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Bulletin of the Mizunami Fossil Museum, no. 37 (2011), p. 37–46, 4 fi gs., 2 tables.
© 2011, Mizunami Fossil Museum
In situ mud shrimps (Decapoda: Axiidea: Callianassidae) preserved
within their burrows from the middle Miocene of the Central Paratethys
Matúš Hyžný
Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University,
Mlynská dolina G1, 842 15 Bratislava, Slovakia<hyzny.matus@gmail.com>
Abstract
In situ preservation of mud shrimps of the family Callianassidae (Decapoda: Axiidea) has rarely been noted in the
fossil record. The present contribution reports body fossils of four members of the family (“Callianassa” almerai,
“C.” pseudorakosensis, “C.” sp. 1 and “C.” sp. 2) which are apparently preserved within their burrows, all from
middle Miocene deposits of Austria, Slovakia and Hungary (Central Paratethys). Description, with detailed figures
for each reported occurrence, is given, followed by a review of fossil “thalassinideans”, mainly Callianassidae,
preserved within burrow structures or associated with burrows. Occurrences of all middle Miocene callianassids of
the Central Paratethys currently known are also summarized.
Key words: Decapoda, Callianassidae, in situ preservation, middle Miocene, Central Paratethys
Introduction
The family Callianassidae has a robust fossil record.
Unfortunately, systematics based on hard-part morphology are still
debated. However, following publication of a paper by Manning
and Felder (1991), which highlighted, besides other “traditional”
characters, the taxonomic significance of major cheliped
morphology, many workers have attempted to assign fossil
callianassids more naturally to extant genera or, alternatively, have
erected new ones. In view of the delicate nature of the callianassid
exoskeleton, only chelipeds (which usually are heavily calcified)
are likely to be preserved in the fossil record (Bishop and
Williams, 2005).
Thirty-four extant callianassid genera are currently known (De
Grave et al., 2009). However, fewer than a quarter of these has a
fossil record which dates back beyond the Pliocene. This can be
ascribed both to preservational and collecting biases. It should also
be noted that many extant genera can be differentiated on the basis
of soft-part morphology only, so they are bound to remain
unrecognized in the fossil record.
Contrary to the rather robust fossil record of callianassid
chelipeds, remains of animals preserved inside their burrows are
relatively rare. However, Bishop and Williams (2005) noted that
the preservation of major and minor chelipeds in close proximity
might represent the remains of burrow-dwelling individuals,
because the chelipeds would have been dispersed if not protected
within a burrow.
The aim of the present contribution is to focus on the in situ
preservation of Callianassidae in the fossil record. For that reason,
no systematic and taxonomic issues relating to the nature of the
body fossils presented herein, are discussed. Detailed
re-descriptions of “Callianassa” almerai Müller, 1993 and
“C.” pseudorakosensis Lőrenthey in Lőrenthey and Beurlen, 1929
with emended diagnoses and a discussion of their systematic
affinities are currently being prepared by the author and will be
published elsewhere.
Decapods as producers of burrows in the fossil record
There are several higher taxa of decapod crustaceans which
independently evolved the construction of permanent burrows or
burrow systems. Unfortunately, identifying decapods as producers
of burrows without direct evidence of in situ preservation is rather
difficult. However, ichnofossils commonly attributed to decapod
crustaceans are very common, yet usually do not contain any body
fossils. Such associations are very rare, as a literature survey
shows.
Species that produce permanent burrows have been identified in
six decapod infraorders sensu De Grave et al. (2009). Among
carideans the family Alpheidae is known to build rather complex
burrow systems. Although no undisputed member of the family has
been reported from the fossil record (Schweitzer et al., 2010),
Radwański et al. (2006) recorded burrow systems from the middle
Miocene of Ukraine and interpreted them as having been
constructed by alpheid shrimps, although no body fossil evidence
had been recovered.
Among astacideans, a burrowing behaviour is quite common
(e.g. families Nephropidae, Cambaridae and Parastacidae).
38
Bedatou et al. (2008) described the ichnogenus Loloichnus from
Jurassic and Cretaceous strata of Argentina and interpreted it as the
remains of crayfish burrows. They regarded their possible maker to
have been a member of the family Parastacidae, again without any
body fossil preserved.
In the Glypheidea, a burrowing behaviour was identified in
extinct members of the Erymidae (Monaco and Garassino, 2001),
Glypheidae (Sellwood, 1971), and Mecochiridae (Neto de
Carvalho and Viegas, 2007; Neto de Carvalho et al. 2010). All
these reports described body fossils preserved in situ.
There are several brachyuran families whose members construct
rather simple burrows (e.g. Gecarcinucidae, Goneplacidae,
Portunidae, Panopeidae, Gecarcinidae, Sesarmidae, Varunidae,
Dotiliidae, Macrophthalmidae, Mictyridae and Ocypodidae).
Reports of fossil brachyurans preserved in situ within their burrows
are limited to goneplacids Ommatocarcinus corioensis (Creswell,
1886) from the lower Miocene–Pliocene of Australia (Jenkins,
1975), and Icriocarcinus xestos Bishop, 1988 from the upper
Campanian or lower Maastrichtian of California, USA (Bishop,
1988; Schweitzer et al., 2007) and longusorbiid Longusorbis
cuniculosus Richards, 1975 from the upper Campanian to lower
Maastrichtian of British Columbia, Canada (Richards, 1975).
The most complex burrow systems are constructed by members
of the former Thalassinidea which was subdivided into two
independent infraorders, Axiidea and Gebiidea (Robles et al.,
2009). Extant members of the Laomediidae, Thalassinidae and
Upogebiidae (Gebiidea), and of Axianassidae and Callianassidae
(Axiidea) are known to construct very complex burrow systems
which can reach more than 1 metre in depth.
The paucity of remains of “thalassinideans” found within their
burrows was already emphasized by Feldmann and Zinsmeister
(1984), Stilwell et al. (1997), and Bishop and Williams (2005).
There are several reports of Upogebia body fossils associated
with burrow structures (Karasawa and Inoue, 1992; Kato and
Koizumi, 1992; Ando and Karasawa, 2010). Fraaije et al. (2006)
reported two species of Upogebia preserved in nodules interpreted
as parts of burrows. The only unequivocal record of body fossils of
Upogebia in burrows is that from the Miocene of Japan (Kato,
1996).
Murray and Hanley (1986) gave an account of the famous
Australian subfossil mudlobster of the genus Thalassina which is
preserved in great numbers. They interpreted them as moults which
underwent rapid carbonate permineralization within the burrows.
The families Ctenochelidae and Callianassidae are sometimes
very difficult to distinguish one from another, especially when
exclusively dealing with fragmented chelipeds.
Of the family Ctenochelidae, Glaessner (1947) described
Ctenocheles bakeri (Glaessner, 1947) from the Eocene of Australia
associated with burrows. Later Rasmussen (1971) reported
Ctenocheles sp. from the lowermost Paleocene of Denmark
associated with burrows and Kato (1996) similarly described an
assemblage of Callianopsis spp. from the Miocene of Japan. Shinn
(1968, Pl. 111, Fig. 3) figured a complete specimen of “a shrimp,
the presumed architect, preserved in the burrow”, from Texas. It is
remarkably similar to Gourretia aquilae (Rathbun, 1935) from the
Turonian of Mexico (Vega et al., 2007). Both occurrences are from
the same lithostratigraphic unit (Eagle Ford Group), and may
therefore be conspecific.
There are several reports of callianassids preserved within their
burrow structures. Protocallianassa faujasi (Desmarest, 1822)
Table 1. Occurrences of fossil callianassid remains preserved within burrow structures or associated with burrows.
Taxon Age Country Remarks Major reference
Protocallianassa antiqua Turonian Czech Republic associated with burrows Kříž and Čech, 1974
Protocallianassa ex aff. antiqua Senonian Germany in a burrow Mertin, 1941
Protocallianassa faujasi Maastrichtian Netherlands in burrows Swen et al., 2001
Protocallianassa faujasi Campanian Germany in burrows Mourik et al., 2005
Protocallianassa faujasi Santonian Germany associated with burrows Förster, 1973
Protocallianassa faujasi Cenomanian Czech Republic claw fragment in a burrow Veselská, 2009
Protocallianassa faujasi Cenomanian Czech Republic associated with burrows Veselská, 2009
Protocallianassa mortoni Campanian Delaware (USA) associated with burrows Picket et al., 1971
Callichirus waagei Maastrichtian South Dakota (USA) cheliped in a burrow Crawford et al., 2006
Callichirus symmetricus Eocene Antarctica in burrows Schweitzer and Feldmann, 2000
Neocallichirus rhinos Eocene Mexico in a burrow Schweitzer and Feldmann, 2002
Vegarthron santiago Eocene Mexico in a presumed burrow Schweitzer and Feldmann, 2002
Podocalichirus grandis Pleistocene Japan in burrows Karasawa et al., 2006
“Callianassa” almerai Miocene Austria in burrows this paper
“Callianassa” burckhardti Maastrichtian–Danian Argentina asscoiated with burrows Feldmann et al., 1995
“Callianassa” pseudorakosensis Miocene Slovakia in supposed burrows this paper
“Callianassa” ?pseudorakosensis Miocene Poland associated with burrows Radwański and Wysocka, 2004
“Callianassa” sp. Campanian Texas (USA) claw fragment in a burrow Beikirch and Feldmann, 1980
“Callianassa” sp. Eocene USA in burrows Bishop and Whitmore, 1986
“Callianassa” sp. 1 Miocene Hungary in a burrow this paper
“Callianassa” sp. 2 Miocene Hungary in a burrow this paper
“Callianassa” sp. Miocene Austria claw fragment in a burrow Ehrenberg, 1938
39
preserved in situ was noted by several authors from the Upper
Cretaceous of Germany, the Netherlands and the Czech Republic
(Förster, 1973; Swen et al., 2001; Mourik et al., 2005; Veselská,
2009). Two species of Callichirus were described from in situ
occurrences in the Maastrichtian of South Dakota (Crawford et al.,
2006) and the Eocene of Antarctica (Stilwell et al., 1997;
Schweitzer and Feldmann, 2000). One more in situ Eocene
occurrence is known (Neocallichirus rhinos Schweitzer and
Feldmann, 2002) from Mexico. Schweitzer and Feldmann (2002)
also recorded Vegarthron santiago Schweitzer and Feldmann, 2002
from the Eocene of Mexico preserved in a presumed burrow
structure. So far, from Miocene strata there has been a single
record of in situ callianassid remains by Ehrenberg (1938). He
reported a claw fragment preserved at the blind end of a tunnel,
which was a part of a burrow system from the lower Miocene of
Austria. The present paper describes several additional examples
from Austria, Hungary, and Slovakia.
All occurrences of Callianassidae preserved within burrow
structures or associated with burrows known to the author,
inclusive of the ones reported herein are summarized in Table 1.
Geological and geographical settings
The material presented herein comes from several localities (Fig. 1):
1. Styrian Basin – Retznei quarry in Austria (Steiermark), where
strata of the Weissen-egg Formation are exposed. The age is early
“Badenian” (Langhian). For sedimentological details see Friebe
(1990). Decapod remains of several species were reported by
Schouppe (1949) and Flügel (1986); however, a large portion of
the decapod fauna present has not yet been described.
2. Vienna Basin – the Dúbravská hlavica locality in Slovakia
(vicinity of Bratislava), where strata of the Sandberg Member
(Studienka Formation) crop out, their age being late “Badenian”
(early Serravallian). For details on sedimentology see Baráth et al.
(1994).
3. Danube Basin – the Gyakorló and Rákos localities in Hungary
(Budapest area), where strata of the Rákos Formation are exposed,
whose age is late “Badenian” (early Serravallian). For
sedimentological data and details on the associated decapod fauna,
see Müller (1984) and references cited therein.
Palaeogeographically, all localities were situated within the
Central Paratethyan Sea during the middle Miocene. For details of
the current status of Miocene Central Paratethys stratigraphy
reference is made to paper by Piller et al. (2007). For details of the
palaeogeography and palaeobiogeography of the Central
Paratethys see Harzhauser and Piller (2007) and references cited
therein.
Middle Miocene Callianassidae of the Central Paratethys
To date, some thirteen species of Callianassidae have been
identified in middle Miocene strata of the Central Paratethys (see
Table 2). Some species have also been reported from other
European regions; however, these are not included here. The list
shows the relative diversity of Callianassidae within the same
palaeogeographical area during roughly the same time (i.e., middle
Miocene). The list should be considered preliminary because it
includes only occurrences mentioned in the literature or examined
personally.
According to personal observation it can also be stated that
many of the species listed can be accommodated within genera
other than Callianassa Leach, 1814 (sensu Ngoc-Ho, 2003), thus,
representing a mixture of several independent genera in several
subfamilies of the Callianassidae. However, such reassignment is
beyond the scope of the present contribution and will be
considered elsewhere. Therefore, the material presented herein is
treated under “Callianassa” as a nomen collectivum in the broadest
sense.
Fig. 1. Map showing the position of the study area
in the Carpathian–Pannonian Basin complex
of the Central Paratethys (modified from
Harzhauser et al., 2003). In grey are pre-
Neogene sediments and basement. Localities
studied: 1 – Retznei (Styrian Basin), 2 –
Dúbravská hlavica (vicinity of Bratislava), 3 –
Rákos and Gyakorló (Budapest area).
40
In situ preservation
The material presented herein is deposited in the collections of
the Natural History Museum of Slovak National Museum in
Bratislava, Slovakia (SNM Z) and of the Hungarian Natural
History Museum at Budapest, Hungary (M), as well as in the
private collections of Miroslav Hornáček at Smolenice, Slovakia
(MH), Pál Müller at Budapest (PM) and Gerhard Wanzenböck at
Bad Vöslau, Austria (GW).
Bishop and Williams (2005), in their review on taphonomy of
“thalassinidean” shrimps, used several terms which mirrored the
nature of their body fossil preservation. Body fossils preserved as
parts of burrows in the form of button-like disks were referred to as
burrow buttons (Bishop and Williams, 2005: 219). The material
presented herein is, however, preserved within these parts of the
burrows which can clearly be identified as trace fossils. Their
shape is not button-like, it rather corresponds with the real shape of
a burrow. Therefore, in such cases I use the term “burrow
structure”, and intentionally avoided using ichnotaxonomic
determination as nearly all the material was not collected by
myself and in most specimens the position of the burrow structure
in the section can no longer be precisely determined.
“Callianassa” almerai Müller, 1993 (Figs. 2A–G, 3A–D)
The material comprises at least five individuals apparently
preserved within their burrow structures and several other
fragmentary specimens from the Retznei quarry (Styrian Basin).
Specimens preserved inside the burrows are represented by
articulated chelipeds (cheliped disassociation unit sensu Bishop
and Williams, 2005), so the characters of ischium, merus, carpus,
propodus and dactylus can be observed (Figs. 2, 3). Remains of
both major and minor chelipeds are preserved in two cases (GW
RET04-010, GW RET01-014, Figs. 2E, 3B). In a single case, two
specimens are preserved within one and the same burrow structure
(Fig. 3B). Both of them are oriented in the same direction. In three
cases (GW RET93-021, GW RET04-010, GW RET01-014; Figs.
2B–F, 3A–B) individuals are preserved on one side of the burrow
structure, so probably the fossils are seen from the underside. A
similar preservation style was reported by Mourik et al. (2005).
One specimen (GW RET05-011, Fig. 2G) represents a propodus
positioned at right angle with the burrow structure. Another one
(GW RET96-009, Fig. 3D) is preserved in the middle of the
burrow tube. For this specimen it is clear that the cheliped of an
individual did not lie on its side during the proces of fossilization,
suggesting that rather might be a live animal that was buried.
Similarly, in one propodus a geopetal texture (sparite) can be
observed (GW RETXX-017, Fig. 3C) documenting that the
specimen did not lie on its side either when fossilized.
The burrow structures are preserved in carbonate facies. Their
cross section is rounded (Fig. 2), although in one specimen it is
rather elliptical (GW RET01-014, Fig. 3A).
The species “Callianassa” almerai was until recently known
only from the middle Miocene of Spain (Müller, 1993). The
present report extends its geographic distribution to Austria, i.e., to
the Central Paratethys.
Material described by Müller (1993) comprises a dozen or so
specimens represented by isolated propodi only. He reported
remains of both right and left major chelae and erroneously
interpreted the species to be isochelous. However, in axiideans
both right- and left-handed individuals occur and “C.” almerai is
no exception. Material from Retznei documents specimens with
both chelipeds preserved and, indeed, minor chelae are also present
(Figs. 2E, 3B). This also allows assignment of two propodi which
were described and figured by Müller (1993) as “Callianassa” sp.
and which come from the same locality as “C.” almerai. Their
morphology corresponds with that of minor chelipeds of “C.”
almerai reported herein.
Table 2. Middle Miocene occurrences of Callianassidae reported from the Central Paratethys (occurrences reported in open nomenclature as “Callianassa”
sp. are not included).
Taxon Geographic distribution within the CP area Major reference
“Callianassa” almerai Müller, 1993 Austria (Styrian Basin) Müller, 1993
“C.” brocchi Lőrenthey, 1898 Hungary (Budapest area), Slovakia (Vienna Basin) Müller, 1984
“C.” chalmasii Brocchi, 1883 Hungary (Budapest area) Müller, 1984
“C.” fl oriana Glaessner, 1928 Austria (Styrian Basin) Glaessner, 1928
“C.” kerepesiensis Müller, 1976 Hungary (Budapest area, Novohrad Basin) Müller, 1984
“C.” cf. kerepesiensis Müller, 1976 Hungary (Budapest area), Poland (Carpathian Foredeep) Müller, 1984
“C.” munieri Brocchi, 1883 Hungary (Budapest area), Austria (Vienna Basin., Styrian Basin) Müller, 1984
“C.” norica Glaessner, 1928 Austria (Styrian Basin) Glaessner, 1928
“C.” pseudorakosensis Lőrenthey in Lőrenthey
& Beurlen, 1929
Hungary (Budapest area, Novohrad B.), Romania (Gr. Hungarian
B.), Slovakia (Vienna Basin), Poland (Carpathian Foredeep) Müller 1984
“C.” cf. pseudorakosensis Lőrenthey in
Lőrenthey & Beurlen, 1929 Austria (Styrian Basin) Glaessner, 1928; Müller, 1998
“C.” roztoczensis Müller, 1996 Poland (Lublin Upland), Hungary (Budapest area) Müller, 1996
“C.” sismondai A. Milne-Edwards, 1860 Austria (Styrian Basin) Glaessner, 1928
“C.” szobensis Müller, 1984 Hungary (Börzsöny Mts.) Müller, 1984
Callichirus bertalani Hyžný and Müller, 2010 Hungary (Bakony Mts.) Hyžný and Müller, 2010
41
“Callianassa” pseudorakosensis Lőrenthey in Lőrenthey
and Beurlen, 1929 (Figs. 4A, B)
The material consists of several burrow structures with preserved
animal remains from the Dúbravská hlavica locality in the vicinity
of Bratislava. In one case the articulated remains of three
individuals are preserved (SNM-Z-21.373, Fig. 4A), in the other
case two individuals are present (MH specimen, Fig. 4B). Both
chelipeds together with scattered remains of other pereiopods are
preserved of one individual (Fig. 4A), which can be characterized
as cheliped and thoracopod disassociation units sensu Bishop and
Williams (2005). Interestingly, individuals are oriented in the same
direction in both depicted specimens. Although the burrow walls
are not discernible, from the arrangement of the individuals it may
be assumed they were preserved lying on one side (possibly on the
bottom of a burrow).
Many more specimens of this species preserved with scattered
remains of pereiopods in close association with chelipeds as
burrow buttons sensu Bishop and Williams (2005) were collected
from the locality.
The burrow structures are preserved in a poorly consolidated
sandstone. Their cross section is difficult to determine; however,
the mode of preservation of the individuals suggests preservation
within the burrow (Bishop and Williams, 2005).
At the locality very abundant remains of “Callianassa” brocchi
Lőrenthey, 1898 co-occur with very rare Calappa heberti Brocchi,
1883.
“Callianassa” pseudorakosensis is very common in the area of
the Central Paratethys. Up to now it has been reported from
Hungary (Lőrenthey and Beurlen, 1929; Müller, 1984), Romania
(Müller, 1984), Austria (Müller, 1998), and Slovakia (the present
contribution). Recently, Radwański and Wysocka (2004) reported a
single chela associated with Ophiomorpha from the middle
Miocene of Poland. However, the morphology of that specimen is
rather unusual for “C.” pseudorakosensis, so it might represent
Fig. 2. “Callianassa” almerai Müller, 1993: A–C – a near-complete right major cheliped with the remains of the minor one within the burrow structure (GW
RET93-021); D–F – remains of the right major (ischium, merus and carpus preserved) and left minor cheliped within a burrow structure (GW RET04-
010); G – an isolated left major propodus preserved perpendicular to the burrow structure (GW RET05-011). Note the rounded cross section of the
burrows. All specimens come from the Retznei quarry (Styrian Basin, Austria). All scale bars represent 10 mm.
42
another species.
“Callianassa” sp. 1 (Figs. 4C, D)
The material from the locality Rákos MR8 (sensu Müller, 1984)
represents one burrow structure with a preserved fragment of
propodus inside it (M 2010.511.1.1; Figs. 4C, D). The cross
section of the burrow is elliptical. The same shape was also
observed in other burrow structures (without any fossilized
inhabitants) collected from the same locality (pers. observation).
Because of the fragmentary nature of the specimen, specific
identification is impossible.
“Callianassa” sp. 2 (Figs. 4E, F)
The material from the locality Gyakorló (in the Budapest area)
represents one burrow structure with the remains of two individuals
(PM specimen, Figs. 4E, F). The burrow structure is preserved in a
sandstone. Its cross section is elliptical (Fig. 4F); however,
compaction cannot be ruled out.
The material is too fragmentary for specific identification,
although it should be mentioned that one individual possessed
equal or subequal chelipeds (not visible on figures). Such chelipeds
are known for instance in the genera Eucalliax Manning and
Felder, 1991 or Calliaxina Ngoc-Ho, 2003 of the subfamily
Eucalliacinae. However, without more complete material a generic
assignment is impossible.
At both Hungarian localities mentioned herein several species of
Callianassidae have been recorded (for details see Müller, 1984).
Discussion and conclusions
The material presented herein represents very rare occurrences
of fossil callianassids preserved in situ within their burrows. As
stated by Bishop and Williams (2005), callianassids might enter the
preservation process as corpses, moults or disassociation units.
They regarded most mortality of these animals as “resulting from
drastic causes such as voluminous shifting sediments, poisoning by
Fig. 3. “Callianassa” almerai Müller, 1993: A–B – two individuals preserved within the same burrow structure (GW RET01-014), note propodi of minor
chelipeds as indicated in the interpretive drawing; C – an isolated left major propodus (GW RETXX-017) with geopetal texture (sparite) in its upper
part (black arrow); D – remains of a left major cheliped preserved within a burrow structure (GW RET96-009). All specimens come from the Retznei
quarry (Styrian Basin, Austria). All scale bars represent 10 mm.
43
red tides, or severe and prolonged changes in oxygen levels,
salinity, etc.” (Bishop and Williams, 2005: 224). In general it is
assumed that members of the Callianassidae moult and die within
their burrow systems, however, often it is virtually impossible to
recognize moults from corpses.
It can be assumed that there are two main types of preservation
in the material described and figured herein:
1) preservation of supposedly dead animals or moults lying on
their side when entering the fossilization process. They are
preserved on one side of the burrow structures, so when observed
the underside is seen. The remains sank down with the heaviest
exoskeleton part, the large cheliped, to the bottom of the burrow
Fig. 4. “Callianassa” pseudorakosensis Lőrenthey in Lőrenthey and Beurlen, 1929: A – three individuals within the same burrow structure (SNM-Z-21.373),
note the propodus in oblique view indicated by a black arrow; B – remains of major chelipeds of two individuals within the same burrow structure (MH
specimen). “Callianassa” sp. 1: C–D – a burrow structure with a fragment of propodus preserved inside it (M 2010.511.1.1). “C.” sp. 2: E–F – propodi
of two individuals within a burrow structure (PM specimen), note the propodus indicated by a white arrow. Localities: A, B – Dúbravská hlavica (Vienna
Basin, Slovakia), C, D – Rákos (Budapest area, Hungary), E, F – Gyakorló (Budapest area, Hungary). All scale bars represent 10 mm.
44
tunnel, so the large claw is always visible. This is the case in
material of “C.” almerai (Figs. 2A–F; 3A, B) and “C.”
pseudorakosensis as well (4A, B). Roughly the same mode of
preservation when the tracemaker is preserved on the side of a
burrow (i.e. at its bottom) was reported and figured by Mertin
(1941: Fig. 30.1), Shinn (1968: Pl. 111, Fig. 3), Swen et al. (2001:
Figs. 5.5, 5.6), and Mourik et al. (2005: Pl. 2).
2) preservation of animals in the middle of the burrow. Some
specimens of “C.” almerai (Figs. 2G, 3D) and “Callianassa” sp. 1
(Figs. 4C, D) represent cheliped fragments which are not preserved
on the side of the burrow structure. In one case (Fig. 3D) the
cheliped is positioned in the burrow longitudinally in its centre. It
may represent an animal which was buried alive, however, the
fragmentary nature of the specimen does not allow further
speculation. Preservation of cheliped parts positioned at right
angles to the burrow tunnel (Figs. 2G, 4C, D) may indicate a
vertical or subvertical shaft of a burrow or a horizontal or
subhorizontal tunnel filled with debris together with exoskeleton
remains. Unless the exact position of a preserved burrow tube
within the section which yielded them is known, no further
conclusions can be drawn. Unfortunately, this information is
unavailable in the case of the specimens presented herein.
The preservation of several in situ individuals in the same
direction has already been reported in Protocallianassa faujasi
(Desmarest, 1822) from the lower Campanian of Germany (Mourik
et al., 2005). They interpreted them as remains of presumably dead
animals, although there are difficulties to explain how such an
arrangement can originate. They hypothesized that the dead
individuals were carried away and stored in dead-end tunnels by
living ones. Such systematic removal of exoskeletal fragments
(moults or corpses) into disposal chambers that subsequently may
be closed off was mentioned also by Bishop and Williams (2005);
however, this behaviour has never been directly observed in extant
species (P. Dworschak, pers. comm., 2010). Unfortunately, the
material presented herein cannot shed new light on this issue due to
the imperfect preservation and limited samples.
Acknowledgements
I wish to thank G. Wanzenböck (Bad Vöslau, Austria), P. M.
Müller (Geological Survey of Hungary, Budapest) and M.
Hornáček (West-Slovakian Museum, Trnava, Slovakia) for
permission to study material in their collections, J. W. M. Jagt
(Natuurhistorisch Museum Maastricht, the Netherlands) for
improving the English of an earlier draft and the final manuscript;
and R. M. Feldmann (Kent State University, Ohio, USA) for
literature items. A. Kroh and M. Harzhauser (Naturhistorisches
Museum, Wien, Austria) helped process the map of the Neogene
basins (Fig. 1). M. Jamrichová and J. Kučerová (Comenius
University, Bratislava) helped with scanning and processing
composite figures. The manuscript benefited from the thorough
review and constructive comments of R. M. Feldmann. This work
has been supported by research grants APVV 0280-07 to D.
Reháková (Comenius University, Bratislava), UK/353/2009 and
PRIFUK/19/2010 to M. Hyžný.
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Manuscript accepted on October 20, 2010
