Contributions to Zoology, 72 (2-3) 173-179 (2003)
SPB Academic Publishing bv, The Hague
New studies of decapod crustaceans from the Upper Jurassic lithographic limestones of southern Germany
Günter Schweigert¹ & Alessandro Garassino²
2 1 Staatliches Museum fiir Naturkunde, Rosenstein I, D-70I91 Stuttgart, Germany; Museo civico di Storia naturale, Corso Venezia 55, 1-20121 Milano, Italy
Keywords:: Crustacea, Decapoda, lithographic limestones, Upper Jurassic, Solnhofen, fossil record, diversity
Abstract Introduction
The Upper Jurassic lithographic limestones ofsouthern Germany TheUpper Jurassic lithographic limestones in south- have long been known for their exceptional preservation of ern Germany outcrop at numerous localities are of decapod crustaceans (Glaessner, 1965), similar to the Upper and differentage setting and span an area of sev- Cretaceous of Lebanon (Hakel, Hadjoula) and the still poorly eral hundreds of kilometers (Fig. 1, Table 1) with known Callovian strataat La Voulte-sur-Rhône (France). In these localities in the Frankische Alb often summarized non-bioturbatedlimestones, the decay of decapod skeletons is
that mineralized and reduced, so besides the heavily chelae as ‘Solnhofen Lithographic Limestones’. Many fos-
often even delicate structures such as pleopods and carapace sils, both in old and new collections (fossil traders),
antennae are preserved. Recently, new decapod material has which is are labeled ‘Solnhofen’, highly mislead- been obtained from both scientific and commercial excavations, ing and precludes recognition of evolutionary trends in part in reopened lithographic limestone quarries.
I. Fossiliferous lithographic limestones in southern Germany.
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Table I. Age assignments for decapodcrustacean-bearing litho- in the Regensburg (Ebenwies) area. An important limestones in southern graphic Germany. is that of where thin locality nearby Brunn, a paper
shale of Late in Kimmeridgian age deposited a very Locality Stratigraphic age
shallow lagoon surroundedby calcareous green algae Mornsheim, Daiting Early Tithonian, Hybonotum meadows, coral reefs, and small islands, is exposed. Zone, uppermostpart is the Slightly younger (still Late Kimmeridgian) Solnhofen, Langenaltheim Early Tithonian, Hybonotum Nusplingen Lithographic Limestone in SW Ger- Zone, upper part famous for its richness in fossil Eichstatt, Zandt Early Tithonian, Hybonotum many, sharks as
Zone, lower well This part as decapod crustaceans. was deposited
Painten, Painten, Schamhaupten, Kimmeridgian/Tithonian in surrounded a c. 100 m deep lagoon, by sponge- Ochselberg boundary beds microbial mounds and islands. East of Eichstatt Nusplingen Late Kimmeridgian, Beckeri (Frankische Alb), several localities have Zone, Ulmense Subzone yielded
crustaceans from limestones Brunn LateLate Kimmeridgian, Beckeri decapod lithographic
Zone, Subeumela Subzone spanning the Kimmeridgian/Tithonian boundary,
LateLate Ebenwies/Naab Kimmeridgian, Pseudo-IPseudo- later depending on a definition of the lower boundary mutabilismutahilis Zone of the Tithonian. Excavations by staff members of
the Jura-Museum Eichstatt in the siliceous litho-
graphic limestones of Schamhaupten have resulted
and local differences in paleoecology. Only an in a proliferation of new decapod crustacean mate-
in-depth knowledge oflocal lithologies allows rec- rial. In contrast, the lithographic limestones ofZandt
ognizing the exact provenance of material. Strati- and the more biodetritic limestones of the nearby
well-documented material have graphically has recently Ochselberg yielded a diverse, excellently pre-
been collected at several scientific excavations, and served decapod crustacean fauna, including sev-
there are a number of private collectors who focus eral new taxa. Other faunas of an age similar to
on special sites. those ofZandt are from quarries north of Eichstatt,
e.g., Blumenberg, Schernfeld, Wintershof. Many
‘old’ taxa were first described from this area.
Results The classic lithographic limestones of Solnhofen
and are of Tithonian thus Langenaltheim Early age,
Repositories of material referred to here are as are younger than those of the Eichstatt area. Apart
from of follows: BSPM - Baycrische Staatssammlung fur mass occurrences planktonic crinoids, these
limestones in fossils. Vertebrate fau- Paliiontologie und historische Geologic, Miinchen; are very poor
- nal and other data more JME Jura-Museum, Eichstatt; NHM - The Natu- suggest a pelagic setting.
ral History Museum, London; SMNS - Staatliches Both the Eichstatt and Solnhofen sections comprise
Museum fur Naturkunde, Stuttgart. micritic limestones deposited in larger, shallow
Schweigert et al. (1996) and Schweigert (2000) basins, the margins of which are poorly known,
worked out the biostratigraphy of the Upper Kim- since at present only the purest limestones in the
meridgian, but it should be noted that our knowl- central basins are quarried. The Solnhofen Litho-
edge of ammonite faunal succession of the Tithonian graphic Limestone Formation is usually overlain
Stage is still in its infancy. The oldest lithographic by a hardground or another discontinuity followed
the limestones, in part not yet dated precisely, occur by more biodetritic and silica-rich Moernsheim
Fig. 2. A, Rosenfeldia oppeli (Woodward), Solnhofen Lithographic Limestones (upper Eichstatt Formation, Lower Tithonian), most
from likely Eichstatt area (NHM 44886, holotype, Haberlein Colin). B, Erymastacus n. sp., Nusplingen Lithographic Limestone
Formation (Upper Kimmeridgian), Nusplingen quarry (SMNS 64872,2002excavation). C, Megachelafrickhingeri Schweigert, Solnhofen
Limestones Peneeid Solnhofen Lithographic (Lower Tithonian), Eichstatt (SMNS 64933, paratype, LudwigColin). D, (n. gen., n. sp.),
Limestones Lithographic (PaintenFormation,Ochselberg Member,Kimmeridgian/Tithonian boundary beds), Ochselberg near Breitenhill
(Burger Colin, Bad Hersfeld). All scale bars equal 10 mm.
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which has also Formation, yielded numerous de- served taxa came to light, which often allowed a capod crustaceans. revision of these.
Future research into decapod crustaceans from
the lithographic limestones of southern Germany
has modern revision of the Discussion many aspects. Firstly, a
is in which the taxa necessary, stratigraphic age,
provenance, and taphonomy is included. Most of The bulk of the decapod crustaceans faunas of the material the type is still available, but some taxa lithographic limestones in southern Germany was in are fact nomina dubia. The bulk of the types described in nineteenth century monographs (von from the Frankische Alb are in museums in Miinchen Schlotheim, 1820,1822; von Munster, 1839;Oppel, (Munster and Oberndorfer collections) and in Ber- 1861, 1862). Discoveries of extant eryonids dur- lin (von Schlotheim and Redenbacher collections). ing deep-sea expeditions triggered the study of fossil Few others survive in the Senckenberg Museum at forms (e.g., von Knebel, 1907). In the first half of Frankfurt am Main (H. von Meyer Colin), at the the twentieth century, both Beurlen and Glaessner Geiseltal Museum (Halle/Saale; Germar Colin), and almost ignored decapods from the lithographic lime- at the Natural History Museum (London; Haberlein stones in their scientific work. In recent years, c. Colin). The types from Nusplingen are almost ex- 56 valid taxa from the Frankische Alb and four others clusively housed at the Staatliches Museum fiir from the isolated locality ofNusplingen (Schwaben) Naturkunde Stuttgart (O. Fraas Colin). have have been described, and only a few of these For many taxa, lectotypes have never been des- been revised Forster (1966, 1967, 1973, briefly by and there still ignated, are some doubtful genera 1977). Following his untimely death in 1987, the and species. New topotypes in better preservation study of fossil decapod crustaceans from Germany and the use of ultraviolet light (Fig. 2A-D) often declined. allow a more detailed redescription. Unfortunately,
From the late 1990s scientific excava- onwards, the use of ultraviolet light does not work with de- tions both in eastern Bayern (Brunn), the southern capods from Zandt, which otherwise includes the Alb Frankische (Schamhaupten) and at Nusplingen, material. best Secondly, a larger amount of new has rekindled research activities (Polz, 1999, 2000; taxa, approximately 15-20, still awaits description, Schweigert 2001 a,b,c; Schweigert & Dietl, 1999; documenting most orders of Jurassic decapods. Some & Schweigert et al., 2000; Schweigert Roper, 2001). of them are easily assigned to described genera, It became obvious that both for paleoenvironmental while others are completely new and demonstrate reconstructions and it is the systematic paleontology, incompleteness of our present knowledge of to the fossil material of each necessary separate decapod diversity in the Upper Jurassic. In several
Another is the mono- locality. strong pulse given by groups, even in those held to be conservative, evo- graphic documentationof‘Solnhofen’ fossils stored trends have been in the lutionary observed, e.g., in various museums around the world as well in as penaeid genus Antrimpos (Schweigert, 2001b), or several private collections 1994, 1999). in the Cancrinos. both correla- (Frickhinger, scyllarid In genera, aim Frickhinger’s was a complete documentation tion between and with provenance stratigraphic age of all of animals and from the ‘Solnhofen’ taxa plants described species has become obvious, suggesting limestones. From this otherwise unavailable data they most likely represent successive chronospecies.
Pool of the existence many new taxa became clear. In others, however, no morphological changes are
material of rare or in of different Additionally, good poorly pre- recognizable populations ages.
Gg. 3. A, Malmuncina wulfi Schweigert & Garassino, Solnhofen Lithographic Limestones (Painten Formation, Ochselberg Member,
K-immeridgian/Tithonian boundary beds), Ochselberg near Breitenhill (JME Ti 2003/1, holotype, Wulf Colin). 15, Udora brevispina von Munster, Solnhofen Lithographic Limestones (Upper Eichstatt Formation, Lower Tithonian), Eichstatt area (BSPM 1964 XXIII
Solnhofen Limestones Eichstatt Lower Eichstatt C, Eukyphid (n. gen., n. sp.), Lithographic (Upper Formation, Tithonian), area
(SMNS Penaeid Limestones Lower 64943, Kiimpel Colin), D, (n. gen., n. sp.), Solnhofen Lithographic (Upper Eichstatt Formation,
Tithonian), Eichstatt area 64945, (SMNS Kiimpel Colin). All scale bars equal 10 mm.
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Reptants are mostly represented by molts, ex- 2003). All three genera are included in the Axiidae,
for in cept Mecochirus longimanatus (von Schlotheim), which they represent a primitive clade joined by
one of the commonest decapods in the Solnhofen a well-developed diaeresis.
and their Within the the recent area, few eryonids (Cycleryon sp.) with natantians, new discover-
the ies last trails and the dead body preserved on sea are enormous. Especially the eukyphids formerly
floor. known few but by very morphologically strikingly
In eryonids, sexual dimorphism has been dem- heterogeneous genera (Hefriga, Udora, Udorella,
onstrated for Cycleryon propinquus (von Schlot- and Blaculla) are much more diverse. Amongst
assumed be the and C. the well-known heim), to male, spinimanus them, only Hefriga serrata von
be the female to be (Germar), thought to (Schweigert, Munster (see Forster, 1967) appears quite
A 2001a). unique, incompletely preserved speci- common, although mostly incompletely preserved
men (Fig. 3A), described by Woodward (1866) as or juvenile, whereas the other taxa are extremely
Fate Eryon oppeli, represents a Jurassic represen- rare. Again, ultraviolet light analysis provides fine
tative of Rosenfeldia, a previously monospecific details otherwise barely visible, such as delicate
genus from the Upper Triassic. Although hundreds spines of the pereiopods or pleopods and uropods
of eryonids have been collected from the Upper in Udora brevispina von Munster (Fig. 2B). One
Jurassic lithographic limestones since the original of the recently discovered new eukyphid forms
description of the type of ‘Eryon’ oppeli, no other is the monospecific Schmelingia, which exhibits
specimen has come to light. prolonged first pereiopods bearing strong chelae
Among the Astacidea, erymids are the common- (Schweigert, 2002). Another strange, yet unnamed,
est. However, apart from the relatively abundant eukyphid decapod, which cannot be assigned to a
several other modern bears feather-like lateral Eryma modestiformis (von Schlotheim) family, large, ap-
species occur, differing mainly in details of their pendices at the pleopods of all somites (Fig. 2C),
whereas the features of the which have had chelae, carapace are must a special function, not only
for differentia- for the not significant enough a specific hatching eggs.
tion. Another of is characterized the habitats of of these group erymids by Obviously, many new
chelae with elongate pincers bearing strong and forms were far away from the place of subsequent
widely spaced teeth. Beurlen (1928) included these entombment, with their exuviae drifted only in ex-
in Erymastacus, whereas Forster (1966) ignored this ceptional cases into the anoxic basins, hostile to
in character and lumped them again Eryma s. sir. benthic organisms. Amongst penaeids, Antrimpos
Although isolated chelae of Erymastacus occur is the most abundant genus, accompanied by sev-
frequently in various Middle and Upper Jurassic eral species ofAeger and Acanthochirana. Also in
beds in southern and the records and Germany elsewhere, penaeids, new species even new genera are
from lithographic limestones are extremely rare and recorded (Fig. 2D; Fig. 3D). Genera that originally
material is mostly incomplete. In addition to Erymas- included more than a single species, often need
‘ For Muns- tacus major (Oppel), another species of this genusi revision. example, ‘Bylgia’ hexadonvon
‘ occurs in the Kimmeridgian at Nusplingen, based ter and ’B.’ haeberleini von Munster are best re-
on a isolated Another ferred a new with B. the single cheliped (Fig 2B). to genus, spinosa as type
astacidean family, the Uncinidae, was previously species, already designated by Oppel (1862). A new
assumed to be confined to the Upper Fiassic; this species of Bylgia, which differs from B. spinosa in
is obviously a collecting bias, because three speci- its larger rostrum bearing more dorsal spines has
of in mens a Fate Jurassic uncinid (Fig. 2A) are now recently been discovered the Oechselberg Shales.
known from Zandt, Ochselberg, and from the Eich- Other rare peneid species such as Koelga curvirostris
statt area (Schweigert & Garassino, 2003). von Munster, originally only known from a single,
Thalassinoids are rarely found in lithographic poorly preserved specimen, have lately been found
in in limestones. In addition to the long-known Magila several specimens and a much better condi- and the tion. Etallonia, new genus Megachela (Fig. 3C) with an extraordinarily prolonged fixed finger oc- Isolated carapaces of prosopids are very abun-
curs sporadically at Eichstatt and Zandt (Schweigert, dant in the ‘normal’ marly limestone lithology of
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in southern Polz H. 1999. Etallonia hoellorum the Upper Jurassic Germany, beyond sp. nov. (Crustacea,
Decapoda, Axiidae) aus dem Oberkimmeridgium der siid- the laminated facies. In the lithographic limestones, lichen Frankenalb. Archaeopteryx 17: 33-39. however, they are almost absent. Only at few lo- Polz H. 2000. viohli Glyphea sp. nov. (Crustacea: Decapoda; calities do (Ochselberg near Breitenhill) prosopids Glypheidae) aus den Solnhofener Plattenkalken. Archae- occur sporadically. are Usually, they poorly pre- opteryx 18: 17-25. served because of their weak mineralization, but Schlothcim EF von 1820. Die Petrefactenkunde auf dem
the jetzigen Standpunktedurch die Beschreibung seiner Samm- occasionally carapaces preserve even pereio- lung versteinerter und fossiler Uberresle des Thier- und pods. PJlanzenreichs der Vorwelt erldutert. Gotha: Becker.
Schlotlicim EF von 1822. Nachtrdge zur Petrefactenkunde,
I. Gotha: Becker.
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