologique de

Belgique, 15, 129–137.-1888b. Recherche sur les poissons des terrains pal�eozo€ıques

de Belgique. Poissons des Psammites du Condroz, Famennian

sup�erieur. Annales de la Soci�et�e G�eologique de Belgique, 15,

112–203.LONG, J. A. 1984. New phyllolepids from the Victoria and the

relationships of the group. Proceedings of the Linnean Society

of New South Wales, 107, 263–308.-and DAESCHLER, E. B. 2013. First articulated phyl-

lolepid placoderm from North America, with comments on

phyllolepid systematics. Proceedings of the Academy of Natural

Sciences of Philadelphia, 162, 33–46.-ANDERSON, M. E., GESS , R. and HILLER, N. 1997.

New placoderm fishes from the Late Devonian of South

Africa. Journal of Vertebrate Paleontology, 17, 253–268.MILES , R. S. 1973. An actinolepid arthrodire from the Lower

Devonian Peel Sound Formation, Prince of Wales Island.

Palaeontographica, 143, 109–118.OBRUCHEV, D. V. 1964. Agnatha and fish. In ORLOV, Yu.

(ed.). Fundamentals of paleontology, Vol. 11. Moscow, 522 pp.

OLIVE, S. in press. Devonian antiarch placoderms from Bel-

gium revisited. Acta Paleontologica Polonica. doi:10.4202/app.

00015.2013

-PRESTIANNI , C. and DUPRET, V. 2015a. A new spe-

cies of Groenlandaspis Heintz, 1932 (Placodermi, Arthrodira),

from the Famennian (Late Devonian) of Belgium. Journal of

Vertebrate Paleontology, 35 (4). doi:10.1080/02724634.2014.

935389

-CL�EMENT, G., DAESCHLER, E. B. and DUPRET,

V. 2015b. Data from: Characterization of the placoderm

(Gnathostomata) assemblage from the tetrapod-bearing local-

ity of Strud (Belgium, upper Famennian). Dryad Digital

Repository. doi:10.5061/dryad.qb550

PRESTIANNI , C. and GERRIENNE, P. 2010. Early seed

plant radiation: an ecological hypothesis. Geological Society,

London, Special Publication, 339, 71–80.-STREEL, M., THOREZ, J. and GERRIENNE, P.

2007. Strud: old quarry, new discoveries. Preliminary report.

43–47. In STEEMANS, P. and JAVAUX, E. (eds). Recent

advances in palynology. Carnet de G�eologie, Brest, Memoir

2007/01, 73 pp.

RITCHIE , A. 1984. A new placoderm, Placolepis gen. nov.

(Phyllolepidae), from the Late Devonian of New South Wales,

Australia. Proceedings of the Linnean Society of New South

Wales, 107, 321–353.-2005. Cowralepis, a new genus of phyllolepid fish (Pisces,

Placodermi) from the Late Middle Devonian of New South

Wales, Australia. Proceedings of the Linnean Society of New

South Wales, 126, 215–259.SCHULTZE, H-P. and CLOUTIER, R. 1996. Comparison

of the Escuminac Formation ichthyofauna with other late

Givetian/early Frasnian ichthyofaunas. 348–368. In

SCHULTZE, H-P. and CLOUTIER, R. (eds). Devonian

OLIVE ET AL . : P LACODERMS FROM STRUD 1001

http://dx.doi.org/10.4202/app.00015.2013

http://dx.doi.org/10.4202/app.00015.2013

http://dx.doi.org/10.1080/02724634.2014.935389

http://dx.doi.org/10.1080/02724634.2014.935389


Fishes and Plants of Miguasha, Quebec, Canada. Verlag Dr.

Friedrich Pfeil, M€unchen, 374 pp.

SEVON, W. D. 1985. Nonmarine facies of the Middle and Late

Devonian Catskill coastal alluvial plain. 79–90. In WOO-

DROW, D. L. and SEVON, W. D. (eds). The Catskill Delta.

Geological Society of America, Special Paper, 201, 246 pp.

SHEAR, W. A. 2000. Gigantocharinus szatmaryi, a new trigono-

tarbid arachnid from the Late Devonian of North America

(Chelicerata, Arachnida, Trigonotarbida). Journal of Paleontol-

ogy, 74, 25–31.STENSI €O, E. 1934. On the Placodermi of the Upper Devonian

of East Greenland. I. Phyllolepida and Arthrodira. Meddelelser

om Grønland, 97, 1–58.-1936. On the Placodermi of the Upper Devonian of East

Greenland. Supplement to Part I. Meddelelser om Grønland,

97, 1–53.-1939. On the Placodermi of the Upper Devonian of East

Greenland. Second supplement to Part I. Meddelelser om

Grønland, 97, 1–33.STREEL , M., HIGGS, K. T., LOBOZIAK, S., RIEGEL,

W. and STEEMANS, P. 1987. Spore stratigraphy and corre-

lation with faunas and floras in the type marine Devonian of

the Ardenne-Rhenish Regions. Review of Palaeobotany and

Palynology, 50, 211–229.THOREZ, J., DREESEN, R. and STREEL , M. 2006. Fa-

mennian. Geologica Belgica, 9, 27–45.VASILIAUSKAS, V. 1963. Phyllolepis tolli sp. nov. and some

problems concerning the stratigraphy of the Famennian

deposits in the Baltic area. 407–429. In GRIGELIS , A. A. and

KARATAJUTE-TALIMAA, V. N. (eds). Geology of Lithua-

nia. Lietuvos TSR Mokslu akademija, Vilnius. [in Russian]

VASILKOVA, J., LUKSEVICS , E., STINKULIS , G. and

ZUPINS, I. 2012. Taphonomy of the vertebrate bone beds

from the Kl�unas fossil site, Upper Devonian T�ervete Forma-

tion of Latvia. Estonian Journal of Earth Sciences, 61, 105–119.

WOODROW, D. L., ROBINSON, R. A. J., PRAVE, A. R.,

TRAVERSE, A., DAESCHLER, E. B., ROWE, N. D. and

DELANEY, N. A. 1995. Stratigraphic, sedimentological and

temporal framework of Red Hill 41 (Upper Devonian Catskill

Formation) near Hyner, Clinton County, Pennsylvania: site of

the oldest amphibian known from North America. p. 16. In

WAY, J. (ed.). Field Trip Guide, 60th Annual Field Conference

of Pennsylvania Geologists. Lock Haven University, Lock Haven.

WOODWARD, A. S. 1891. Catalogue of the fossil fishes in the

British Museum of Natural History. Part II. Containing the

Elasmobranchii (Acanthodii), Holocephali, Ichthyodorulites,

Ostracodermi, Dipnoi, and Teleostomi (Crossopterygii), and

chondrostean Actinopterygii. British Museum of Natural

History, London, 567 pp.

-1915. Preliminary report on the fossil fishes from Dura

Den. Reports of the British Association for Advancement of

Science, Australia, 84, 122–123.YOUNG, G. C. 2005a. An articulated phyllolepid fish (Placo-

dermi) from the Devonian of central Australia: implications

for non-marine connections with the Old Red Sandstone con-

tinent. Geological Magazine, 142, 173–186.-2005b. A new phyllolepid placoderm occurrence (Devonian

fish) from the Dulcie Sandstone, Georgina Basin, central Aus-

tralia. Proceedings of the Linnean Society of New South Wales,

126, 203–213.-2005c. New phyllolepids (placoderm fishes) from the

Middle-Late Devonian of Southeastern Australia. Journal of

Vertebrate Paleontology, 25, 261–273.ZIEGLER, P. A. 1990. Geological atlas of Western and Central

Europe, Second edition. Shell Internationale Petroleum

Maatschappij B.V. & Geological Society of London, Avon,

239 pp.


Characterization of the placoderm (Gnathostomata) assemblage from the tetrapod-bearing locality of Strud (Belgium, Upper Famennian)

Documents