Journal of South American Earth Sciences 16 (2003) 155–161

www.elsevier.com/locate/jsames

Late Devonian acanthodians from Colombia

Carole J. Burrowa,*, Philippe Janvierb,c, Carlos Villarroeld
a
Department of Zoology and Entomology, University of Queensland, Brisbane 4072 Queensland, Australia
b
UMR 8569 du CNRS, Laboratoire de Paléontologie, Muséum National d’Histoire Naturelle, 8 rue Buffon, 75005 Paris, France
c
Department of Palaeontology, The Natural History Museum, SW7 5BD London, UK
d
Departamento de Geociencias, Universidad Nacional de Colombia, Aptdo Aéreo 56833, Bogotá, Colombia
Received 1 February 2002; accepted 1 December 2002

Abstract
Acanthodian remains occur in micaceous siltstone lenses (presumed to have been deposited during a marine incursion) in the Cuche
Formation (?Frasnian) of northeast Colombia. The acanthodians are represented by patches of scales from climatiidid Nostolepis sp. cf.
N. gaujensis and a fin spine and scales from a new diplacanthid. Type material of N. gaujensis is from the Frasnian Sventoji regional stage in
the Baltic, and Nostolepis sp. cf. N. gaujensis has been recorded in the Frasnian of Iran, as well as from Colombia. The new diplacanthid
taxon shows affinity to Baltic and Antarctic diplacanthids. The fauna thus shows possible links to both Gondwanan and Euramerican
acanthodian assemblages.
q 2003 Elsevier Science Ltd. All rights reserved.
Keywords: Acanthodii; Cuche Formation; frasnian; Devonian; Colombia

1. Introduction 2. Occurrence and stratigraphy

The vertebrate fauna from the Late Devonian Cuche
Formation of Colombia has been briefly described by Janvier
and Villarroel (1998, 2000). It is known to comprise antiarchs
(probably two Bothriolepis species and a large Asterolepis
species), an arthrodire (presumably either a groenlandaspidid
or a primitive brachythoracid), an actinopterygian (a Mimia-
like form), at least three sarcopterygians (a rhizodontid, a
holoptychiid, and a large cosmine-covered osteolepiform), a
chondrichthyan (Antarctilamna sp.), and acanthodians. The
assemblage includes forms comparable to coeval Euramer-
ican taxa, as well as to others known from Gondwana (Janvier
and Villarroel, 2000). Most of the material consists of
scattered scales and dermal bones. The acanthodians, which
are represented by isolated scales, patches of squamation, and
isolated spines, are described here in detail. The material
described herein is housed in the collection of the Department
of Geosciences, Universidad Nacional de Colombia, Bogota,
Colombia (UN-DG-V) and the Museo Geologico of the
Caldas University, Manizales, Colombia (MGCU).
* Corresponding author. Tel.: þ 61-7-33654825; fax: þ 61-7-33651655.
E-mail address: cburrow@zen.uq.edu.au (C.J. Burrow).
0895-9811/03/$ - see front matter q 2003 Elsevier Science Ltd. All rights reserved.
doi:10.1016/S0895-9811(03)00026-9
The Cuche Formation outcrops around the igneometa-
morphic core of the Floresta Massif in the Boyacá
department of Colombia (Fig. 1), where it overlies to
various extents two other Devonian formations, the El Tibet
and Floresta formations (for extensive geological infor-
mation, see Janvier and Villarroel, 2000; Berry et al., 2000;
Moreno-Sanchez, 2001). Unlike the Floresta Formation,
which yields a rich marine invertebrate fauna of Emsian
age, the Cuche Formation consists of reddish sandstones and
is almost barren except for some clayey lenses that yield
plant remains, ostracods, bivalves, lingulids, and fish
remains. These fossiliferous lenses principally outcrop in
the southwestern part of the Floresta Massif, approximately
5.2 km southwest of the Floresta village in the Quebrada
Potrero Rincón, and are rather conspicuous in that they
appear as yellowish patches on the dominantly reddish
background of the Cuche Formation. They are evidence of
slight marine incursions, because they yield inarticulate
brachiopods that are otherwise lacking in the Cuche
Formation.
Acanthodian remains have been found in only one of
these lenses (Potrero Rincon 1 [PR1] in Janvier and
Villarroel, 2000), which consists of thin-bedded, micaceous
156 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161
Fig. 1. Locality map (Janvier and Villarroel, 2000, Fig. 1).
siltstone in which delicate fish (acanthodian and actinopter-
ygian) and plant remains are preserved. This kind of facies
suits the preservation of articulated specimens, and the
acanthodian spine and associated portion of squamation
described herein from this outcrop probably belong to the
same individual.
The age of the Cuche Formation (at least its fish-bearing
part) has been discussed by Janvier and Villarroel (2000)
and considered as late Frasnian or early Famennian on the
basis of the fish fauna. If it is Frasnian, the abundance
of Strepsodus-like rhizodontid remains suggests a latest
Frasnian age, because these rhizodontids occur elsewhere
in the Famennian and Carboniferous. This dating also
agrees with the late Frasnian age of the plant remains in
the same outcrops (Berry et al., 2000; Moreno-Sanchez,
2001).
3. Description of the Acanthodians
Class Acanthodii [Owen, 1846]
Family Climatiidae [Berg, 1940]
Genus Nostolepis [Pander, 1856]
Type species Nostolepis striata [Pander, 1856]
Nostolepis gaujensis [Valiukevicius, 1998]. The species
is diagnosed by having large scales with four to six broad
ridges on a diamond-shaped crown. The neck is low and
stout, the base is large and strongly convex (Valiukevicius,
1998).
Nostolepis sp. cf. N. gaujensis (Figs. 2a and 3a –d).
UN-DG-VJ1 (three small siltstone flakes) has one piece
with a possibly articulated scale patch and two pieces with
patches of disarticulated scales. MGCU-B17 is a larger rock
with a large patch of disarticulated scales (Fig. 2a).
The scale morphology (Fig. 2a– c) shows large acantho-
dian scales 0.5 – 1.2 mm wide, 0.8 – 0.0 mm long, and
, 1.0 mm high. The crown is relatively flat and bears four
or five subparallel ridges that extend the length of the crown.
The ridges are wide with a rounded crest and separated by
wide, shallow troughs. The neck is short, with a shallow
slope from the anterior crown edge toward the neck/base
rim; the base protrudes in front of the crown. The posterior
part of the crown overhangs the base by up to half the length
of the crown. The globular base is of low to moderate
height.
Structurally (Fig. 2d), most scales have been reminer-
alized or replaced, though the outer layers in some scales are
preserved as a phosphatic shell with some histological
structure discernible. A horizontal section of the crown
shows cross-cut dentine tubules rising in the anterior crown,
remnants of vascular canals running back between the
ridges, and horizontal dentine tubules branching off
perpendicular to the canals.
Nostolepid scales are rare in Middle Devonian and
younger deposits; only two species—Nostolepis kernaven-
sis [Valiukevicius, 1985] and N. gaujensis—have been
erected for late Middle – Late Devonian material. Morpho-
logically, the Colombian scales compare well with the type
material of N. gaujensis from the Sventoji Formation and
Gauja and Amata regional substages (Frasnian) of the
Sventoji regional stage in the Baltic countries (Valiukevi-
cius, 1998); and scales assigned to Nostolepis sp. cf. N.
gaujensis from Frasnian? deposits near Chahriseh (Ghola-
malian et al., 2000; Turner et al., 2002, figs. 9, 10), and Bidu
1 (Chanaruh; Frasnian), north of Kerman (Janvier, 1974,
1977), Iran. Because none of the type scales of N. gaujensis
was sectioned, histological comparison is not possible,
though we assume that the type scales were deemed to have
Nostolepis-type histology. Longitudinal and transverse thin
sections of the Iranian Nostolepis sp. cf. N. gaujensis scales
show Stranggewebe (a specialized mesodentine with thin,
closely spaced, parallel tubules/lacunae; Gross, 1971) in the
inner layers of the crown (Tumer et al, 2002, Ag. 10 A), but
because the Colombian scales are hollow shells with their
histological structure only preserved in the outer layers,
their innermost structure is unknown. The scales of
Nostolepis kernavensis (upper Eifelian, Narva regional
substage, East Baltic) differ from those of Nostolepis
gaujensis and Nostolepis sp. cf. N. gaujensis in that they
are larger and the medial crown ridges are usually elevated
above the rest of the crown.
Family Diplacanthidae [Woodward, 1891]
Florestacanthus new genus
Type species—Florestacanthus morenoi new species
Diagnosis the same as for the type species, by monotypy.
Etymology after the formation in which the type material
was found.
Florestacanthus morenoi new species (Figs. 2b – d, 3e –i,
and 4a –i)
 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161 157

Fig. 2. Acanthodian-bearing specimens from the Cuche Formation. (a) Micaceous siltstone block MGCU-B.17 with patch of Nostolepis sp. cf. N. gaujensis
scales. (b–d) Micaceous siltstone block MGCU-B.20 with fin spine and scales of Florestacanthus morenoi nov. gen. et sp. (b) View of whole specimen (plus
plant fragments, palaeoniscoid scales, ostracods, ?sponges). Scale bar ¼ 10 mm. (c) Close up of fin spine impression approximately 25 mm from the tip. Scale
bar ¼ 1 mm. (d) Close up near the middle of the eroded fin spine, showing the radial trabeculae of the middle layer. Scale bar ¼ 0.02 mm.

2000 Cheiracanthoides? sp. indet. Janvier and Villarroel,
pp. 736– 8, text fig. 5B, C
Diagnosis a diplacanthid acanthodian with slender,
relatively straight unpaired fin spines with a subcircular
cross-section. The exserted part of the spine is ornamented
with six or more sharp longitudinal ridges per side and a broad
ridge along the leading edge. The central pulp cavity of the
spine is wide and extends almost to the distal tip. The scales
have a relatively flat crown bearing 10 – 20 close-set
subparallel ridges on the anterior half, and some have a
narrow, smooth rim in front of the ridges. The height of the
neck is approximately one-third total scale height, and the
scale base is deepest centrally. The scales have centrifugal
dentine tubules in the top layers of the crown and lack wide
vascular canals and durodentine/enameloid.
Description — The holotype specimen MO – CU – B.20
(Fig. 2b) comprises a fin spine, disarticulated scales, and
a small patch of articulated scales that are probably part
of the fin web.
The scale morphology (Figs. 3e – f and 4d –i) shows that
the flank scales are 0.5 – 1.0 mm wide, 0.5 –1.0 mm long,
and 0.4 –0.7 mm high. The crown of most scales is flat;
some are slightly depressed centrally. The crown extends
slightly beyond the base. Ten to 20 close-set subparallel
ridges ornament the anterior half of the crown and are
narrow and unbranched. The anterior crown edge has a
rounded outline; some scales have a narrow smooth rim in
front of the ridges. There is no indication of pores opening
between the ridges. The neck is strongly concave all around
and constitutes approximately one-third of the height of the
scale. A marked rim separates the base and neck. The base is
rounded and deepest at the center (i.e. it is not pushed
forward). Small, presumably fin web scales are approxi-
mately 0.3 mm wide and 0.2 mm long with shapes
comparable to those of the flank scales.
The scale histology (Fig. 3g) revealed by the horizontal
section (Fig. 3h) of the crown shows some preservation of the
outer, relatively narrow growth zones. Dentine tubules are
centrifugal; some vertical dentine canals have been cut
through in the anterior growth zones. The vertical longitudinal
section (Fig. 3h) shows the delineation between the base and
crown: The base apex forms a very low angle pyramid.
Vertically directed, fine-caliber dentine canals and tubules are
preserved in the crown. No wide vascular canals, durodentine,
or enameloid are visible. The base has some preserved
remnants of Sharpey’s fibers and canals of Williamson.
The fin spine morphology (Figs. 2b– c and 4a – c) indicates
that the whole 67 mm length of the spine is preserved eithier as
an impression or the original hard tissue; the exposed surface
represents the eroded trailing, or posterior, face. The spine is
relatively slender and perfectly straight with a subcircular
cross-section; it is widest (3.5 mm) near the insertion/
exsertion boundary. Because the spine is symmetrical, we
presume it is from an unpaired fin. The insertion area is
approximately 20 mm long and semicircular in cross-section,
tapers only gradually toward the proximal end, and is
158 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161

Fig. 3. Acanthodian scales from the Cuche Formation, Colombia. (a–d) Nostolepis sp. cf. N. gaujensis. (a) MGCU-B.17.1, crown view. (b) MGCU-B.17.2,
crown view. (c) MGCU-B.17.2, lateral view (most of the base is missing). (d) MGCU-B.17.3, horizontal thin section of crown, anterior corner. Scale
bar ¼ 0.1 mm. (e –i) Florestacanthus morenoi n. gen. et sp. (e) Scale MGCU-B.20.1, crown view. (f) Scale MGCU-B.20.1, anterior view. (g) Vertical
longitudinal ground thin section of scale MGCU-B.20.2. (h) Horizontal ground thin section of scale crown MGCU-B.20.3. (i) Cross-section of spine 10 mm
from the tip of MGCU-B.20.4. Scale bar ¼ 0.1 mm. adt ¼ ascending dentine tubules; cc ¼ central longitudinal cavity; r ¼ longitudinal ridges; and
vc ¼ vascular canal. Arrow is anteriad.

ornamented with fine, close-set, parallel striations. Each side
of the exserted part bears six or more sharp, narrow ridges
separated by wider, shallow grooves; the leading edge is broad
and appears ornamented with approximately 16 fine-caliber
parallel nodose ridges of comparable width to those on the
inserted part.
The fin spine internal structure (Figs. 2d and 3i)
shows a wide central pulp cavity extending almost to the
distal tip of the spine. The hard tissue is oriented mainly
longitudinally for the proximal two-thirds; 10 radially
directed trabeculae characterize the mid-length portion.
The spine appears to be formed of a thin, dense outer
layer, a trabecular middle layer, and a thin, dense inner
layer surrounding the pulp cavity.
Etymology—The species is named after Dr Mario
Moreno-Sanchez, University of Manizales, who discov-
ered the specimens described herein.
Holotype—A patch of squamation associated with a
fin spine is presumed to derive from the same individual
(MGCU-B.20)
Type locality and stratiography a clayey lens referred to
as PR1 in Janvier and Villarroel (2000, fig 1), Floresta,
Boyacá Department, Colombia; Cuche Formation latest
Frasnian or early Famennian.
Reffered material—Impressions of isolated scales on
sample UN-DG-PAL V40 (Janvier and Villarroel, 2000,
figs. 5C, D, 12), and scale MGCU-B20.1, ground thin
sections of scales MGCU-B20.2 and MGCU-B20.3, and a
ground thin section of fin spine fragment MGCU-B20.4
from the holotype.
Discussion—Scales of the same type as those of
Florestacanthus morenoi were assigned to Cheira-
canthoides? sp. by Janvier and Villarroel (2000); these are
preserved as impressions on the sample UN-DG-PAL V40
(from the same clayey lens as the specimens described
here), along with actinopterygian remains. The scales
superficially resemble those of Cheiracanthoides comptus
[Wells, 1944]. Although poorly preserved, they appear to
differ from the latter (and other Cheiracanthoides spp.) in
lacking Stranggewebe and a regular network of widened
 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161 159

Fig. 4. Spine and scales of Florestacanthus morenoi nov. gen. et sp., Cuche Formation, Colombia. (a –c) Reconstruction of MGCU-B.20, unpaired fin spine. (a)
Trailing edge view. (b) Leading edge view. (c) Lateral view with outlines of cross-sections at the level of the insertion area and distally. Scale bar ¼ 10 mm.
(d–i) Camera lucida drawings of scales from MGCU-B.20. (d) MGCU-B.20.5, crown view. (e) MGCU-B.20.5, basal view. (f) MGCU-B.20.5, lateral view. (g)
MGCU-B.20.6, crown view. (h) MGCU-B.20.6, basal view. (i) MGCU-B.20.6, lateral view. Scale bar ¼ 0.1 mm. Arrow is anteriad.

vascular canals in the crown and in having canals of
Williamson in the base. The only other scales of a similar
age from South America that have been assigned to the
Acanthodii are from the upper member of the Campo Chico
Formation (middle Frasnian) of Venezuela (Young and
Moody, 2002, figs. 16A – C). However, the structure of these
scales indicates that they are from a protacrodont chon-
drichthyan rather than an acanthodian.
Scales from several other Middle –Late Devonian taxa
from other regions look similar to the Colombian scales.
Those of diplacanthidid Rhadinacanthus balticus [Gross,
1973] (Eifelian Narova beds, Estonia) differ in having
bifurcating ridges on the crown. Scales of Devononchus
tennuispinus and D. concinnus [Gross, 1930] (e.g. Valiu-
kevicius, 1998, pl. 8, figs. 17 – 20) have many more crown
ridges, which are only weakly and variably developed,
whereas those of D. ketleriensis [Gross, 1947] have a low
base and crown ridges that converge posteriorly (Denison,
1979). Of the scales with a smooth anterior rim on the
crown, the Colombian ones most closely resemble those of
the cheiracanthidids Cheiracanthus brevicostatus and
Markacanthus parallelus [Valiukevicius and Karatajute-
Talimaa, 1986], but as in all acanthodid scales, they have
well-developed enameloid layers in the center of the crown,
and those of the latter taxon (Valiukevicius and Karataju-
te-Talimaa, 1986, pl. II, Figs. 10 –11; pl. III, fig. 1) have
ridges covering the whole length of the crown, a relatively
deep neck, and a somewhat pointed base.
Several Gondwanan acanthodian taxa of a similar age are
known. The west Venezuelan fish fauna described by Young
and Moody (2002) includes Machaeracanthus fin spines
from the early Frasnian and smaller acanthodian fin spines
plus one Acanthodes-type scale from the middle Frasnian of
the Campo Chico Formation. None of these fin spines
compares closely to the Colombian example. Semi-
articulated diplacanthidids from the ? middle Givetian of
Antarctica (Young, 1989) have scales that resemble those of
Florestacanthus morenoi, but the former have a pore canal
system in the anterior crown and some bifurcating crown
ridges. The fin spines of the Antarctic fish match those of
Byssacanthoides debenhami [White, 1968] (?late Givetian,
Antarctica), which have a subcircular cross-section and
similar structure but also have radial trabeculae developed
for most of the length of the spine and an ornament that
comprises relatively broad, flat ridges separated by narrow
grooves.
Rare scales from deposits in the Chahriseh region of Iran
superficially resemble both the Antarctic and Colombian
160 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161
scales, but they have a deep neck and appear to have pore
openings in the crown grooves. Thus, they could be
conspecific with the Antarctic rather than the Colombian
taxon. Culmacanthid diplacanthoids have been described
from the oldest and youngest Aztec faunas (middle
Givetian) from Antarctica, and the latest Givetian Tagg-
erty-Howitt and Pambula River faunas of Australia (Young,
1993). The Colombian acanthodian is unlikely to be a
culmacanthid, however; none of the distinctive culma-
canthid cheek plates have been recorded in association with
the scales found in this locality. Also, scales from the only
articulated species, Culmacanthus stewarti [Long 1983],
have fewer and weaker crown ridges, and the fin spines have
broad smooth ridges separated by narrow grooves.
Several Middle – Late Devonian acanthodian taxa have
been erected for spines with radiating trabeculae that form
the middle layer for some or all of the exserted length. Of
these, Archaeacanthus quadrisulcatus [Kade, 1858]
(Middle Devonian, Baltic countries) is of a similar size to
the spine on MGCU-B.20; it has four or five smooth
longitudinal ridges per side, predominantly longitudinal
canals running through the middle layer proximally
(Lyarskaya, 1975), and an open pulp cavity for the proximal
third of spine length. However, spines of A. quadrisulcatus
are laterally flattened and sometimes have a subcostal canal.
Devononchus concinnus [Gross, 1940] (early Frasnian,
upper Gauja and Podsnetogor stages, Baltic countries) has
five smooth longitudinal ridges and an open pulp cavity for
the proximal third of the length but also a subcostal canal.
Because both the scales and spine from colombia differ from
previously described species, they are assigned to a new
taxon, Florestacanthus morenoi.
4. Conclusions
Janvier and Villarroel (2000) noted that the fish taxa
represented in the Cuche Formation (and the type of
assemblage in terms of relative abundance) were rather
suggestive of Euramerican vertebrate faunas of comparable
age, notwithstanding the presence of certain taxa hitherto
known essentially from Gondwanan regions (e.g. Antarc-
tilamna). They suggested that this faunal assemblage could
indicate a close geographical link between Euramerica and
northwestern Gondwana by Late Devonian times, in
accordance with geophysical models proposed by Dalziel
et al. (1994). In contrast, Young et al. (2000) described a
Bothriolepis-dominated Late Devonian fish fauna from
western Venezuela and concluded that it was of dominantly
Gondwanan affinity. Considering that the two localities,
Colombian and Venezuelan, belong to the same tectonic unit,
the Eastern Andean Block, the discrepancy between these
two conclusions seems surprising. However, Late Devonian
fish faunas from Gondwana and Euramerica are not radically
different. Only a few taxa seem restricted to one or the other
areas; e.g. the absence of the antiarch Asterolepis in
Gondwana, except possibly for Colombia. One Chinese
species that was assigned to Asterolepis sinensis is now
regarded as significantly different from the Euramerican
species and referred to the genus Jiangxilepis (Zhang and
Liu, 1991). One of the most striking examples of likely
Gondwanan endemism is that of phyllolepid placoderms,
which seem to have been restricted to Gondwana until well
after the Frasnian – Famennian boundary and then suddenly
invaded Euramerica by the late Famennian. With regard to
the acanthodians described here, the two forms closely match
both Euramerican and Gondwanan forms, but more precise
biogeographic relationships cannot be based on the new
material. The nektonic lifestyle of most acanthodians
probably assisted their wide dispersal, enhancing their
biostratigraphic potential but limiting their biogeographical
usefulness. Most of the Euramerican acanthodian taxa which
we have compared with the Colombian material are from the
late Middle – early Late Devonian of the Main Devonian field
(cf. Esin et al., 2000) on the East European platform, which
was in the tropics during the Late Devonian (Li et al., 1993);
the strata comprise clastic sediments, originating from the
Baltic Shield, with shallow water carbonate intercalations.
Notably, the depositional environment and latitude of those
Baltic ‘Old Red Sandstone’ facies and the Nostolepis sp. cf.
N. gaujensis-bearing red beds at Chanaruh, Iran, were
probably very similar.
It seems the composition of the Late Devonian fish
assemblages from Euramerica and the northeastern Gond-
wanan margin was chiefly constrained by their tropical
distribution. The Devonian fish assemblages of Colombia
and Venezuela appear radically different from those known
from the Early – Late Devonian of Bolivia, which are
generally associated with Malvinokaffric invertebrate com-
munities (Janvier and Suarez-Riglos, 1986; Gagnier et al.,
1988). The Devonian vertebrate faunas of Bolivia are
dominated by chondrichthyans (some of them strongly
endemic) and acanthodians, and placoderms are extremely
rare, with only an Eifelian rhenanid and a late Famennian
dunkleosteid known. Bothriolepid-dominated assemblages
are totally unknown in Bolivia, though favorable rock facies
occur. The same marked difference between northwestern
and central-southern South America is also clear when Late
Devonian plants are considered (Berry et al., 2000).
Whether this difference is due to tectonics (the Eastern
Andean Block may have been an allochtonous terrane that
linked Euramerica and western Gondwana in the Late
Devonian) or a climatic gradient is still undecided.
Moreno-Sanchez (2001), however, believes that both
factors are equally plausible and may have coexisted.
Acknowledgements
Part of the material described herein has been collected
in the field thanks to grant #6373-98 of the National
Geographic Society. The authors are grateful to Dr Mario
 C.J. Burrow et al. / Journal of South American Earth Sciences 16 (2003) 155–161
Moreno-Sanchez (Caldas University, Manizales) for the
loan of the material which he collected. CJB acknowledges
the support of an Australian Research Council Post-
Doctoral Fellowship.
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