New remarkable Late Jurassic teleosts from southern Germany: Ascalaboidae n. fam., its content, morphology, and phylogenetic relationships

Abstract

Complete morphological descriptions, as preservation permits, are provided
for a new Late Jurassic fish taxon (Ebertichthys ettlingensis n. gen. et n. sp.) and a revision of
some morphological features of Ascalabos voithii Graf zu Münster from the Solnhofen limestones,
southern Germany. A new family, Ascalaboidae, is erected to include the two
species. The new family is supported by numerous synapomorphies, e.g.,
maxilla with external row of small conical teeth increasing in size
posteriorly, absence of gular plate, low number of vertebrae (34 to 39),
deep and narrow supracleithrum – deeper than opercle, and vertebral centrum
formation of caudal region including paired chordacentra
(pseudo-diplospondyly) that fuse in early ontogeny forming one chordacentrum
that is later surrounded by an autocentrum. A phylogenetic analysis based on
173 characters and 42 taxa was performed. Following the phylogenetic
hypothesis, the sister-group relationship [Ascalabos + Ebertichthys] + more advanced teleosts
stands above the node of Leptolepis coryphaenoides plus more advanced teleosts and below the node of
Tharsis plus more advanced teleosts, and the new taxa are interpreted as extinct and
primitive forms within Teleostei. The new genus and species is endemic and
restricted to one Upper Jurassic locality – Ettling – whereas Ascalabos is known
from different localities in the Solnhofen limestones, with the exception of Ettling.

Full text

Foss. Rec., 19, 31–59, 2016
www.foss-rec.net/19/31/2016/
doi:10.5194/fr-19-31-2016
© Author(s) 2016. CC Attribution 3.0 License.
New remarkable Late Jurassic teleosts from southern
Germany: Ascalaboidae n. fam., its content, morphology,
and phylogenetic relationships
G. Arratia
Biodiversity Institute, University of Kansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, Kansas 66045, USA
Correspondence to: G. Arratia (garratia@ku.edu)
Received: 3 September 2015 – Revised: 6 December 2015 – Accepted: 9 December 2015 – Published: 18 January 2016
Abstract. Complete morphological descriptions, as preser-
vation permits, are provided for a new Late Jurassic fish
taxon (Ebertichthys ettlingensis n. gen. et n. sp.) and a re-
vision of some morphological features of Ascalabos voithii
Graf zu Münster from the Solnhofen limestones, southern
Germany. A new family, Ascalaboidae, is erected to in-
clude the two species. The new family is supported by nu-
merous synapomorphies, e.g., maxilla with external row of
small conical teeth increasing in size posteriorly, absence
of gular plate, low number of vertebrae (34 to 39), deep
and narrow supracleithrum – deeper than opercle, and ver-
tebral centrum formation of caudal region including paired
chordacentra (pseudo-diplospondyly) that fuse in early on-
togeny forming one chordacentrum that is later surrounded
by an autocentrum. A phylogenetic analysis based on 173
characters and 42 taxa was performed. Following the phy-
logenetic hypothesis, the sister-group relationship [Ascala-
bos+Ebertichthys]+more advanced teleosts stands above
the node of Leptolepis coryphaenoides plus more advanced
teleosts and below the node of Tharsis plus more advanced
teleosts, and the new taxa are interpreted as extinct and prim-
itive forms within Teleostei. The new genus and species is
endemic and restricted to one Upper Jurassic locality – Et-
tling – whereas Ascalabos is known from different localities
in the Solnhofen limestones, with the exception of Ettling.
1 Introduction
The fossiliferous localities of Bavaria, southern Germany,
represent one of the richest fossil fish localities and one of the
most extraordinary fossil Lagerstätte in the world (Arratia et
al., 2015). They are a succession of Upper Jurassic localities
(Kimmeridgian, Tithonian; Schweigert, 2015) with hundreds
of nominal plant and animal species (Schultze, 2015, table 5).
Unfortunately, as a common practice in the past, many of the
museum specimens are identified as from “Solnhofen”, with-
out a precise locality.
Late Jurassic fishes of Bavaria have been known for
over 200 years. The first illustrations appeared in the work
of Knorr (1755: pl. 23: figs. 2, 3; pl. 26a: figs. 1–4;
pl. 28: fig. 2; pl. 29: figs. 2–4). Later, fishes illustrated by
Knorr were giving scientific names by de Blainville (1818).
For further details see Arratia (1997) and Tischlinger and
Viohl (2015). The publication from de Blainville (1818)
opened the door to intensive research on fishes during the
19th century (e.g., Agassiz, 1833–1843; Graf zu Münster,
1834, 1839a, b, 1842; Wagner, 1861, 1863; Vetter, 1881;
Woodward, 1895) that slowed down in the first half of the
20th century with few contributions, e.g., Eastman (1914),
Biese (1927), and Weitzel (1933). After a gap in the study
of Jurassic fossil fishes of the Solnhofen limestones, Ny-
belin (1964, 1967, 1974) and Arratia (1987a, b) proposed
fundamental changes to the taxonomy of teleosts from the
region, and Taverne (1975a, b, 1981) and Patterson and
Rosen (1977) provided additional morphological informa-
tion of some of the teleostean species. Lambers (1992) listed
41 teleostean species, including pachycormiforms and aspi-
dorhynchiforms. Since 1992 that number has increased to at
least 62 nominal species (Schultze, 2015) and many others
remain to be described.
Some of the new teleosts are known from a locality – Et-
tling – that was modestly explored during the last years of the
20th century. Its first known fossil was a fish (Orthogoniklei-
thrus hoelli; Arratia, 1997), whose description was based
Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin.

32 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
on the 23 specimens recovered at that time. Now, the same
species is known from thousands of specimens, and other
species, most of them new, have been excavated and are cur-
rently under study. The first Jurassic crossognathiform from
the Northern Hemisphere (Bavarichthys incognitus; Arratia
and Tischlinger, 2010) and a new aspidorhynchiform (Aspi-
dorhynchus sanzenbacheri; Brito and Ebert, 2009) are also
known only from this locality. For a list of materials and
species from Ettling, see Ebert et al. (2015).
The goal of this contribution is to describe a new fish fam-
ily from the Solnhofen limestones that includes Ascalabos
Graf zu Münster and a new genus and species. The phyloge-
netic position of the new family is studied and discussed.
2 Materials and methods
The specimens here described are listed under their corre-
sponding descriptive sections. For a list of material used
in the phylogenetic analysis, see Arratia (2013). Specimens
here studied are deposited in the following institutions: CM,
Carnegie Museum, Section of Vertebrate Paleontology, Pitts-
burg, Pennsylvania; CMMH, Cleveland Museum of Natu-
ral History, Cleveland, Ohio; JME ETT, Jura Museum Eich-
stätt, Bavaria (ETT indicates that the fish was collected in
Ettling); JME SOS, Jura Museum Eichstätt, Bavaria (SOS in-
dicates that the fish was collected at an unspecified locality in
the Solnhofen limestones); NHM, Natural History Museum,
London; and SMNH, Swedish Museum of Natural History,
Department of Paleozoology, Stockholm.
Abrasive machines or chemicals were not used in the
preparation of specimens from Ettling because of their frag-
ile nature. All specimens studied herein were mechanically
prepared.
The author, using Wild FM 8 and Leica MZ9 stereomicro-
scopes equipped with a camera lucida, prepared illustrations
of the specimens. Thus, illustrations are based directly on
specimens rather than on photographs. Photographs are not
retouched with Photoshop. The latter was only used to label
figures.
2.1 Phylogenetic methodology
The phylogenetic analysis was conducted using PAUP
(Phylogenetic Analysis Using Parsimony) software (ver-
sion 4.0b10) for 32 bit Microsoft (Swofford, 2000). The char-
acter matrix was constructed using MacClade for the analy-
sis to run in PAUP. All characters are unweighted, unordered,
and considered to be independent of one another. The phylo-
genetic analysis used the list of characters and coding of Ar-
ratia (2013) with the addition of seven new characters (Sup-
plement 1). The outgroups used in the analysis are fossils and
extant neopterygians, e.g., the basal parasemionotiform Wat-
sonulus, the amiiforms Amia calva and A. pattersoni, and the
lepisosteiforms Obaichthys and Lepisosteus. The data matrix
with coding of characters is presented in Supplement 2.
2.2 Anatomical terminology
The terminology of the skull roof bones follows West-
oll (1943), Jollie (1962), and Schultze (2008). This termi-
nology is based on study of bones in Osteichthyes and dif-
fers from traditional skull roof terminology, which is funda-
mentally based on human anatomy and is not homologous.
To avoid confusion, all figures show in square brackets the
names of bones in the traditional terminology, e.g., parietal
bone [=frontal]: pa [=fr]. The terminology of the palato-
quadrate and urohyal follows Arratia and Schultze (1990,
1991).
The structure of the vertebral column is one of the ma-
jor sources of characters in the evolutionary history of
holosteans and teleosts (Schultze and Arratia, 1989; Arra-
tia et al., 2001). The term “vertebra” includes all serially
repeated ossified, cartilaginous, and ligamentous elements
around the notochord, consisting of the centrum, neural arch
and spine, and haemal arch and spine (Schultze and Arratia,
1988; Arratia et al., 2001). Vertebrae formed by a single cen-
tral element are called monospondylous, and those formed
by more than one element are called diplospondylous (see
Arratia et al., 2001, for further details). Mineralized or cal-
cified portions that develop inside the fibrous sheath of the
notochord or ossified portions that surround the notochord
can form the centrum. The centrum is termed chordacen-
trum, arcocentrum, or autocentrum, depending on its origin
and structure (Arratia et al., 2001, 103–106). These terms are
used here in the descriptions of the vertebral centra and also
in the lists of characters used in the phylogenetic analysis.
The terminology of the caudal endoskeletal elements
(e.g., preural centrum, ural centrum, and parhypural) and of
types of caudal skeletons (e.g., polyural or diural) follows
Nybelin (1963), Schultze and Arratia (1986, 1989, 2013),
and Arratia and Schultze (1992). “True” uroneurals are mod-
ified ural neural arches; “uroneural-like” elements are mod-
ified preural neural arches. These terms are included here
in the phylogenetic analysis, and their usage follows Arratia
and Lambers (1996) and Arratia and Schultze (2013). The
terms fin rays, scutes, fulcra, procurrent rays, epaxial rudi-
mentary rays, and principal rays follow definitions provided
by Arratia (2008, 2009). A rudimentary ray, as defined by
Arratia (2008), is a small ray with a short base not reach-
ing endoskeletal bones. It is positioned between the epaxial
basal fulcra and the first principal ray of the caudal fin. A
rudimentary ray may be distally segmented or not. This in-
terpretation of a rudimentary ray differs from that of Grande
and Bemis (1998, figs. 84, 86).
The terminology of scales follows Schultze (1966, 1996).
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 33
Figure 1. Distribution of Plattenkalk basins and reef areas in the southern Franconian Alb during the Early Tithonian (slightly modified from
Viohl, 1996). The new fish described herein was recovered in Ettling.
3 Systematic paleontology
Actinopterygii Cope, 1887
Teleostei sensu Arratia, 1999
Ascalaboidae n. fam.
Diagnosis (based on a unique combination of charac-
ters; uniquely derived features among basal teleosts are
identified with an asterisk [*]): Small teleosts of about
115mm maximum length. Maxilla with external row
of small conical teeth increasing in size posteriorly [*].
Supramaxilla 2 with a large, broad body and a narrow, long
dorsoanterior process covering most of dorsal border of the
long supramaxilla 1. Large, narrow opercle, almost 4 times
deeper than subopercle. No gular plate. Low number of ver-
tebrae, 34 to 39 [*]. Deep and narrow supracleithrum, deeper
than opercle [*]. Vertebral centrum formation of caudal re-
gion including paired chordacentra (pseudo-diplospondyly)
that fuse forming one chordacentrum that is later surrounded
by an autocentrum [*]. Broad first dorsal pterygiophore with
a peculiar fan-like shape with three or more anteroventral
processes [*]. Preural ural centrum 1 and ural centrum 1
with short neural spines. High number of hypurals, nine or
more. Six to eight uroneurals. Cycloid scales with circuli in
anterior field and no radii.
Derivation of name: The family name derives from the genus
name Ascalabos.
Content: Ascalabos voithii Graf zu Münster, 1839b and Eber-
tichthys ettlingensis, n. gen. et n. sp. (A few specimens pre-
liminarily identify as Ascalabos-like have been recovered in
Wattendorf and are in the process of being described. An-
other possible candidate is Anaethalion cirinensis Gaudant,
1968 from Cerin, France, which currently is under revision.)
Geographical distribution and age: Southern Germany,
Bavaria (Fig. 1). Upper Jurassic, Kimmeridgian (Wat-
tendorf), Kimmeridgian–Lower Tithonian? (Ettling), and
Tithonian (e.g., Eichstätt).
Ebertichthys n. gen.
Diagnosis (based on a unique combination of charac-
ters; uniquely derived features among primitive teleosts
are identified with an asterisk [*]): Small teleosts of about
90mm maximum length with dorsal-fin origin placed
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34 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
posterior to pelvic origin and equidistant to pelvic and anal
fins [*]. Large head, about 33% of standard length [*].
Large eye, about 35% of head length [*]. Large, heavily
ossified supraorbital bone, anteroventrally expanded. Large,
elongate antorbital [*]. Infraorbital 1 long, narrow, and
slightly broader anteriorly. Infraorbital 2 long and narrow,
forming most of the ventral orbital margin [*]. Infraorbital 3
relatively small, leaving most of quadrate exposed [*]. Elon-
gate ectopterygoid with a small tooth patch. Entopterygoid
with small conical teeth. Triangular, narrow preopercle with
ventral and posterior margins forming an angle of about
90◦[*]. Dorsal arm of preopercle almost reaching lateral
margin of skull roof. Preopercle with an expanded, rounded
flange at the level of infraorbital 3 [*]. Preopercular canal
with elongate, narrow sensory tubules in ventral and dorsal
arms. A massive, well-ossified cleithrum with dorsal arm
shorter than ventral one [*]. First anal pterygiophore long,
broadly expanded, and almost rectangular in shape [*].
Neural spine of preural centrum 1 shorter than neural spine
of preural centrum 2. Ural centrum 1+2 bearing two neural
arches with short spines. Eight or nine hypurals. First three
anterior uroneurals longer than posterior uroneurals 4 to 6.
Uroneural 1 extending anteriorly and reaching preural
centrum 2. One “urodermal” present. Body covered with
large cycloid scales.
Derivation of name: Ebertichthys is given in recognition of
the dedicated work that Martin Ebert (Eichstätt) has done
recently at Ettling, recovering fossil fishes and preparing
them with the suffix – ichthys (Greek) for fish. M. Ebert
collected the specimens described herein.
Ebertichthys ettlingensis n. sp.
Figures 2–13
2008 Undescribed small teleost. – Ebert and Kölbl-Ebert,
fig. 6.
2011 Ascalabos-like. – Ebert and Kölbl-Ebert, fig. 8.
2015 Teleost sp. 1. – Ebert et al., p. 21, fig. 13b; p. 22,
fig. 14a.
Diagnosis: Same as generic diagnosis.
Derivation of name: The specific name ettlingensis refers to
the locality of Ettling and its rich and beautifully preserved
fossils, from which the fishes were recovered.
Holotype: JME ETT 108a and 108b is preserved in part and
counterpart. It is a complete, beautifully preserved specimen
of about 75mm total length including soft anatomy preser-
vation (part of digestive system) and squamation (Fig. 2a).
The specimen is heavily ossified, so that it is interpreted as
an adult although the presence of unfused hypurals 1 and 2
and haemal arch of preural centrum 1 with their respective
centrum.
Paratypes: JME ETT 11, JME ETT 24 (Fig. 2c), JME
ETT 60, JME ETT 61, JME ETT 64a, JME ETT 132a, b
(Fig. 2b), JME ETT 148, and JME ETT 847a.
Type locality and age: Ettling (Plattenkalks I–III), Bavaria
(Arratia and Tischlinger, 2010; Ebert et al., 2015). Upper
Jurassic, probably upper Kimmeridgian (Ebert et al., 2015)
to lower Tithonian. However, the age of the locality is ap-
proximated due to the fact that well-preserved ammonoids
have not been recovered.
Description
General description: The fish is small, below 90mm total
length, slightly fusiform, and with dorsal, pelvic, and anal
fins producing a triangular outline, with the dorsal fin almost
equidistant to the pelvic and anal fins (Fig. 2a–c). Dorsal fin
insertion placed posterior to the midpoint of standard length
(61 to 64%). Pelvic fin insertion about at the midpoint of
standard length (about 52 to 55%). Anal fin insertion 69 to
73% of standard length (SL). Caudal peduncle deep, about
half of the maximum body depth. The head is proportionally
large, about 33% of standard length. In a young individual
(JME ETT 61) of 34.7mm total length, the head is compar-
atively larger (37% of SL). Eye large, 34 to 36% of head
length.
Skull roof and braincase: The skull roof is incompletely pre-
served or partially damaged in the studied specimens. Addi-
tionally, the skull is laterally compressed from burial, so that
it is difficult to describe particular elements. All bones of the
skull roof have smooth surfaces and are unornamented.
The main element of the skull roof (Fig. 3a–b) is the pari-
etal bone (i.e., frontal bone of traditional terminology) that
occupies most of the preorbital region and part of the postor-
bital region, which is short. The postorbital region is about
one-third of the preorbital length. Anteriorly, the parietal
bones suture with a broad and short mesethmoid (Figs. 3b
and 4b). The latter bone presents two short and narrow lat-
eral processes. Posteriorly, the mesethmoid sutures with the
parietals throughout several interdigitating projections of dif-
ferent lengths. The parietals apparently leave a long, narrow
space between them as a cranial fenestra, a condition that
also has been observed in Ascalabos voithii (Arratia, 1997).
A nasal bone is laterally placed to the anterior part of the pari-
etal bone. The bone seems to be narrow and elongate, mainly
carrying the anterior section of the supraorbital canal. Due to
poor preservation, the limits between the parietal, postpari-
etal, pterotic, and autosphenotic are not discernable, but the
postparietal bones seem to be small and sutured to each other
medially. The pterotic is very short and is the main element
that articulates with the hyomandibula. The pterotic sutures
with a small, triangular, and well-ossified autosphenotic ante-
riorly and with the parietal and postparietal medially. Its pos-
terior region is covered by an elongate, narrow extrascapula.
A short section of the middle pit line is observed, but it is
unclear whether the pit line crossed the pterotic or not. The
supraoccipital seems to be very small, with a very low crest.
Boundaries between these bones (Fig. 3a–b) are difficult to
discern due to the fragility and transparency of the bones.
Anteriorly and below the parietal, the autosphenotic su-
tures with a well-developed and ossified chondral bone, the
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 35
Figure 2. Ebertichthys ettlingensis n. gen. et n. sp. in lateral views. (a) Holotype JME ETT 108. (b) Paratype JME ETT 132a. (c) Paratype
JME ETT 24, specimen with a fish prey in its mouth (small individual of Orthogonikleithrus). Scales equal 1cm. Photographs courtesy of
M. Ebert.
pterosphenoid (Fig. 3b). Anterior to the pterosphenoid is
another chondral bone that has a median position, the or-
bitosphenoid. The latter extends anteriorly and ventrally, pro-
ducing an incomplete interorbital septum. The lateral eth-
moid is well developed, but its preservation does not allow
a proper description. A section of the parasphenoid is visi-
ble throughout the large orbit. It has a well-developed ascen-
dent process suturing with the pterosphenoid. Anteriorly, the
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36 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
parasphenoid sutures with the well-developed posterior pro-
cess of vomer (Fig. 3b); however, the suture between both
bones is unclear in other specimens (Fig. 4a–b). There are
numerous, minuscule teeth scattered below the parasphenoid,
vomer, and entopterygoid (Figs. 3b and 4b). Because of the
angle at which some of those teeth are preserved in JME
ETT 132a, it is possible to assume that not only the en-
topterygoid carried teeth but also the vomer and parasphe-
noid. This assumption should be tested when more speci-
mens become available. It is unclear whether a basipterygoid
process was present or not.
The supraorbital canal is a simple canal with a few
tubules. The short parietal branch (Fig. 3b) is rudimentary
and does not extend posteriorly into the postparietal bone.
The supraorbital canal does not produce a lateral tubule near
the autosphenotic; thus, a connection between supra- and in-
fraorbital canals is absent. Due to condition of preservation,
the otic canal was not observed in the studied material.
Orbit and circumorbital series: The fish has a large orbit
(Figs. 2a–c and 3a–b), so that the postorbital distance be-
tween the margin of the orbit and the preopercle is very nar-
row. This space is partially covered by the narrow and small
infraorbitals 4 and 5, which do not reach the preopercle pos-
teriorly, so that the dorsal part of the hyomandibula is ex-
posed laterally (Figs. 3b, 4a–b, and 5).
The series of circumorbital bones apparently encloses
the large orbit completely, although a dermosphenotic is
not preserved in the studied specimens. However, given the
space between the most posterodorsal infraorbital and the
autosphenotic, it is assumed that the bone was small. The
series includes a supraorbital, antorbital, and five (occasion-
ally six?) infraorbitals (Figs. 3b, 4a–b, and 5). Specimen JME
ETT 148 seems to have six infraorbitals (Fig. 5); however,
and by comparison with other specimens, I prefer to inter-
pret this 6th bone as result of a fracture of infraorbital 5, and
not as a dermosphenotic, because the infraorbital canal does
not turn posteriorly to join the otic canal. There are two large
sclerotic bones occupying anterior and posterior positions.
The supraorbital bone (Fig. 4a–b) is large, well ossified,
and expanded anteriorly. Only its anteriormost portion is pre-
served in the holotype (Fig. 3b). Because of its size and it
being displaced in all specimens, it is interpreted here as a
bone that covers most of the dorsal margin of the orbit.
The antorbital (Fig. 3b) is a slightly elongate, triangular
bone that is comparatively larger than antorbitals present in
other Jurassic teleosts. An antorbital branch of the infraor-
bital canal has not been observed.
Infraorbital 1 (Figs. 3b, 4a, and 5) is an elongate, narrow
bone that is slightly broader at its anterior margin than pos-
teriorly. Infraorbital 2 (Figs. 3b, 4a, and 5) is long and nar-
row, but becomes broader at its posteroventral corner, joining
infraorbital 3. The bone is so long that it is the main ele-
ment forming the ventral margin of the orbit. Infraorbital 3
(Figs. 3b, 4a–b, and 5) is a small, slightly square bone at the
posteroventral corner of the orbit. Its posterior margin just
reaches the anterior margin of the preopercle or both bones
are separated by a short distance. Due to the small size of
infraorbital 3, the quadrate is partially exposed laterally. In-
fraorbitals 4 and 5 form the posterodorsal margin of the orbit.
The infraorbital canal (Figs. 3b, 4a–b, and 5) is enclosed
by thin bone and is of simple type. The main canal is rela-
tively broad, with a few branching sensory tubules. At least
three tubules are counted in infraorbital 1. They open close
to the ventral margin of the bone. No sensory tubules branch
off in infraorbitals 2, 4, and 5. Infraorbital 3 has only one or
two sensory tubules that end at the middle region of the bone.
Upper jaw: Premaxilla, maxilla, and two supramaxillae form
the upper jaw. The premaxilla (Figs. 3b and 4a–b) is a slightly
triangular bone, with a short, broad ascendent process and an
elongate oral margin, which is as long as the elongate ante-
rior articulatory region of the maxilla. The oral margin bears
small, conical teeth. It is unknown how many rows of teeth
were present.
The maxilla (Figs. 3b and 4a–b) is elongate, partially cov-
ering the lateral aspect of the quadrate and extending close
to the posterior margin of the orbit. Its articulatory anterior
region is about a third of the maxillary length. The ven-
tral margin is slightly convex, as is its posterior margin. A
row of small, conical teeth that increase in size posteriorly
is present. The teeth are comparatively larger in younger
states than in older (larger) specimens. The posterolateral
side of the maxilla – which is well preserved in the holotype
– presents small, rounded tubercles (Fig. 3b), an uncommon
feature in Late Jurassic teleosts, which lack ornamentation.
Two supramaxillae (Figs. 3b, 4a–b, and 5) lie on the dor-
sal margin of the maxilla. Both bones together occupy the
whole length of the maxillary blade. Supramaxilla 2 has a
broad, expanded body and a narrow, long anterodorsal pro-
cess that almost covers the whole dorsal margin of supramax-
illa 1, which is slightly ovoid.
Lower jaw: The jaw (Figs. 3b and 4a–b) is moderately long,
with its articulatory region for the quadrate placed at the level
of the posterior half of the orbit. The jaw is formed later-
ally by two bones, the dentary and the angular. The suture
between both bones is not visible because of preservation,
although discontinuous sections are observed in some speci-
mens. From a narrow mandibular symphysis, the dentary ex-
pands posteriorly, producing together with the angular a mas-
sive and high coronoid process. A “leptolepid” notch has not
been observed, because the maxilla covers the region where
the notch would be placed. A retroarticular has not been ob-
served in any specimen, so that it is assumed that the angular,
articular, and retroarticular are fused in the medial side of the
jaw. The postarticular process is elongate and well ossified.
A narrow section of a chondral bone is medial to the ven-
tral margin of the angular in specimens JME ETT 60 and is
interpreted here as part of a branchial arch.
A surangular is not present at the posterodorsal corner of
the jaw. Coronoid bones are not present either.
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 37
Figure 3. Ebertichthys ettlingensis n. gen. et n. sp. in lateral view (holotype JME ETT 108). (a) Photograph of cranium and anterior part
of body. Photograph courtesy of M. Ebert. Scale equals 1cm. (b) Drawing of cranium and anterior part of body. Abbreviations: ant, an-
torbital; br, branchiostegal rays; cl, cleithrum; cor, coracoid; dpa, dermopalatine; ent, entopterygoid; exc, extrascapula; hy, hyomandibula;
io1-5, infraorbitals 1–5; iop, interopercle; l.de, left dentary; leth, lateral ethmoid; l.pmx, left premaxilla; met, mesethmoid; mx, maxilla;
op, opercle; orb, orbitosphenoid; pa[=fr], parietal [=frontal bone of traditional terminology]; par, parasphenoid; pcl1-3, postcleithra 1–3;
pop, preopercle; ppa[=pa], postparietal [=parietal bone of traditional terminology]; pt, pterotic; ptsp, pterosphenoid; ptt, posttemporal; qu,
quadrate; sc, scales; sca, scapula; scl, supracleithrum; scl.b, broken sclerotic bone; smx1–2, supramaxillae 1–2; sn, supraneurals; sorb.b,
broken supraorbital bone; sop, subopercle; sy, symplectic; r.de, right dentary; r.pmx, right premaxilla; vo, vomer.
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38 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 4. Ebertichthys ettlingensis n. gen. et n. sp. Crania in lateral views illustrating specific morphological features. (a) JME ETT 60.
(b) JME ETT 24. Arrow points to a small toothed patch. Abbreviations: ang, angular; aup, autopalatine; b.ant, broken antorbital; a.cer,
anterior ceratohyal; br, branchiostegal rays; cl, cleithrum; de, dentary; ect, ectopterygoid; ent, entopterygoid; ghy, glossohyal; d.hy, dorsal
hypohyal; iop, interopercle; hy, hyomandibula; io1–5, infraorbitals 1–5; l.ang, left angular; l.de, left dentary; l.pmx, left premaxilla; met,
mesethmoid; mtg, metapterygoid; mx, maxilla; op, opercle; par, parasphenoid; pmx, premaxilla; pop, preopercle; ptt, posttemporal; qu,
quadrate; ra, remnant of a branchial arch; r.de, right dentary; r.pmx, right premaxilla; scl, supracleithrum; scl.b, broken sclerotic bones;
smx1–2, supramaxillae 1–2; sorb, supraorbital; sop, subopercle; ur, urohyal; v.hy, ventral hypohyal; vo, vomer.
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 39
Figure 5. Ebertichthys ettlingensis n. gen. et n. sp. Infraorbital bones, preopercle, suspensorium, and part of upper jaw (JME ETT 148).
Abbreviations: hy, hyomandibula; io1–4, infraorbitals 1–4; io5?, broken infraorbital 5?; mtg, metapterygoid; mx, maxilla; pop, preopercle;
qu, quadrate; smx1–2, supramaxillae 1–2.
The mandibular canal is positioned near the ventral margin
of the jaw. Pores have not been observed in the posteroventral
region of the dentary and angular, so that it is assumed here
that the mandibular canal exits medially.
Palatoquadrate, suspensorium, hyoid arch, glossohyal, and
urohyal: Most of these bones are partially hidden by other
bones, so that a complete description is restricted to a few
elements. A small, somewhat square autopalatine (Fig. 4b)
is placed lateral to the vomer, and slightly dorsal to a small
dermopalatine. The latter bears very small, conical teeth.
The autopalatine and quadrate are separated by a thin, elon-
gate entopterygoid (Figs. 3b and 4b) that is covered medi-
ally with tiny conical teeth, at least at the anterior part of
the entopterygoid and below the orbit. The ectopterygoid
(Fig. 4b) is lateral to the entopterygoid and in front of the
quadrate. The bone is elongate, boomerang-like, and has a
small patch of tiny, conical teeth close to the palatine re-
gion. The quadrate (Figs. 3b, 4b, and 5) is slightly triangu-
lar, with its dorsal margin straight or slightly rounded, and
a comparatively small, slightly rounded condyle for articula-
tion with the lower jaw. Its posterodorsal margin sutures with
the metapterygoid. The length of the posterodorsal process of
the quadrate is unknown because the process is covered by
the anterior margin of the preopercle. A small portion of the
metapterygoid (Figs. 3b, 4a, and 5) can be observed below
the small infraorbital 3. Only a portion of the upper region
of the hyomandibula (Figs. 3b and 4b) and a portion of the
ventral region of the symplectic are observed, because those
bones are covered laterally by the posterodorsal infraorbitals
and the preopercle. The dorsal portion of the hyomandibula
(Figs. 3b, 4a–b, and 5) is narrow and apparently has one elon-
gate articular region with the braincase.
The lower part of the hyoid arch has preserved the hypo-
hyals, anterior ceratohyal, and a small portion of the poste-
rior ceratohyal. The posterior ceratohyal is partially observed
below the preopercle in specimen JME ETT 24. The dorsal
and ventral hypohyals (Fig. 4a–b) are slightly rectangular,
and the ventral one is slightly larger than the dorsal one. The
anterior ceratohyal (Fig. 4a–b) is rectangular, with an ovoid
“beryciform” foramen in its middle region, below the groove
for the hyoidean artery. The anterior and posterior borders of
the anterior ceratohyal are almost straight with no articular
interdigitations.
The glossohyal (Fig. 4a) is a small and elongate narrow
bone extending anterodorsally to the dorsal hypohyals. An
elongate, triangular urohyal (Fig. 4a–b) is preserved in speci-
men JME ETT 24. The bone is narrow anteriorly and expands
posteriorly producing three processes, the longest being the
middle one.
Opercular and branchiostegal series and gular plate: The pre-
opercle (Figs. 3b, 4a–b, and 5) is triangular-shaped, with
its posterior and ventral margins forming an angle of about
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40 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
90◦. The bone is not expanded posteroventrally, but it has a
characteristically slightly rounded flange just anterior to the
curvature of the preopercular canal. Its dorsal arm is long,
almost reaching the posterolateral margin of the pterotic,
whereas the ventral arm is short. All margins are smooth.
Due to conditions of preservation, the preopercular canal
and its tubules (Figs. 3b, 4b, and 5) are incomplete in most
specimens, with the exception of JME ETT 60 with sen-
sory tubules preserved in both the ventral and dorsal regions
(Fig. 4a). Apparently, the preopercular canal gives off seven
or eight tubules in the ventral arm and at least five or six
tubules in the dorsal arm. The tubules are very delicate, sim-
ple, and narrow, and they open close to the margins of the
bone. They are difficult to observe due to the transparency
of the bone, and this may explain the differences observed
among specimens illustrated herein.
The opercle (Figs. 3b and 4a–b) is large and dorsally is
broadly separated from the skull roof bones and the brain-
case. The dorsal margin is gently rounded, whereas the an-
terior and posterior margins are almost straight, and the ven-
tral margin is markedly oblique. The surface of the bone is
smooth with its anterior margin thickened and heavily ossi-
fied.
The subopercle (Figs. 3b and 4a–b) is a moderately narrow
bone. Its depth is 4 times less than in the opercle. Its ventral
margin is gently curved, and its anterodorsal process is short
and rounded. The interopercle (Figs. 3b and 4a) is covered
by the preopercle (or is not preserved) in most specimens,
so that a description cannot be presented. Apparently, the in-
teropercle is as long as the ventral margin of the preopercle
as shown by the holotype.
There are 10 incompletely preserved branchiostegal rays
(Fig. 4a) in specimen JME ETT 60; six narrow and short
branchiostegals articulate with the anterior ceratohyal, and
four broader ones articulate with the posterior ceratohyal.
Eleven rays are counted in the holotype, the first ones nar-
row and partially destroyed. Nevertheless, it is possible that
the number was much higher than 11. A gular plate has not
been observed.
Vertebral column and intermuscular bones: There are 38 or
39 vertebrae, including preural centrum 1; from these, 21 or
22 are abdominal or precaudals, so that the caudal region is
shorter than the abdominal one. The first four or five verte-
brae are covered laterally by the opercle. All vertebrae are
well ossified and with smooth lateral surfaces (Figs. 2a–c,
6a–b, 7, and 8). The centra are almost square, as deep as
long, with the exception of the preural centra that are slightly
deeper than long and of the ural centra, which are reduced in
size. All centra lack pre- and postzygapophyses.
The neural arches of the abdominal vertebrae are autoge-
nous, and the halves of each arch are unfused medially. The
neural arches are comparatively narrow and sit on the mid-
dle of the dorsal region of each centrum. Each neural arch
(Figs. 3b and 6a–b) is slightly expanded proximally, and it
has a slightly expanded anterior flange that gives the arch
a bilobate aspect. Each epineural process (Fig. 6b) emerges
at the posterolateral margin of the arch. Most of the neural
spines of the abdominal region are moderately inclined to-
ward the horizontal, and they are moderately long, just reach-
ing the ventral tips of the supraneurals. The spines are shorter
than the epineural processes. The parapophyses (Figs. 3b
and 6a–b) are fused to the anterolateral portion of the cen-
trum, near its ventral margin. Each is formed by a thin chon-
dral ossification with a prominent bony edge surrounding a
small articulatory cavity placed ventroposteriorly in the para-
pophysis. The head of each rib articulates with the small ar-
ticular cavity.
The neural arches of the caudal vertebrae (Figs. 2a–c, 7,
and 8) are fused to their centra, with the exception of the
first two or three caudal vertebrae (vertebrae 23 and 24) that
have autogenous neural arches. All haemal arches are fused
to their respective centra. The neural and haemal spines are
narrow and end in an acute tip, with the exception of those
of the preural centra. The neural and haemal spines are mod-
erately inclined toward the body axis, with slightly greater
inclination caudally. The haemal spines (Fig. 7) are short,
not extending between the anal pterygiophores, except for
the first two. The neural and haemal spines of the midcaudal
region are completely ossified (membrane bone), lacking an
internal core of cartilage; however, the condition changes in
the preural region (chondral bone; see below).
There are 18 or 19 unquestionable pairs of ribs. The first
two articulate with centra that are laterally covered by the
opercle, and the last pair is positioned lateral to the first
anal pterygiophore. All ribs (Figs. 2a–c, 3b, 6a–b, and 7) are
slightly curved, but the curvature is more evident in the ros-
tral direction. The ribs are well ossified and narrow along
their length, but slightly expanded at their proximal, small,
articulatory heads. Ventrally, they do not reach the margin
of the body. There is an unclear pattern between the last
centrum bearing a pair of ribs (Fig. 7), which is placed an-
terolaterally to the first anal pterygiophore, and the first cen-
trum with a complete haemal arch and spine that is posi-
tioned above the third anal pterygiophore (Figs. 7 and 8). Be-
tween those centra, there are three vertebrae with ventrolat-
eral structures that resemble parapophyses more than haemal
arches. Each is associated to an elongate, median structure
that is bifid at its proximal head and ends distally in two elon-
gate processes extending between the anal pterygiophores. If
these are modified ribs, then they occupy a median position,
but if these are modified haemal spines, then they are unique,
because they are bifid distally. I have not observed similar
structures in other fossil teleosts.
A complete series of about 16 supraneural bones (Figs. 2a–
c, 3b, and 6a) extends between the posterior part of the cra-
nium and the processes of the first dorsal pterygiophore. The
anteriormost supraneurals are larger and expanded antero-
posteriorly than the following ones, which become narrower
and smaller posteriorly. The supraneurals are slightly sig-
moid or curved. They are placed above or just between the
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 41
Figure 6. Ebertichthys ettlingensis n. gen. et n. sp. Details of certain abdominal vertebrae and pectoral fin. (a) Anterior abdominal region and
pectoral girdle and fin (holotype JME ETT 108). (b) Enlargement of a few abdominal vertebrae illustrating details of the epineural processes
(holotype JME ETT 108). (c) Details of pectoral fin and radials (JME ETT 132a). Abbreviations: cl, cleithrum; cor, coracoid; d.ra, distal
radials; epin.p, epineural process; na, neural arch; ns, neural spine; paph, parapophyses; pcl1–3, postcleithrum 1–3; pec.f, pectoral fin; pec.ra,
proximal radials supporting pectoral rays; pel.p, pelvic plate; ptt, posttemporal; ri, ribs; sc, anterior field of cycloid scales; sca, scapula; scl,
supracleithrum; sn, supraneurals; vc, abdominal vertebral centra; 1st. pec, first pectoral ray.
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42 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 7. Ebertichthys ettlingensis n. gen. et n. sp. Dorsal and anal fins and their relationships to the vertebral column (JME ETT 132a). Ab-
breviations: Ant, anterior; an.r, principal anal rays; epl.b, epipleural bones; dor.r, dorsal principal rays; epin.p, epineural processes; ha, haemal
arch; hs, haemal spine; la.pt, last anal pterygiophore; ld.pt, last dorsal pterygiophore; na, neural arch; ns, neural spine; paph, parapophyses;
pr.anr, anal procurrent rays; pr.dr, dorsal procurrent rays; ri, ribs; sn, supraneural bones; V21, vertebra 21; 1st a.pt, first anal pterygiophore;
1st d.pt, first dorsal pterygiophore.
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 43
Figure 8. Ebertichthys ettlingensis n. gen. et n. sp. Detail of vertebrae 21–23 and associated intermuscular bones (JME ETT 132a). Small
arrows point to a median bone that is bifid distally. Abbreviations: Ant, anterior; epin.p, epineural processes; epl.b, epipleural bones; ha,
haemal arch; hs, haemal spine; na, neural arch; ns, neural spine; V21, vertebra 21.
tips of the neural spines, but do not extend into the space
between neighboring spines.
The epineural processes of the neural arches (Figs. 2a–c,
3b, 6a–b, and 7) extend along the abdominal region ending
close to the end of the dorsal fin. The last pair may arise from
the neural arch of vertebra 22 or 23. The epineural processes
are long, extending laterally along the space occupied by five
or more centra. They are thin, heavily ossified, and curve pos-
terodorsally, following the angle of the neural spines, with
the exception of the last ones, which lie closer to the dorsal
margin of the centra than to the tips of the neural spines.
A short series of epipleural bones (Figs. 7 and 8) lies lat-
erally to vertebrae 18 to 26, in the hypaxial musculature. The
epipleurals are very thin, heavily ossified, and extend ventro-
caudally to the ventrolateral surface of the last ribs, haemal
arches and spines. They can be as long as the epineural pro-
cesses.
Ontogenetic development of vertebral centra: Each adult ab-
dominal centrum of Ebertichthys ettlingensis is formed by
a chordacentrum surrounding and constricting the notochord
and an autocentrum around the chordacentrum. In contrast,
each adult caudal centrum is formed by the chordacentrum,
the autocentrum, and the dorsal and ventral arcocentra that
are fused to the autocentrum producing a compact verte-
bra. In young specimens of Ebertichthys ettlingensis, the
caudal centra are unusual, because they present a pseudo-
diplospondyly. Each centrum presents two small chordacen-
tra or one large chordacentrum. The two hemicentra are sep-
arated vertically, and early in ontogeny they fuse into a sin-
gle chordacentrum. In contrast, the only centrum that forms a
chordacentrum in certain vertebrae presents a vertical separa-
tion that is lost early in ontogeny, so that only one centrum is
observed (Fig. 9a and b). This pseudo-diplospondyly is still
observed occasionally in some autocentra showing a verti-
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44 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 9. Ebertichthys ettlingensis n. gen. et n. sp. (a) Photograph of young specimen (JME ETT 61) in lateral view. Photograph courtesy of
H. Tischlinger, Scale equals 1cm. (b) Detail of last caudal vertebrae illustrating pseudo-diplospondyly and caudal endoskeleton incompletely
preserved. Abbreviations: chcPU4, chordacentrum of preural centrum 4; chcU1–3, chordacentra of ural centra 1–3; E, epurals (the first one
is missing); H1–7, hypurals 1–7; hsPU2, haemal spine of preural centrum 2; PH, parhypural; ‘UD’, “urodermal”; UN1–6, uroneurals 1–6.
cal “fracture” separating incompletely anterior and posterior
regions in a centrum.
A series of caudal chordacentra (Fig. 8b) is illustrated in a
specimen of 34.7 mm standard length of Ebertichthys n. gen.
A few centra show an elongate vertical gap on the lateral
wall of the chordacentra. Each caudal chordacentrum is pro-
portionally large, but its size reduces strongly caudally. Hy-
purals 1 to 4 are supported each by a chordacentrum, reveal-
ing a polyural condition in early ontogeny. Ural chordacen-
tra 1 and 2 fuse during ontogeny, then are surrounded by the
autocentrum producing a compound ural centrum 1+2 that
articulates with hypurals 1 and 2. Then, the three elements
become fused (see below).
Pectoral girdles and fins: The bones of the pectoral girdle
and fins (Figs. 3b, 6a and c) are very well preserved in a few
specimens, including the holotype, and this permits a detailed
description. The posttemporal is a large bone with an almost
rectangular body and a long and broad dorsal process articu-
lating with the cranium. The ventral arm is apparently short
because it was not observed in any of the specimens due to
the position of the preserved bone. The main lateral line ap-
parently is positioned along the main body of the posttempo-
ral and exits at its posteroventral margin. No sensory tubules
have been observed, only the main canal.
The supracleithrum (Figs. 3b, 4b, and 6a) is a narrow, long
bone extending from just at the level of the dorsal margin of
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 45
Figure 10. Ebertichthys ettlingensis n. gen. et n. sp. Pelvic plate and pelvic fin (JME ETT 108). Abbreviations: Ant, anterior; pel.r, pelvic
rays; p.pl, pelvic plate or basipterygium; p.sp, pelvic splint.
the opercle to the level of the subopercle, below the suture
between the opercle and subopercle and far below the level
of the vertebral column. Its posterior margin is slightly crenu-
lated in JME ETT 24. The lateral line is placed in the upper
middle of the bone and exits about the level of the vertebral
column.
The cleithrum (Figs. 3b and 6a) is a compact, heavily os-
sified bone with a short dorsal limb, a slightly expanded pos-
teroventral corner, and a narrower, short ventral limb inclined
anteroventrally. The anteromedial flange of the cleithrum is
expanded, almost rounded, giving the bone a characteristic
shape, unique among Jurassic teleosts.
Three postcleithra (Figs. 3b and 6a) are present. The first
one is placed at the junction between the supracleithrum and
cleithrum, the second one is partially covered by the pos-
teroventral expansion of the cleithrum, and the third one is
ventral to postcleithrum 2. Postcleithra 1 and 2 are ovoid
shaped, whereas postcleithrum 3 is styliform.
The scapula and coracoid (Fig. 3b) are preserved in the
holotype, but not the proximal and distal radials. The cora-
coid is slightly L-shaped and ends anteriorly in a moderately
broad and straight symphysis. Four proximal radials and a
few distal radials are preserved in JME ETT 132a (Fig. 6c).
The pectoral fin (Figs. 2a–c, 3b, and 6b–c) has a low po-
sition on the flank, near to the ventral margin of the body.
The total number of pectoral rays is unknown, but speci-
mens JME ETT 132a and JME ETT 148 have 15 rays pre-
served; the first one is spine-like and thicker than all other
rays (Fig. 6c), and with scarce segmentation only at its dis-
tal end. This ray is fused with the propterygium. All other
rays have very long bases and are only distally segmented
and branched. The inner rays become progressively shorter.
Pelvic girdles and fins: The pelvic girdles (Figs. 2a–c, 6a,
and 10) are well preserved in JME ETT 132. The basiptery-
gium (triangular pelvic plate) presents a thickened lateral
margin that expands anteriorly, while the inner region is
thinly ossified bone. The posterior part of the basipterygium
is thick and retains a large core of cartilage. A posterior pro-
cess is apparently missing. No proximal radials have been
observed. At least a short splint and nine rays are preserved
in JME ETT 132. A short splint and disarticulated rays are
preserved in the holotype (JME ETT 108a). Similar to the
pectoral rays, the pelvic rays have very long bases and are
only distally segmented and finely branched.
Dorsal and anal fins: The dorsal fin has three procurrent rays
and 12 principal rays. The first two procurrent rays are very
thin and short; the third one is almost the half of the size of
the first principal dorsal ray, which is only segmented and is
the largest of the series of rays. All principal rays have long
bases and are finely segmented and branched distally. The
first dorsal pterygiophore (Figs. 7, 9a, and 11a) expands an-
teroventrally and has one to three processes that are preceded
by a flat, almost rectangular bony flange that gives a charac-
teristic shape to the first pterygiophore. This element sup-
ports the procurrent rays, whereas the second pterygiophore
supports the first principal dorsal ray. Pterygiophores 2–4 are
of similar length, and they do not project ventrally between
the neural spines. Pterygiophores 5–12 decrease slightly in
size and thickness posteriorly, and the last pterygiophore is
markedly expanded, with a moderately long and narrow base.
As usual in teleosts, the last pterygiophore bears two small
rays, but these rays articulate in different positions with the
pterygiophore (Fig. 11b). The basal portion of the pterygio-
phores, except the first and last ones, have a lanceolate as-
pect, with thin, small anterior and posterior bony flanges.
They are apparently fused with the middle portion or middle
radial, whereas the distal portion or distal radial is partially
ossified in some of the last pterygiophores (Fig. 7).
The anal fin has, commonly, two or three procurrent and
11 principal anal rays that are supported by 11 pterygio-
phores. The procurrent rays (Fig. 7) are comparatively much
shorter than the first principal, which is only segmented. All
principal rays have long bases and are distally segmented and
finely branched. The first anal pterygiophore is a long, broad,
and flat rectangular bone that may support only the procur-
rent rays or the procurrents and part of base of the first prin-
cipal ray. The next two pterygiophores are slightly narrower
and shorter than the first one, and the remaining pterygio-
phores are much shorter and do not reach the distal tips of
the haemal spines. They are thinner than the first two ptery-
giophores, and because of the shape of their bases, they are
interpreted as formed by fusion with the proximal and mid-
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46 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 11. Ebertichthys ettlingensis n. gen. et n. sp. (JME ETT 64a). (a) Details of first two dorsal pterygiophores and first dorsal-fin rays.
(b) Detail of last dorsal pterygiophores. Arrows point to the two articular areas for fin rays. Abbreviations: Ant, anterior; 1st d.pt, first dorsal
pterygiophore; 2nd d.pt, second dorsal pterygiophore; ld.pt, last dorsal pterygiophore.
dle radials. The distal radials are mostly lost – probably be-
cause they were cartilaginous. The last anal pterygiophore is
broadly expanded distally and bears two minuscule rays in
comparison to the first principal rays.
Caudal fin and endoskeleton: The caudal fin and endoskele-
ton are preserved in several specimens, so that a detailed
description can be presented and intraspecific variation re-
ported. The caudal fin (Figs. 2a–c, 12, and 13) is deeply
forked with very short middle principal rays in comparison
to the long, leading marginal rays.
Five preural vertebrae support the caudal rays. All these
vertebrae are characterized by their smooth surfaces, and
their broad dorsal and ventral arcocentra are fused to their re-
spective centrum in larger specimens. They may retain rem-
nants of cartilage in some vertebrae. The neural spines of
preural vertebrae 2–5 are slightly expanded, and they have
a central core of cartilage surrounded by a thin perichon-
dral ossification and slightly developed anterior and posterior
bony flanges. In the vertebrae that are completely preserved,
it is possible to observe the anterior elongate processes at the
base of neural spines 1 to 4. Neural spines 2 to 5 are long,
whereas the neural spine of preural centrum 1 is shorter than
the preceding ones. The haemal spines of preural centra 1–5
are broader than their respective neural spines. However, the
haemal spine of preural vertebra 5 is narrower. The anterior
processes located between the base of the haemal arch and
spine are slightly rounded. The haemal spines of preural ver-
tebrae are chondral elements; some retain a core of cartilage.
In most specimens the neural and haemal arches of preural
vertebra 1 are fused to their centrum, but in specimen JME
ETT 108a, the haemal arch is still unfused. A complete neu-
ral arch, with its spine, is present on preural centrum 1. This
spine is shorter than the preceding spines. The haemal arch
and its broad parhypural are fused to the centrum in most
specimens (e.g., Fig. 13). A hypurapophysis on the lateral
wall of the haemal arch of preural centrum 1 was not ob-
served in any specimen.
Three or four ural centra (of the polyural terminology) are
associated with their respective hypurals. The first ural cen-
trum that bears hypurals 1 and 2 results from the fusion of
ural centra 1 and 2 (Figs. 9, 12, and 13). Ural centra 3 and 4
are associated with hypurals 3 and 4, respectively. An arch
and its spine and an incomplete arch are present above ural
centrum 1+2. A third incomplete arch is present in other
specimens (Fig. 13).
The complete number of uroneurals is unclear. Some spec-
imens have four (Fig. 12) or five (Fig. 13) or possibly six
(JME ETT 132a) uroneurals preserved. The first uroneural,
the longest of the series, extends anteriorly, reaching the
lateral surface of preural centrum 2. The second uroneural
reaches the lateral surface of preural centrum 1, and the third
uroneural reaches the lateral surface of ural centrum 1+2.
The fourth uroneural is short, reaching anteriorly the base
of hypural 6 or 7. There are other two smaller, elongate ele-
ments that I interpret as uroneurals 5 and 6. The first uroneu-
ral may have a small membranous outgrowth. There are three
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 47
Figure 12. Ebertichthys ettlingensis n. gen. et n. sp. Caudal fin and endoskeleton in lateral view (holotype JME ETT 108). Abbreviations:
Ant, anterior; d.scu, dorsal caudal scute; ebfu, epaxial basal fulcra; ffu, fringing fulcra; H1–7, hypurals 1–7; hsPU4, haemal spine of preural
centrum 4; nsPU2–4, neural spines of preural centra 2–4; naU1+2, neural arches of ural centrum 1+2; PH, parhypural; prp, procurrent
rays; PU1, 4, preural centra 1, 4; U1+2, 3, ural centrum 1+2 and 3 (polyural terminology) fused with the bases of hypurals 1 and 2; ‘UD’,
“urodermal”; UN1–4, uroneurals 1–4; PR1–19, principal rays 1–19; v.csu, ventral caudal scute.
elongate epurals occupying the space between the neural
spines 1 and 2 and the uroneurals; the first one is the longest.
At least nine hypurals are present. Hypurals 1 and 2 are
continuous at their bases, and they fuse to the autocentrum of
ural centrum 1+2 during ontogeny. Hypural 1 is the largest
element of the series, and hypural 2 is comparatively narrow.
Hypurals 2 and 3 are contiguous to each other, so that the
hypural diastema is very narrow or non-existent. Hypural 3
is the broadest among hypurals 3–9, which decrease in size
posterodorsally. Hypurals 2 and 3 are partially covered by the
expanded bases of the middle principal rays (e.g., 10 and 11).
There are eight or nine epaxial basal fulcra, three fring-
ing fulcra, 19 principal rays, three hypaxial procurrent-
segmented rays, and four or five hypaxial basal fulcra
(Figs. 12 and 13). One long and slightly fusiform or ovoid
dorsal scute and a ventral scute precede the epaxial and hy-
paxial series of basal fulcra, respectively.
The anterior epaxial basal fulcra in specimen JME
ETT 108a are apparently formed by paired elements
(Fig. 12); however, the anterior epaxial basal fulcra are un-
paired but with two separate, ventral projections in other
specimens. The basal fulcra are elongate, leaf-like elements
that expand laterally, partially covering the next fulcrum. The
three elongate fringing fulcra lie on the dorsal margin of the
first principal ray. They can be elongate and of similar size
(Fig. 13), or the first one is comparatively longer than the
other two (Fig. 12). The bases of the posteriormost basal ful-
crum and the first principal caudal ray produce an angle as
described for other fishes, mainly teleosts (Arratia, 2008).
Ten principal caudal fin rays are articulated with at least
hypurals 3 to 6. Hypural 6 supports the first principal ray,
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48 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 13. Ebertichthys ettlingensis n. gen. et n. sp. Caudal fin and endoskeleton in lateral view (JME ETT 60). Abbreviations: Ant, anterior;
d.scu, dorsal caudal scute; ebfu, epaxial basal fulcra; ffu, fringing fulcra; H1–6, hypurals 1–6; hsPU4, haemal spine of preural centrum 4; mo,
membranous outgrowth on anterodorsal margin of first uroneural; nsPU1–4, neural spine of preural centra 1–4; PU1,4, preural centrum 1, 4;
U1+2, ural centrum 1+2 (polyural terminology) fused with the bases of hypurals 1 and 2; ‘UD’, “urodermal”; UN1–5, uroneurals 1–5;
PR1–19, principal rays 1–19; v.csu, ventral caudal scute.
and the other nine rays are supported by hypurals 3 to 5. Nine
principal rays are supported by hypurals 1 and 2, parhypural,
and haemal spine 2. The articulation between segments of
the leading rays is mainly Z- or step-like, whereas the articu-
lation between segments of the inner principal rays is mainly
straight. Some of the bases of the middle principal caudal
rays are broken, but those that are preserved show that they
were expanded (Figs. 12 and 13). Dorsal processes associ-
ated with the bases of the middle principal rays of the upper
lobe have not been observed in the available material.
One elongate “urodermal” (sensu Arratia and Schultze,
1992) lies between the bases or close to the bases of the first
and penultimate principal rays. The “urodermal” is a thin,
oval bone (Figs. 12 and 13).
Scales: Thin, large cycloid scales with circuli at their ante-
rior field (Figs. 2a–b, 3a–b, and 6) cover the whole body.
No radii are observed. The scales are so large that approxi-
mately six rows of scales covered the skin of the flank below
the vertebral column and between pectoral and pelvic fins.
No remnants of scales are observed on the fin rays.
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 49
Ascalabos Graf zu Münster, 1839b
Diagnosis (emended from Nybelin, 1974, and Arratia,
1997). (The diagnosis is based on a unique combination of
characters. Uniquely derived features among basal teleosts
are identified with an asterisk [*].) Small teleosts of about
115 mm maximum length. Broad, simple cephalic sensory
canals with pointed sensory tubules [*]. With low number of
vertebrae, 34 to 39. Caudal centra, mainly midcaudals, with
a prominent lateral crest [*]. Seven or eight uroneurals, first
four longer than posterior series of three or four. First two
uroneurals reaching preural vertebra 3. Ten or 11 hypurals.
Few epaxial segmented procurrent rays posterior to last
epaxial basal fulcrum [*]. One rudimentary epaxial ray.
Caudal fringing fulcra absent.
Content. Only one species known, Ascalabos voithii.
Ascalabos voithii Graf zu Münster, 1839b
Figures 14–19
1839b Ascalabos Voithii – Graf zu Münster, p. 112, pl. XII,
fig. 5.
1843 Leptolepis Voithii – Agassiz, p. 131, 165, 295, pl. LXI,
figs. 2–3.
1843 Leptolepis polyspondylus – Agassiz, p. 133, pl. LXI,
figs. 7–8.
1843 Leptolepis paucispondylous (nomen nudum) – Agassiz,
p. 134.
1848 Leptolepis Voithii – Giebel, p. 143.
1848 Tharsis radiatus – Giebel, p. 146.
1863 Leptolepis Voithii, Wagner, p. 131, 134.
1895 Leptolepis voithii – Woodward, p. 512.
1974 Ascalabos voithi – Nybelin, 173–180, text-figs. 35, 36,
pl. XXIX, figs. 1–5, pl. XXX.
1975a Leptolepis (Ascalabos)voithi – Taverne, 233–243,
figs. 1–6.
1977 Ascalabos voithii – Patterson and Rosen, 151–152,
figs. 52–53.
1991 Ascalabos voithi – Arratia, 268–271, fig. 9.
1997 Ascalabos voithii – Arratia, 31–39, figs. 14–21.
Neotype: The holotype is lost. Nybelin (1974) designated as
neotype a specimen identified as Eichstätt I (currently JME
SOS 537).
Additional specimens: CM 9491. CMMH 9491. JME
SOS 2362, JME SOS 2364, JME SOS 2365, JME SOS 2458,
JME SOS 2483, JME SOS 2496, JME SOS 2497, JME
SOS 2886 (peels of scales), and many other specimens de-
posited at the JME. NHM 3672, NHM 3673a, NHM 37062,
and NHM 37080. SHL, collection of H. Leich (Bochum,
Germany) currently deposited in Tierpark und Fossilium
Bochum (Bochum). SMNH P5683. TM 6651, TM 10307,
and TM 10325. Additionally, one uncatalogued specimen be-
longing to U. Eller (Dümpelfeld, Germany) was included be-
cause of its excellent preservation.
Locality and geological time: Blumenberg, Eichstätt, Hof-
stetten, Kelheim, and Wintershof (Fig. 1) in Bavaria,
southern Germany. Upper Jurassic, Tithonian (Meyer and
Schmidt-Kaler, 1989, 1990; Schweigert, 2015). Ascalabos
voithii has also been cited from Cerin (France; de Saint-
Seine, 1949; Wenz et al., 1993), which is interpreted as Kim-
meridgian in age. This material is currently under revision.
Description
A description of Ascalabos voithii can be found in Arra-
tia (1997). Here, I only describe and illustrate additional in-
formation after of a re-study of the neotype and other speci-
mens. I describe some characters that add new interpretations
or a more complete understanding of certain structures. Some
are compared with the structures in Ebertichthys n. gen.
General description: Ascalabos voithii has a head proportion-
ally large, about 30% of standard length; however, the head
in Ebertichthys is comparative larger, about 34 % of SL. The
diameter of the eye is large, 22 to 25% of the head length
(Figs. 14b and 15b), whereas the eye in Ebertichthys ettlin-
gensis is even larger, 34 to 36 % of the head length. The in-
sertion of the dorsal fin (Figs. 14a, 15a, and 16) is above the
level of the insertion of the pelvic fins, about 52 to 58% of
SL. In contrast, the insertion of the dorsal fin is posterior to
that of the pelvic fins in Ebertichthys ettlingensis.
Cranial bones: The cranial bones are unornamented with
the exception of the premaxilla, which bears some small,
tubercle-like ornaments (Fig. 14c) covered with a thin layer
of ganoine in the neotype. The braincase of the neotype is
poorly preserved and does not add new information.
Circumorbital series: The supraorbital is incompletely pre-
served in most specimens or not preserved at all. It has an
expanded anteroventral region, similar to that found in Eber-
tichthys ettlingensis. Infraorbital 1 is an elongate bone that
joins with an elongate infraorbital 2 that is not as long as that
of Ebertichthys (compare Figs. 3b, 4a, and 5 with Fig. 14b).
Infraorbital 3 is a small bone at the posteroventral corner of
the orbit (Fig. 14b). Because of its reduced size, a significant
part of the quadrate is exposed. Although dorsoposterior in-
fraorbitals 4 and 5 as the dorsal part of the preopercle are not
preserved in the neotype, space left indicates that the region
between the inner margin of infraorbitals 3–5 and the anterior
margin of the preopercule was narrow.
Upper jaw: It contains the premaxilla, maxilla, and two
supramaxillae (Fig. 14b), which are all preserved in the neo-
type. The re-study of the neotype confirms that the premax-
illa has a short and small ascendent process as in the premax-
illa in Ebertichthys n. gen. However, the articulatory anterior
process of the maxilla is not as long as that in Ebertichthys, so
that the posterior blade of the premaxilla is shorter. The max-
illa is long and partially covers the anterior part of quadrate.
The two supramaxillae of the neotype are very similar to
that of Ebertichthys ettlingensis (compare Figs. 3b and 12b).
Supramaxilla 2 is broadly expanded posteriorly, forming the
main body of the bone. The anterodorsal process is very long
and narrow. Supramaxilla 1 is elongate and ovoid-shaped.
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50 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 14. Ascalabos voithii (neotype, JME SOS 537). (a) Photograph of neotype in lateral view. Photograph courtesy of M. Ebert. Scale
equals 1cm. (b) Cranium in lateral view and enlargement of premaxilla (c). Abbreviations: ang, angular; cl, cleithrum; de, dentary; ent,
entopterygoid; io1–3, infraorbitals 1–3; mx, maxilla; par, parasphenoid; pcl1, postcleithrum 1; pmx, premaxilla; pop, preopercle; ptt, post-
temporal; qu, quadrate; scl, supracleithrum; scl.b, broken sclerotic bone; smx1–2, supramaxillae 1–2; sop, subopercle; sy, symplectic.
Both supramaxillae cover the dorsal margin of the maxillary
blade.
Lower jaw: The mandible (Fig. 14b) is strongly ossified, and
although it is largely hidden by the maxilla and supramax-
illae, the large and high coronoid process is visible in the
neotype. The articulation of the mandible with the quadrate
is placed at the posterior margin of the orbit in the neotype
(specimen with almost closed mouth), which is in contrast
to an anterior position in specimens with the mouth open. A
“leptolepid” notch is present in the ascending dorsal margin
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 51
Figure 15. Ascalabos voithii from Schernfeld in the Eichstätt Basin. Uncatalogued specimen from U. Eller (Dümpelfeld). (a) Photograph of
specimen in lateral view. (b) Enlargement of the head. Note the peculiar shape and distribution of the sensory tubules of the preopercular
canal. Photographs courtesy of U. Eller.
of the dentary in specimen JME SOS 3228. This notch is not
observed in the neotype, because the maxilla covers this re-
gion. The “leptolepid” notch is narrow and partially forms
the anterior wall of the massive coronoid process.
Opercular bones: The preopercle (Figs. 14b and 15b) is in-
completely preserved in the neotype, and it is probably some-
how displaced. The important aspect to be noted is that the
bone presents a slightly rounded bony flange at its ante-
rior margin, similar to that in Ebertichthys n. gen. (compare
Figs. 3b and 4a with Fig. 14b). Since this feature has been
observed only in a few specimens due to poor preservation,
I have not considered it as a diagnostic character of the fam-
ily, but this feature should be re-evaluated when more speci-
mens of Ascalabos become available. The characteristic sen-
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52 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 16. Ascalabos voithii. Restoration in lateral view, updated from Arratia (1997).
sory tubules of the preopercular sensory canal are nicely pre-
served in the neotype (Fig. 14b), as well as in an additional
specimen illustrated in Fig. 15b.
Vertebral column: A long and prominent lateral crest
(Fig. 17) characterizes the centra of the caudal region of the
vertebral column in large specimens. The prominent crest
is still present on the lateral surface of preural centrum 2
(Fig. 18). This is very different to the condition in Eber-
tichthys with smooth centra. Ascalabos may have from 34 to
39 vertebrae (commonly 36), whereas in Ebertichthys the
number is commonly 38 or 39. These represent the lowest
counts among teleosts from the Solnhofen limestones (see
Arratia and Schultze, 2015). There are 16 to 19 pairs of ribs
(Fig. 16), whereas 18 or 19 pairs of ribs are present in Eber-
tichthys ettlingensis. A series of about 14 supraneurals ex-
tend between the occiput and below the first dorsal ptery-
giophores, whereas 16 supraneurals are commonly found in
Ebertichthys, but they do not extend below the first dorsal
pterygiophores.
The early formation of the centra of Ebertichthys ettlin-
gensis (Fig. 9) is similar to that of Ascalabos voithii, so
that the description above stands for both species. My ob-
servations (here and in Arratia, 1991, p. 257) and interpre-
tations differ from those of Schaeffer and Patterson (1984),
who interpreted the centra of young Tharsis,Ascalabos,
and lycopterid osteoglossomorphs as diplospondylous, each
formed by a precentrum and a postcentrum perichondrally
ossified, and that the chordacentrum is then reabsorbed or
suppressed in the middle of the centrum. These interpreta-
tions are not supported by my observations in young Tharsis,
Ascalabos, and Ebertichthys n. gen.
Pelvic plate or basipterygium: The basipterygium of Ascal-
abos is characterized by its slightly triangular shape and
its posterior expansion that has a short and sharp medial
posterior process (Arratia, 1997, Fig. 19c). In contrast, the
basipterygium of Ebertichthys ettlingensis lacks the posterior
expansion, and a medial process has not been observed.
Dorsal and anal fins and their supports: The first dorsal ptery-
giophore is characteristically expanded and with a few an-
teroventral processes in Ebertichthys ettlingensis (Figs. 7
and 11a), and a similar pterygiophore (Fig. 16) is present
in the neotype of Ascalabos. A first pterygiophore bearing
only two processes (JME SOS 2886) and another partially
broken one (HL 309) were illustrated for Ascalabos by Ar-
ratia (1997, Fig. 19a–b). Currently, I interpret these bones
as partially incomplete. According to the evidence gathered
here, the characteristic shape of the first dorsal pterygiophore
is now proposed as a diagnostic feature of the new family As-
calaboidae.
The first anal pterygiophore is long, broadly expanded and
almost rectangular in outline in Ebertichthys. In contrast, the
pterygiophore is thinner and more rod-like in Ascalabos as
well as in other Jurassic teleosts (see Arratia, 1981, 1997,
2000).
Caudal skeleton and fin: The caudal skeleton of the neo-
type confirms previous descriptions by Arratia (1991, 1997),
but adds some new information about variability. While in
some specimens the first two uroneurals extend anteriorly
to the lateral surface of preural centrum 3 (Fig. 18), they
reach preural centrum 2 in the neotype (Fig. 19). The neotype
presents eight uroneurals, whereas in other specimens only
seven have been observed, but this could be a result of poor
preservation. The fourth uroneural is broken in the neotype,
so that it is unclear whether it reached ural centrum 1+2
or not, as it does in other specimens. Uroneurals 5 to 8 re-
duce in size caudally. The neural spine of preural centrum 2
is slightly shorter than that of preural centrum 3, which is
broken distally. The neural arch of preural centrum 1 is in-
completely preserved.
There is an incomplete series of epaxial basal fulcra,
two segmented procurrent rays, one epaxial rudimentary ray,
19 principal caudal rays, three hypaxial procurrent rays, and
five hypaxial basal fulcra in the neotype (Fig. 19). Fringing
fulcra are absent. In contrast, other specimens have five or six
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 53
Figure 17. Ascalabos voithii (neotype, JME SOS 537). Midcaudal vertebrae illustrating the presence of a lateral crest and grooves on the
lateral side of the vertebral centra. Arrows point to a lateral longitudinal crest. Abbreviations: na, neural arch; ns, neural spine; ha, haemal
arch; hs, haemal spine; vc, vertebral centrum.
epaxial basal fulcra and seven hypaxial basal fulcra (Fig. 18).
Dorsal and ventral scutes precede the epaxial and hypaxial
basal fulcra, respectively.
Scales: The scales of Ascalabos are incompletely known, be-
cause it is difficult to observe isolated scales. The strongly
imbricate scales are usually preserved in situ. They are
slightly rectangular, with incomplete circuli in the ante-
rior field and with transverse lines that do not completely
cross the middle field of the scale. In contrast, Ebertichthys
n. gen. has large, slightly oval scales (Fig. 6b) with few cir-
culi in the anterior field. No transverse lines have been ob-
served.
4 Phylogenetic position of Ascalaboidae, n. fam.
To assess the phylogenetic relationships of Ascalabos and
Ebertichthys, I have performed a cladistic analysis of 42 ad-
vanced neopterygian taxa and 173 cranial, postcranial, and
scale characters. The list of characters (Supplement 1) is
from Arratia (2013) and includes additional taxon sampling
(Ebertichthys) and seven new characters. The coding of
173 characters is presented in Supplement 2. Polarization of
characters is based on outgroup comparison (five outgroups,
including fossil and recent taxa).
A parsimony analysis was performed using PAUP 4.0
beta 10 (Swofford, 2000), using ACCTRAN character-
state optimization, a heuristic search using a random ad-
dition sequence with 500 replicates, and the three bisec-
tion and reconnection (TBR) branch-swapping algorithm.
The parsimony analysis recovered one equally parsimonious
tree of 463 steps (retention index=0.7807; consistency in-
dex=0.4462; Fig. 20).
Figure 20 represents the only tree found. The topology
of this tree differs from that of Arratia (2013) in the rela-
tionships among the outgroups and in Node J (now includ-
ing Ebertichthys). Halecomorphs plus lepisosteiforms (out-
groups) appeared as sisters to each other in Arratia (2013,
fig. 95), whereas they appear now in a trichotomy with the
teleosteomorph clade. For some characters with a few ques-
tion marks, the parsimony analysis set forward some predic-
tions or assumptions as potential synapomorphies of certain
nodes. These assumptions are identified below. Characters
supporting nodes are listed in the caption of Fig. 20. Here I
only discuss the nodes representing the branching of Ascal-
aboidae n. fam.
Node J represents the branching of Ascalabos+Ebertichthys
plus more advanced teleosts and is supported by 10 synapo-
morphies: toothless parasphenoid (Ch. 25[1]); canals for oc-
cipital arteries in basioccipital bone absent (Ch. 29[1]*);
spiracular canal absent (Ch. 30[2]); no suborbital bone
(Ch. 47[0]); one supraorbital bone (Ch. 49[1]); absence of a
well-developed protruding lateral bony ridge extending along
an elongate dentary that separates dental and splenial regions
(Ch. 70[0]); midcaudal vertebral autocentra thick and sculp-
tured (Ch. 97[1]); walls of midcaudal centra with cavities for
adipose tissue (Ch. 98[1]*); notochord strongly constricted
by the walls of the centra (Ch. 99[1]); and epaxial basal ful-
cra or epaxial procurrent rays in close proximity to neural
spines, epurals, and posterior uroneurals (Ch. 145[1]).
The Node J1 corresponds to the branching of Ascala-
bos+Ebertichthys (Ascalaboidae n. fam.) and is supported
by eight synapomorphies, four of which are interpreted as
uniquely derived by the parsimony analysis: maxilla with
external row of small conical teeth increasing slightly in
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54 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
Figure 18. Ascalabos voithii. Caudal skeleton in lateral view slightly modified from Arratia (1997). Abbreviations: Ant, anterior; dpr,
dorsal or epaxial procurrent rays; d.scu, dorsal caudal scute; ebfu, epaxial basal fulcra; E1–3, epurals 1–3; H1–11, hypurals 1–11; hsPU2,
haemal spine of preural centrum 2; mo, membranous outgrowth on anterodorsal margin of first uroneural; mo’, membranous outgrowth on
anterodorsal margin of second uroneural; nsPU2–4, neural spine of preural centra 2–4; PR1,19, principal caudal rays 1, 19; rer, rudimentary
fin ray; U1+2+H1–2, ural centrum 1+2 (polyural terminology) fused with the bases of hypurals 1 and 2; ‘UD’, “urodermals”; UN1–7,
uroneurals 1–7; v.scu., ventral caudal scute.
size posteriorly (Ch. 170[1]*); less than 40 vertebrae present
(171[1]*); pseudo-diplospondyly present in the vertebral col-
umn of young growth stages (172[1]*); and first dorsal ptery-
giophore with three or more anteroventral processes, the
first one broadly expanded (Ch. 173[1]*). Other synapomor-
phies supporting this node are the presence of an antorbital
without antorbital sensory branch (Ch. 39[1]); median gular
plate absent (93[1]); a compound first pectoral ray not fused
with basal fulcra (Ch. 112[0]); hypaxial basal fulcra present
(Ch. 144[0]); and cycloid scales with circuli crossed by trans-
verse lines in the middle field (Ch. 157[1]). Although Eber-
tichthys is coded with questions marks for character 157, the
parsimony analysis predicts that transverse lines in the mid-
dle field of the scales cross the circuli. A similar prediction is
inferred from the parsimony analysis concerning the compo-
sition of the first pectoral ray in Ascalabos that is coded with
a question mark in the matrix.
The phylogenetic analysis interprets the following characters
as autapomorphies of Ebertichthys ettlingensis n. gen. et sp.:
midcaudal vertebral autocentra thick and smooth (Ch. 97[2])
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G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 55
Figure 19. Ascalabos voithii. Caudal skeleton in lateral view with only two epaxial basal fulcra preserved (neotype, JME SOS 537). Abbre-
viations: Ant, anterior; d.scu, dorsal caudal scute; dpr, dorsal or epaxial procurrent rays; ebfu, epaxial basal fulcra; E1–3, epurals 1–3; H1–4,
hypurals 1–4; hbfu, hypaxial basal fulcra; hsPU4, haemal spine of preural centrum 4; naPU1, neural arch of preural centrum 1; nsPU2–4,
neural spine of preural centra 2–4; PH, parhypural; PU1,4, preural centrum 1, 4; rer, rudimentary fin ray; sc, dorsal and ventral caudal scutes;
U1+2, ural centrum 1+2 (polyural terminology) fused with the bases of hypurals 1 and 2; ‘UD’, “urodermals”; UN1–8, uroneurals 1–8;
PR1–19, principal rays 1–19; vpr, ventral procurrent rays; v.scu, ventral caudal scute.
as opposed to the thick and sculptured centra typical in
most Late Jurassic teleosts or the thin and smooth centra
present in Leptolepides, and one tendon-bone “urodermal”
(Ch. 152[2]), in contrast to two found in Ascalabos and other
Late Jurassic teleosts.
The phylogenetic analysis interprets the following characters
as autapomorphies of Ascalabos voithii: epaxial basal ful-
cra or epaxial procurrent rays in close proximity to epurals
and posterior uroneurals (a reversal, Ch. 145[0]) and epaxial
procurrent rays present (Ch. 146[1]).
5 Final comments
A few of the most primitive Late Jurassic teleosts from
the Solnhofen limestones are known by single species that
are not currently assigned to families, such as Ascalabos
and Tharsis. Previously, and following the tradition of the
time, both genera were assigned to the family Leptolepidae
(e.g., Agassiz, 1843; Woodward, 1895) until the year 1974,
when Nybelin removed Leptolepis voithii and Leptolepis du-
bius from the genus Leptolepis and assigned them to new
genera, Ascalabos and Tharsis, respectively, within the fam-
ily Leptolepidae. Later, both genera were removed from Lep-
tolepidae and were left as incertae sedis among basal teleosts
(Arratia, 1997, 1999). The present study demonstrates that
Ascalabos voithii and Ebertichthys ettlingensis n. gen. et
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56 G. Arratia: New remarkable Late Jurassic teleosts from southern Germany
†
Anaethalion
Megalops
Heterotis
Hiodon
†
Lycoptera
†
Domeykos
†
Luisichthys
†
Protoclupea
†
Varasichthys
†
Ascalabos
†
Tharsis
†
Leptolepis coryphaenoides
†
Ichthyokentema
†
Dorsetichthys bechei
†
Ankylophorus
†
Siemensichthys macrocephalu
s
†
Siemensichthys siemensi
†
Lehmanophorus
†
Catervariolus
†
Eurycormus
†
Annaichthys
†
Knerichthys
†
Pholidorhynchodon
†
Pholidophorus gervasuttii
†
Pholidophorus latiusculus
†
Parapholidophorus
†
Pholidophoretes
†
Pholidoctenus
†
Prohalecites
†
Aspidorhynchus
†
Vinctifer
†
Belonostomus
†
Hypsocormus
†
Orthocormus
†
Pachycormus
Elops
Amia calva
†
Amia pattersoni
Watsonulus
†
Lepisosteus
Obaichthys
Elopocephala
Teleostei
†
Pholidophoridae
Teleosteomorpha
†
Ascalaboidae
†
Ebertichthys
F
E
D
C
B
A
M
L
K
J
H
I
G
L1
L2
L3
M1
M2
J1
C1
F1
A1
A2
A3
n. fam.
Figure 20. Hypothesis of phylogenetic relationships of the Late Jurassic Ascalabos and Ebertichthys n. gen. among the most primitive
teleosts (highlighted in bold). Synapomorphies supporting the main nodes are listed below; for a complete list see Arratia (2013, 114–
118). Uniquely derived characters are identified with an asterisk [*]. See Supplement 1 for descriptions of characters listed below. Node A
(teleosteomorphs or total-group teleost): 10[0]*, 24[1]*, 40[0]*, 47[2], 50[1], 94[0]*, 108[1], 112[1], 142[1]*, 164[1]*, 165[1]*, 168[1]*,
and 169[1]*. Characters 142 and 164–168 are soft anatomical features that are unknown in fossils, but the parsimony analysis predicts that
they were present at this phylogenetic level. Node B: 25[1], 34[1], 53[1], 55[1], 56[1], 59[1], 100[1], 109[1], 122[1], 125[1], and 162[1].
Node C: 47[1], 48[1], 59[2], 62[1], 63[1], 65[1], 69[1], 118[1], 148[2], and 153[1]. Node D: 78[1]*, 79[1], 102[1], 132[1], 150[1], and
151[1]. Node E: 13[1], 18[1], 56[0], 80[1]*, 96[1], 110[1], and 131[1]. Node F: 48[0], 66[1], 70[1], 82[1]*, 105[1], 108[0], 109[0], 127[1],
and 129[1]. Node G: 3[1], 25[0], 42[1], 64[1], 68[1]*, 108[3], 111[1]*, and 162[0]. Node H: 28[1], 30[1], 32[1], 56[1], 67[1], 69[2], 83[1]*,
100[0], 103[1]*, 121[1], 124[1], 140[1], and 149[1]*. Character 103[1] was incorrectly listed as a synapomorphy of node E in Arratia (2013).
Node I: 50[2], 107[1], 114[1], 119[1], 120[1], 144[1], 148[3], 152[1], 155[1], and 156[2]. Node J (Ascalaboidae, n. fam.): 25[1], 29[1]*,
30[2], 47[0], 49[1], 70[0], 97[1], 98[1]*, 99[1], and 145[1]. Node K: 86[1], 134[1], and 143[1]. Node L: 31[1], 33[1], 113[1], 116[1], 122[0],
141[1], and 146[1]. Node M: 36[1], 63[0], 65[0], 86[0], 104[1], 117[2], 132[3], 143[0], 136[1], 150[0], and 152[0]. Node M1: 25[0], 51[1],
56[0], 128[1], 133[1], 135[1], and 152[2]. Node M2: 31[0], 38[1], 41[1], 45[1], 49[2], 50[0], 59[1], 75[1], 91[1], 93[1], 112[0], 118[0],
123[0], 129[2], and 148[0].
Foss. Rec., 19, 31–59, 2016 www.foss-rec.net/19/31/2016/

G. Arratia: New remarkable Late Jurassic teleosts from southern Germany 57
n. sp. belong together in a new family, Ascalaboidae, that
is supported by numerous synapomorphies. Tharsis, a genus
that is currently under study, is not assigned to a family
yet. Specimens that were assigned to Leptolepis voithii from
Cerin, France, by de Saint-Seine in 1949 (in part according
to Nybelin, 1974) and to Anaethalion cirinensis by Gaudant
in 1968 are not considered until they are re-studied. Spec-
imens that have been preliminarily identified as Ascalabos-
like from Wattendorf wait for study and clarification of their
taxonomic status.
The Supplement related to this article is available online
at doi:10.5194/fr-19-31-2016-supplement.
Acknowledgements. I thank the following individuals and institu-
tions for permission to study material under their care: D. Berman
(CM); H. Bjerring (SMNH); M. Kölbl-Ebert and G. Viohl (JME);
A. Longbottom and the late C. Patterson (NHMUK). I especially
thank H. Leich (Bochum) and U. Eller (Dümpelfeld) for permission
to study some specimens belonging to their collections. Thanks to
the following individuals and institutions for permission to study
specimens included in the phylogenetic analysis: D. Berman (CM);
H. Bjerring (SMNH); R. Böttcher (SMNS); D. Butt (UCLA);
C. H. von Daniels (BGHan); W. Eschmeyer and D. Catania
(CAS); W. L. Fink and D. Nelson (UMMZ); H. Furrer (PIMUZ);
U. Göhlich (NHMW); L. Grande, W. Simpson, M. Westneat, and
M. A. Rogers (FMNH); H. Jahnke (GOE); M. Kölbl-Ebert and
G. Viohl (JME); the late K. Liem, K. Hartel, the late F. Jenkins,
and J. Cundiff (UMCZ); M. Louette and the late G. Teugels
(MRAC); D. Markle (OS); the late L. Martin and D. Miao (KUVP);
J. McEacharan and M. Retzer (TCWC); W. Mette and W. Resch
(Innsb.); A. Paganoni (MCSNB); L. Parenti and J. Williams
(USNM); the late C. Patterson and A. Longbottom (NHMUK);
T. Robins (UF); M. Röper (BMM-S); R. Rosenblatt (SIO);
W. Saul (ANSP); F. J. Schwartz (UNC); A. Simons and V. Hirt
(JFBM); D. Stacey and E. Holm (ROM); J. D. Stewart (formerly
at LACM); M. Stiassny and B. Brown (AMNH); R. Stucky
(DMNH); A. Tintori (University of Milan, Italy); P. Wellnhofer,
O. W. M. Rauhut and M. Moser (BSPG); F. Westphal and the late
R. Reif (GPIT); F. Witzmann (MB); E. O. Wiley and A. Bentley
(KUNHM); J.-Y. Zhang and M. Zhu (IVVP); and Irene Zorn
(GBA). Martin Ebert (JME) took the photographs of Ebertichthys
n. gen., as well as of the neotype of Ascalabos. J.-P. Mendau
(Berlin, Germany) prepared part of the final line illustrations based
on my original drawings. T. J. Meehan prepared the electronic
submission of all figures and revised the style of the manuscript. I
thank Jürgen Kriwet and an anonymous reviewer for reviewing the
manuscript and Hans-Peter Schultze for comments and suggestions.
Edited by: F. Witzmann
Reviewed by: J. Kriwet and another anonymous referee
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