Bite marks on tests of the echinoid genus Echinocorys LESKE, 1778 from the Maastrichtian of
Hemmoor (NW-Germany)*
Detlev Thies
Institute of Geology and Paleontology of the University Hanover
Callinstra
DFe 30, D-3000 Hanover 1
translated by Richard Forrest
Abstract
Feeding traces caused by fish are found on echinoid tests of the genus Echinocorys from the
Maastrichtian of Hemmoor (NW-Deutschland). These traces were first described by GRIPP (1929)
and interpreted as the bite-marks of ray. A re-examination of the traces is undertaken on the basis
of 28 marked tests, which have been collected more recently in Hemmoor. The ray theory is
refuted, and it is demonstrated that the marks were more probably made by a bony fish of the genus
Enchodus or Cimolichthys (Family Enchodontidae).
I. Introduction
Echinoids are attacked by a wide range of predators, especially fish and crustaceans, and
have developed a number of different strategies (for example spines, endobenthonic lifestyle, etc)
to avoid them (SMITH 1984). The attacks of predators are not always successful. In cases of
failure, traces of the predatory attack can remain on the surface of the echinoid test ("Traces of
predation", BISHOP 1975). In the living echinoids, such traces are known on the sand dollar (genera
Encope and Leodia) (KIER & GRANT 1965). Feeding traces on fossil echinoids were described by
GRIPP some time ago (1929) on test of Echinocorys from the Campanian of Lägerdorf and the
Maastrichtian of Hemmoor (NW Germany). At that time GRIPP observed two different types of
traces on the echinoid test. While the tests from the Campanian of Lägerdorf show many short,
irregular notches at the edge between oral surface and flank, tests from the Maastrichtian of
Hemmoor show a different pattern of injuries. This characteristic pattern is of partly punctuated
and partly continuous tracks, found on the sides of the test near the apex. Extensive new
collections of Echinocorys tests from the Maastrichtian of Hemmoor (collection F. SCHMID,
Niedesächsishes Landesampt für Bodenforschung in Hanover) confirms GRIPP’s observations
respecting the characteristic feeding traces on Echinocorys from Hemmoor. However, GRIPP’s
interpretation of these tracks can as injuries sustained by repeated bites of a ray – a view on which
ABEL (1935) had already declared doubts – is no longer adequate. A new interpretation of the
finding needs to be sought.
II. Material
28 tests of Echinocorys were used for the study of feeding traces. All the specimens come
from the non-operational quarry of Hemmoor Zement AG (TK25 2220 Cadenberg and 2320
Lamstedt). The section profile of SCHULZ & SCHMID (1983) shows a sequence from the sumensis
zone (Lower Maastrichtian) to the baltica/danica zone (Upper Maastrichtian). The tests bearing
*
Original citation: Thies, D. 1985. Bisspuren an Seeigel-Gehdusen der Gattung Echinocorys LESKE, 1778 aus dem
Maastrichtium von Hemmoor (NW-Deutschland). Mitt. Geol. Paläont. Inst. Univ. Hamburg; 59:71-82.
feeding traces were distributed throughout the sequence, the majority of finds being from the
argentea/junior zone (24 examples from the sections F 915 to F 938 of the sequence of S CHULZ &
SCHMID 1983).
All specimens described here are the in the collection of F. SCHMID (NLfB, Hanover). This
collection contains approximately 930 coronae of Echinocorys from Hemmoor. The proportion of
injured tests is about 3%.
III. Description of the Marks
A very detailed description of the traces is found in GRIPP (1929). Briefly, these are in part
furrows and in part individual punctures on the surface of the echinoid test, located around the
apex. (Plate 1, Fig. 1, 2, 4, 5; Plate 2, Fig. 1). Both the furrows and the punctures are normally
found in groups of three (more rarely to two-or four).
Furrows
The furrows have a width of approximately 1 mm. They are found generally in groups of
three, parallel to each other, and spaced at about 2-3 mm.
Punctures
The punctures are spindle-shaped in profile with a longitudinal diameter of approximately 1
mm, and found in groups of three spaced about 2-3 mm apart.
GRIPP noted that the channels and punctures have an average spacing of 3.5 mm. I do not
consider this difference to be significant. Presupposing that the same predator on Echinocorys is
represented in all cases, the different sizes of the feeding traces can be explained by different ages
and sizes of the predator.
Without exception, each group of three impressions is matched by a second group. If one
looks from apically at the corona, it is some centimeters separated on the opposite side (Plate 1,
Fig. 5; Plate 2, Fig. 1; A-A, B-B). It can itself be two groups of furrows (Plate 1, Fig. 4) or two
groups of punctures (Plate 2, Fig. 1; also Plate 1, Fig. 1) or a single group of furrows or punctures
(Plate 1, Fig. 5). A standard unit therefore always consists of two groups of three symmetrically
arranged, and of the same type. The furrows are vertically, while the punctures are aligned parallel
to the axis. In the present material, the maximum spacing between the two groups is approximately
5.5 cm (Plate 1, Fig. 4). Several standard units always overlap on the echinoid corona (seen
especially on Plate 1, Fig. 1).
GRIPP’s description can be supplemented with following observations:
It is noticeable that a group of three punctures (even taking into account the curvature of the
echinoid test) never lies completely in a level. If one connects the two opposite puncture groups,
the middle-line is displaced towards the apex (Plate 2, Fig. 3) or to the oral side of the corona (Plate
1, Fig. 2). In groups of furrows the middle furrow is much longer than the outside ones (Plate 1,
Fig. 6). The newly collected reveals from Hemmoor show that is the characteristic feeding traces
are found not only on the apical but also on the oral surfaces of some tests (Plate 2, Fig. 2).
As GRIPP determined, the tracks have healed almost without exception. This is shown by
necrotic growths on the surface of the damaged calcareous plates (Plate 1, Fig. 3). The predator has
not been able to penetrate or break the echinoid test. Either the aggressor broke off the attack, or
the echinoid found some alternative means of escape, so that it survived the attack with a largely
undamaged test. In an example from the Hemmoor material it can be seen that the echinoid had a
chance of survival even if the test was largely destroyed. Plate 2, Fig. 2 shows a specimen shows
furrowed feeding-traces that stop suddenly at an edge on the oral surface on both sides of a pit. The
test area between the traces was obviously broken away. However it regenerated, though not to the
level of the surface of the test, leaving a clear scar.
Feeding traces on the Echinocorys test that demonstrate a successful attack are almost
unknown in the Hemmoor material. This can be explained in that a successful attack of this
predator on Echinocorys was likely to lead to the total destruction of the test. (In this respect states
the predominance of healed tests tells us nothing about the success rate of the predator. Only in
exceptional cases can a more or less completely test destroyed can give information only on the
cause of death of the echinoid). An example from the material present here (Plate 2, Fig. 4) has a
part of the apical surface of the test is indented. The tracks on the preserved sides of the corona
order are arranged around the depressed area and healed. They consist of single groups of
punctures and furrows – their respective counterparts would be to on the destroyed part of the test.
The broken rim of the test is overgrown with ephisites (bryozoans) (Plate 2, Fig. 5). It is significant
that the growth of ephisites on the edge of the break has damaged the test before the entombment in
the sediment. The unhealed tracks show that this echinoid was the victim of an attack it did not
survive.
Finally, it must be noted that the traces described above appear only the on test of the genus
Echinocorys. The genera Cardiaster FORBES, 1850 and Galerites LAMARCK, 1801, which are also
to be found in the Maastrichtian of Hemmoor, do not show these injuries.
IV. Interpretation of the Marks
Where traces consist of two opposite parts they can be interpreted easily as bite marks of a
fish, with which the two injuries are the tooth rows of upper and lower jaw. While the punctures
are the consequence of the penetration of teeth into the echinoid test, the furrows are caused by a
bite slipping off the test.
Some characteristics of the biting fish can be derived directly from the type of the injuries:
1) The spindle-shaped outline of the punctures shows that the teeth that penetrated into the shell
were shaped lance-shaped.
2) The longitudinal diameter of the punctures of approximately 1 mm show that the teeth were
of considerable size.
3) Because of the similarity of the damage, the upper and lower jaws must have had similar
dentitions.
4) Since itself the impressions (punctures) of the tooth row of upper and lower jaws on the
echinoid test in rows are parallel to the axis of the bite, these teeth must have been in the
most anterior part of a rounded, not a sharp or beak shaped jaw.
5) The notable spacing up to approximately 5.5 cm between the marks of upper and lower jaws
shows a wide gape, on big jaws and with it on a relatively big animal.
The fish-fauna of the northwest German Upper Chalk consists of sharks, rays and bony fish.
To which of these three groups can the fish which produced the bite marks be assigned? The fact
that no more than 3-4 teeth of a jaw have never penetrated simultaneously into the curved echinoid
test and that the upper and lower jaws were several centimeters wide led GRIPP (1929) to the
assumption that only big and laterally oriented jaws could have produced the tracks. He thought
that the bearers of such jaws must be certain ray-like fishes. However, this assumption can no
longer be upheld for several reasons. Admittedly, rays (= Superorder Batoidea) possess jaws of a
certain size orientated laterally; however more detailed investigation shows the symphyseal region
of the upper jaws is incurved, while the lower jaw at this position is curved outwards (S TEHMANN
1978). Jaws of this structure would not leave uniform traces as found here. Furthermore, most rays
possess exclusively blunt crushing teeth. Where pointed teeth are found, these have a pointed,
conical crown, which would not have left a spindle-shaped impression (BIGELOW & SCHRÖDER
1953, CAPPETTA 1972). Study of selachian teeth from the Maastrichtian of Hemmoor by H ERMAN
(1982) has shown moreover that the Batoidea here are extremely rare and represented only by the
family Rhinobatidae (banjo fish, up to now only a single tooth). The banjo fish have crushing teeth
with a small indented crown and prefer small fish, crabs and mollusks as food (BIGELOW &
SCHRÖDER 1953). This precludes rays as originators of the bite marks. More significantly, it is
already a candidate for the originator of the marks in the sharks. Superficially, a predator/prey
relationship between a shark and Echinocorys is possibly, especially since HERMAN (1982)
identified the teeth of 17 shark-genera with more or less lance-shaped teeth in the chalk of
Hemmoor. In the majority of cases, the morphology of these teeth (several cusps per tooth with
Notidanus, diagonally-orientation of the cusps and diagnostic heterodont with the Squaliforms)
does not match the marks on the echinoid tests. In other forms (Parasquatina, Scyliorhinus) it is
known that living representatives of these taxa do not feed on echinoids (BIGELOW & SCHRÖDER
1948, CADENAT & BLACHE 1981). Paraginglymostoma and Hemiscylliums from Hemmoor can be
eliminated on the basis of their small size alone. Heterodontus is the only serious candidate as
possible predator on Echinocorys. Heterodontid sharks, as durophageous feeders typically prey on
shelled molluscs and crustaceans as well as echinoids (RIPE 1976, WHITLEY 1940). The extreme
rarity of Heterodontus in the Maastrichtian of Hemmoor (up to now only 2 teeth have been found,
HERMAN 1982) clearly refutes this assumption.
Consequently, only the bony fish remain as conjectural originators of the bite marks.
Observations of living bony fish show that these animals regularly prey on echinoids; in some
genera they even form the main component of the food (FRICKE 1971, EDGE-ALL 1967). A
particular study of the bony fish fauna of the Maastrichtian of Hemmoor is not yet available.
HERMAN (1982) mentions however numerous enchodontid teeth (Fam. Enchodontidae) from this
locality. The actinopterygian remains from the Campanian and Maastrichtian of the adjoining
Westphalian and Dutch-Belgian area have however been studied in detail (ALBERS & WEILER
1964; MARCK 1858, 1863, 1876,1885; SIEGFRIED 1954). WOODWARD (1902-1911) covered the
bony fish of the English Upper Chalk. Outside Europe, rich bony fish faunas are known from the
Maastrichtian of North-Africa and the Congo (ARAMBOURG 1952, DARTEVELLE & CASIER 19431959). For a closer approach to the question, a list of characteristic of the fish is considered. A
combination of the characteristics 1, 2, 4 and 5, big fish with wide gape and big, lance-shaped teeth
in the most anterior of the jaw is found among the bony fish of the Upper Chalk in the family
Enchodontidae, and I assume therefore that the originator of the bite marks belongs to this family.
(Characteristic 3 – corresponding dental structure in upper- and lower jaws – is often lost through
incomplete conservation of fossil fish.) The family Enchodontidae is represented in the Campanian
and Maastrichtian of northwest Europe by the genera Enchodus, Palaeolycus, Cimolichthys and
Apateodus (ALBERS & WEILER 1964, SIEGFRIED 1954). Palaeolycus, with its pointed muzzle, is
characteristically an ambush predator. By analogy to living ambush predators, for example the garpikes (Fam. Belonidae), its food will have consisted predominantly of smaller fish and nektonic
invertebrates (NIKOL’SKII 1961). With Apateodus, only small, inconspicuous teeth are fond in the
anterior part of the jaw; larger teeth are found only in the posterior part (KRUIZINGA 1923,
WOODWARD, 1902-1911). I consider it therefore likely that the producer of the marks is belongs
either to the genus Cimolichthys or the genus Enchodus. In contrast to Apateodus, the most anterior
parts of the jaws contain large, dagger-like teeth in these two genera (SIEGFRIED 1954, WOODWARD
1902-1911). An unambiguous generic identification appears impossible at the moment. It needs
detailed knowledge of the dental morphology of the individual Enchodus and Cimolichthys types,
which cannot be learned from the predominantly fragmentarily preserved fossil remains.
V. Acknowledgements
Professor Dr. F. SCHMID (NLfB Hanover) inspired the work on bite marks. The
photographs were taken by H. AXMANN (NLfB Hanover). D. J. WARD (Orpington, England) has
corrected the English legends.
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